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Systematics, phylogeny and biogeography
RESEARCH ARTICLE

A preliminary phylogeny for the pseudoscorpion family Garypinidae (Pseudoscorpiones: Garypinoidea), with new taxa and remarks on the Australasian fauna

Mark S. Harvey https://orcid.org/0000-0003-1482-0109 A B *
+ Author Affiliations
- Author Affiliations

A Collections & Research, Western Australian Museum, 49 Kew Street, Welshpool, WA 6106, Australia.

B School of Biological Sciences, University of Western Australia, Crawley, WA 6009, Australia.

* Correspondence to: mark.harvey@museum.wa.gov.au

Handling Editor: Gonzalo Giribet

Invertebrate Systematics 37(9) 623-676 https://doi.org/10.1071/IS23029
Submitted: 10 June 2023  Accepted: 7 August 2023   Published: 26 September 2023

© 2023 The Author(s) (or their employer(s)). Published by CSIRO Publishing.

Abstract

The pseudoscorpion family Garypinidae is globally distributed with 79 species in 21 genera and several species represented by Mesozoic and Eocene fossils. This was recently included with the family Larcidae in a unique superfamily, Garypinoidea but there are no phylogenetic hypotheses for the group. Sequence data were obtained for 14 species in 8 genera and numerous outgroup taxa that formed the basis for a preliminary molecular phylogeny. A new subfamily classification is proposed with Protogarypininae, subfamily nov. comprising five genera mostly found in the southern hemisphere, Amblyolpiinae subfamily nov. comprising two genera and Garypininae for the remaining genera. Several new taxa are described including the first Australian species of Aldabrinus, A. rixi sp. nov., a new genus from South-East Asia, Nobilipinus, comprising Nobilipinus nobilis (With, 1906), N. vachoni (Redikorzev, 1938) (that is removed from the synonymy of G. nobilis) and five new species, N. affinis, N. galeatus, N. karenae, N. kohi and N. tricosus, and Solinus pingrup sp. nov. from south-western Australia. Paraldabrinus Beier, 1966 is newly synonymised with Aldabrinus, and Indogarypinus Murthy and Ananthakrishan, 1977 is newly synonymised with Solinus. The holotype of Garypinus mirabilis With, 1907 from Hawaii is redescribed but found to be a tritonymph, rendering the generic identity uncertain.

ZooBank: urn:lsid:zoobank.org:pub:E15E4705-0697-4208-9338-A778343996CA

Keywords: Aldabrinus, classification, Garypinus, morphology, new taxa, Nobilipinus, Solinus, taxonomy.

References

Arabi J, Judson MLI, 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 |

Bedoya-Roqueme E, Pérez-Agudelo M, Zaragoza JA, Quirós JA (2017) Nuevos reportes de Pseudoescorpiones (Arachnida) de Bosques de Manglar en Córdoba, Caribe colombiano. Revista Ibérica de Aracnología 30, 25-36 [In Spanish].
| Google Scholar |

Beier M (1930) Die Pseudoskorpione der Sammlung Roewer. Zoologischer Anzeiger 91, 284-300 [In German].
| Google Scholar |

Beier M (1932a) Pseudoscorpionidea II. Subord. C. Cheliferinea. Tierreich 58, 1-297 [In German].
| Google Scholar |

Beier M (1932b) Pseudoscorpionidea I. Subord. Chthoniinea et Neobisiinea. Tierreich 57, 1-258 [In German].
| Google Scholar |

Beier M (1936) Zoologische Ergebnisse einer Reise nach Bonaire, Curaçao und Aruba im Jahre 1930. No. 21. Einige neue neotropische Pseudoscorpione. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere 67, 443-447 [In German].
| Google Scholar |

Beier M (1937) Pseudoscorpione aus dem baltischen Bernstein. In ‘Festschrift zum 60. Geburtstage von Professor Dr Embrik Strand. Vol. 2’. pp. 302–316. (Izdevniecība “Latvija”: Riga, Latvia) [In German]

Beier M (1944) Über Pseudoscorpioniden aus Ostafrika. Eos, Madrid 20, 173-212 [In German].
| Google Scholar |

Beier M (1947) Zur Kenntnis der Pseudoscorpionidenfauna des südlichen Afrika, insbesondere der südwest- und südafrikanischen Trockengebiete. Eos, Madrid 23, 285-339 [In German].
| Google Scholar |

Beier M (1951) Die Pseudoscorpione Indochinas. Mémoires du Muséum National d’Histoire Naturelle, Paris, Nouvelle Série 1, 47-123 [In German].
| Google Scholar |

Beier M (1954a) Report from Prof. T. Gislén’s expedition to Australia in 1951–1952. 7. Pseudoscorpionidea. Acta Universitatis Lundensis, Nova Series (2) 50, 1-26.
| Google Scholar |

Beier M (1954b) Zwei neue Pseudoscorpione aus Ostafrika. Zoologischer Anzeiger 152, 84-88 [In German].
| Google Scholar |

Beier M (1955a) Pseudoscorpionidea. In ‘South African animal life. Results of the Lund Expedition in 1950–1951. Vol. 1’. (Eds B Hanstrom, P Brinck, G Rudebeck) pp. 263–328. (Almquist and Wiksell: Stockholm, Sweden)

Beier M (1955b) Über Pseudoscorpione aus Syrien und Palästina. Annalen des Naturhistorischen Museums in Wien 60, 212-219 [In German].
| Google Scholar |

Beier M (1958) The Pseudoscorpionidea (false-scorpions) of Natal and Zululand. Annals of the Natal Museum 14, 155-187.
| Google Scholar |

Beier M (1959a) Zur Kenntnis der Pseudoscorpioniden-Fauna des Andengebietes. Beiträge zur Neotropischen Fauna 1, 185-228 [In German].
| Google Scholar |

Beier M (1959b) Pseudoscorpione aus dem Belgischen Congo gesammelt von Herrn N. Leleup. Annales du Musée du Congo Belge, Sciences Zoologiques 72, 5-69 [In German].
| Google Scholar |

Beier M (1959c) Zur Kenntnis der Pseudoscorpioniden‐Fauna Afghanistans. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere 87, 257-282 [In German].
| Google Scholar |

Beier M (1962) Pseudoscorpioniden aus der Namib-Wüste. Annals of the Transvaal Museum 24, 223-230 [In German].
| Google Scholar |

Beier M (1963) Ordnung Pseudoscorpionidea (Afterskorpione). In ‘Bestimmungsbücher zur Bodenfauna Europas. Vol. 1’. (Akademie-Verlag) [In German]

Beier M (1964a) Die Pseudoscorpioniden-Fauna Chiles. Annalen des Naturhistorischen Museums in Wien 67, 307-375 [In German].
| Google Scholar |

Beier M (1964b) Weiteres zur Kenntnis der Pseudoscorpioniden-Fauna des südlichen Afrika. Annals of the Natal Museum 16, 30-90 [In German].
| Google Scholar |

Beier M (1964c) Some further nidicolous Chelonethi (Pseudoscorpionidea) from Malaya. Pacific Insects 6, 312-313.
| Google Scholar |

Beier M (1964d) Pseudoscorpione von der Insel San Ambrosio. Annalen des Naturhistorischen Museums in Wien 67, 303-306 [In German].
| Google Scholar |

Beier M (1966a) Ergebnisse der österreichischen Neukaledonien-Expedition 1965. Pseudoscorpionidea. Annalen des Naturhistorischen Museums in Wien 69, 363-371 [In German].
| Google Scholar |

Beier M (1966b) Die Pseudoscorpioniden der Salomon-Inseln. Annalen des Naturhistorischen Museums in Wien 69, 133-159 [In German].
| Google Scholar |

Beier M (1966c) Über Pseudoskorpione der Insel Rhodos. Annalen des Naturhistorischen Museums in Wien 69, 161-167 [In German].
| Google Scholar |

Beier M (1967a) Contributions to the knowledge of the Pseudoscorpionidea from New Zealand. Records of the Dominion Museum 5, 277-303.
| Google Scholar |

Beier M (1967b) Pseudoskorpione aus dem tropischen Ostafrika (Kenya, Tansania, Uganda, etc.). Annalen des Naturhistorischen Museums in Wien 70, 73-93 [In German].
| Google Scholar |

Beier M (1967c) Ergebnisse zoologischer Sammelreisen in der Türkei. Annalen des Naturhistorischen Museums in Wien 70, 301-323 [In German].
| Google Scholar |

Beier M (1970a) Die Pseudoscorpione der Royal Society Expedition 1965 zu den Salomon-Inseln. Journal of Natural History 4, 315-328 [In German].
| Google Scholar |

Beier M (1970b) Ergänzungen zur Pseudoskorpionidenfauna der Kanaren. Annalen des Naturhistorischen Museums in Wien 74, 45-49 [In German].
| Google Scholar |

Beier M (1971a) Pseudoskorpione unter Araucarien-Rinde in Neu-Guinea. Annalen des Naturhistorischen Museums in Wien 75, 367-373 [In German].
| Google Scholar |

Beier M (1971b) Pseudoskorpione aus dem Iran. Annalen des Naturhistorischen Museums in Wien 75, 357-366 [In German].
| Google Scholar |

Beier M (1973) Weiteres zur Kenntnis der Pseudoscorpioniden Südwestafrikas. Cimbebasia, A 2, 97-101 [In German].
| Google Scholar |

Beier M (1975) Neue Pseudoskorpione aus Australien und Neu-Guinea. Annalen des Naturhistorischen Museums in Wien 78, 203-213 [In German].
| Google Scholar |

Beier M (1976) The pseudoscorpions of New Zealand, Norfolk and Lord Howe. New Zealand Journal of Zoology 3, 199-246.
| Crossref | Google Scholar |

Beier M (1977) Pseudoscorpiones. In: La faune terrestre de l’île de Sainte-Hélène IV. Annales du Musée Royal de l’Afrique Centrale, Zoologie (8) 220, 2-11 [In French].
| Google Scholar |

Beier M (1978) Pseudoskorpione von den Galapagos-Inseln. Annalen des Naturhistorischen Museums in Wien 81, 533-547 [In German].
| 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 |

Benedict EM (1978) False scorpions of the genus Apocheiridium Chamberlin from western North America (Pseudoscorpionida, Cheiridiidae). Journal of Arachnology 5, 231-241.
| Google Scholar |

Benedict EM, Malcolm DR (1978) Some garypoid false scorpions from western North America (Pseudoscorpionida: Garypidae and Olpiidae). Journal of Arachnology 5, 113-132.
| Google Scholar |

Brown RW (1956) ‘Composition of scientific words’, revised edn. (Smithsonian Institution Press: Washington, DC, USA)

Callaini G (1982) Étude comparative de la membrane pleurale des pseudoscorpions au microscope électronique à balayage. Memorie, Atti della Società Toscana di Scienze Naturali Residente in Pisa B 88(supplemento), 16-26 [In French].
| Google Scholar |

Chamberlin JC (1923) New and little known pseudoscorpions, principally from the islands and adjacent shores of the Gulf of California. Proceedings of the California Academy of Science, 4th Series 12, 353-387.
| Google Scholar |

Chamberlin JC (1924) The Cheiridiinae of North America (Arachnida - Pseudoscorpionida). Pan-Pacific Entomologist 1, 32-40.
| Google Scholar |

Chamberlin JC (1930) LXI.—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).5. Annals and Magazine of Natural History 5(30), 585-620.
| Crossref | Google Scholar |

Chamberlin JC (1931a) The arachnid order Chelonethida. Stanford University Publications, Biological Sciences 7(1), 1-284.
| Google Scholar |

Chamberlin JC (1931b) Parachernes ronnaii, a new genus and species of false scorpion from Brazil (Arachnida – Chelonethida). Entomological News 42, 192-195.
| Google Scholar |

Chamberlin JC (1932) On some false scorpions of the superfamily Cheiridioidea (Arachnida – Chelonethida). Pan-Pacific Entomologist 8, 137-144.
| Google Scholar |

Chamberlin JC (1943) The taxonomy of the false scorpion genus Synsphyronus with remarks on the sporadic loss of stability in generally constant morphological characters (Arachnida: Chelonethida). Annals of the Entomological Society of America 36, 486-500.
| Crossref | Google Scholar |

Cockerell TDA (1920) XXXVI.—Fossil arthropods in the British Museum.—I. Annals and Magazine of Natural History 5(27), 273-279.
| Crossref | Google Scholar |

Daday E (1889) A Magyar Nemzeti Muzeum álskorpióinak áttekintése. Természetrajzi Füzetek 11, 111–136, 165-192 [In Hungarian].
| Google Scholar |

Dashdamirov S (1996) Notes on the false scorpion genus Solinus in the Caucasus (Pseudoscorpionida: Olpiidae). Revue Suisse de Zoologie, Hors Série 1, 117-123.
| Google Scholar |

Dashdamirov S (2007) A new genus and species of false scorpion from Vietnam showing remarkable chelal modifications (Arachnida: Chelonethida). Acta Biologica Benrodis 13, 219-229.
| Google Scholar |

Dashdamirov S, Schawaller W (1993) Pseudoscorpions from Middle Asia, Part 2 (Arachnida: Pseudoscorpiones). Stuttgarter Beiträge zur Naturkunde (A) 496, 1-14.
| Google Scholar |

Ellingsen E (1910) Die Pseudoskorpione des Berliner Museums. Mitteilung aus dem Zoologischen Museum in Berlin 4, 357-423 [In German].
| Google Scholar |

Ellingsen E (1914) On the pseudoscorpions of the Indian Museum, Calcutta. Records of the Indian Museum 10, 1-14.
| Google Scholar |

Feio JLA (1945) Novos pseudoscorpiões de região neotropical (com a descriçao de uma subfamilia, dois géneros e sete espécies). Boletim do Museu Nacional Rio de Janeiro, n.s. Zoologia 44, 1-47 [In Portuguese].
| Google Scholar |

Gardini G (1983) Larca italica n. sp. cavernicola dell’Appennino Abruzzese (Pseudoscorpionida, Garypidae) (Pseudoscorpioni d’Italia XV). Bollettino della Società Entomologica Italiana 115, 63-69 [In Italian].
| Google Scholar |

Gardini G, Gavalas I (2021) The pseudoscorpions of the island of Iraklia (Cyclades, Greece). Parnassiana Archives 9, 43-78.
| Google Scholar |

Gardini G, Galli L, Zinni M (2017) Redescription of Geogarypus minor, type species of the genus Geogarypus, and description of a new species from Italy (Pseudoscorpiones: Geogarypidae). Journal of Arachnology 45, 424-443.
| 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 norarabithern Myanmar. Cretaceous Research 130, 105030.
| Crossref | Google Scholar |

Hadži J (1933) Beitrag zur Kenntnis der Pseudoskorpionen-Fauna des Küstenlandes. Bulletin International de l’Académie Yugoslave des Sciences et des Beaux-arts 27, 173-199 [In French].
| Google Scholar |

Harms D, Roberts JD, Harvey MS (2019) Climate variability impacts on diversification processes in a biodiversity hotspot: a phylogeography of ancient pseudoscorpions in southwestern Australia. Zoological Journal of the Linnean Society 186, 934-949.
| Crossref | Google Scholar |

Harvey MS (1985) The systematics of the family Sternophoridae (Pseudoscorpionida). Journal of Arachnology 13, 141-209.
| Google Scholar |

Harvey MS (1986) The Australian Geogarypidae, new status, with a review of the generic classification (Arachnida: Pseudoscorpionida). Australian Journal of Zoology 34, 753-778.
| Crossref | Google Scholar |

Harvey MS (1987a) A revision of the genus Synsphyronus Chamberlin (Garypidae: Pseudoscorpionida: Arachnida). Australian Journal of Zoology, Supplementary Series 126, 1-99.
| Crossref | Google Scholar |

Harvey MS (1987b) Redescriptions of Geogarypus bucculentus Beier and G. pustulatus Beier (Geogarypidae: Pseudoscorpionida). Bulletin of the British Arachnological Society 7, 137-141.
| Google Scholar |

Harvey MS (1988) Pseudoscorpions from Krakatau Islands and adjacent regions, Indonesia (Chelicerata: Pseudoscorpionida). Memoirs of the Museum of Victoria 49, 309-353.
| Crossref | Google Scholar |

Harvey MS (1990) New pseudoscorpions of the genera Americhernes Muchmore and Cordylochernes Beier from Australia (Pseudoscorpionida: Chernetidae). Memoirs of the Museum of Victoria 50, 325-336.
| Crossref | Google Scholar |

Harvey MS (1992) The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Systematics 6, 1373-1435.
| Crossref | Google Scholar |

Harvey MS (2000) From Siam to Rapa Nui – the identity and distribution of Geogarypus longidigitatus (Rainbow) (Pseudoscorpiones: Geogarypidae). Bulletin of the British Arachnological Society 11, 377-384.
| Google Scholar |

Harvey MS (2010) Two new species of Synsphyronus (Pseudoscorpiones: Garypidae) from southern Western Australian granite landforms. Records of the Western Australian Museum 26, 11-22.
| 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, Du Preez G (2014) A new troglobitic ideoroncid pseudoscorpion (Pseudoscorpiones: Ideoroncidae) from southern Africa. Journal of Arachnology 42, 105-110.
| Crossref | Google Scholar |

Harvey MS, Muchmore WB (1989) 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, Wynne JJ (2014) Troglomorphic pseudoscorpions (Arachnida: Pseudoscorpiones) of northern Arizona, with the description of two new short-range endemic species. Journal of Arachnology 42, 205-219.
| Crossref | Google Scholar |

Harvey MS, Ratnaweera PB, Udagama PV, Wijesinghe MR (2012) A new species of the pseudoscorpion genus Megachernes (Pseudoscorpiones: Chernetidae) associated with a threatened Sri Lankan rainforest rodent, with a review of host associations of Megachernes. Journal of Natural History 46, 2519-2535.
| 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, 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 |

Henderickx H, Vets V (2002) A new Larca (Arachnida: Pseudoscorpiones: Larcidae) from Crete. Bulletin of the British Arachnological Society 12, 280-282.
| Google Scholar |

Hernández-Corral J, Zaragoza JA, Micó E (2018) New species of Pseudoscorpiones (Arachnida) from tree hollows in a Mediterranean oak forest in Spain. Zootaxa 4497, 201-225.
| Crossref | Google Scholar |

Heurtault J (1970) Pseudoscorpions du Tibesti (Tchad). I. Olpiidae. Bulletin du Muséum National d’Histoire Naturelle, Paris, 2e Série 41(5), 1164-1174 [In French].
| Google Scholar |

Heurtault J (1982) Le développement postembryonnaire chez deux espèces nouvelles de Pseudoscorpions Olpiinae du Venezuela. Revue de Nordest Biologie 3, 57-85 [In French].
| Google Scholar |

Hlebec D, Podnar M, Kučinić M, Harms D (2023) Molecular analyses of pseudoscorpions in a subterranean biodiversity hotspot reveal cryptic diversity and microendemism. Scientific Reports 13, 430.
| Crossref | Google Scholar |

Hoff CC (1950) Pseudoescorpionidos nuevos o poco conocidos de la Argentina (Arachnida, Pseudoscorpionida). Arthropoda, Buenos Aires 1, 225-237 [In Spanish].
| Google Scholar |

Hoff CC (1952) Two new species of pseudoscorpions from Illinois. Transactions of the Illinois State Academy of Sciences 45, 188-195.
| Google Scholar |

Hoff CC (1956) Diplosphyronid pseudoscorpions from New Mexico. American Museum Novitates 1780, 1-49.
| Google Scholar |

Hoff CC (1961) Pseudoscorpions from Colorado. Bulletin of the American Museum of Natural History 122, 409-464.
| Google Scholar |

Hoff CC (1964) The pseudoscorpions of Jamaica. Part 3. The suborder Diplosphyronida. Bulletin of the Institute of Jamaica, Science Series 10(3), 1-47.
| Google Scholar |

Johnson J, Loria SF, Joseph MM, Harms D (2022) Biogeographical and diversification analyses of Indian pseudoscorpions reveal the Western Ghats as museums of ancient biodiversity. Molecular Phylogenetics and Evolution 175, 107495.
| Crossref | Google Scholar |

Judson MLI 1993 African Chelonethi: studies on the systematics, biogeography and natural history of African pseudoscorpions (Arachnida). PhD thesis, University of Leeds, Leeds, UK.

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 (2005) Baltic amber fossil of Garypinus electri Beier provides first evidence of phoresy in the pseudoscorpion family Garypinidae (Arachnida: Chelonethi). In ‘European Arachnology 2003 – Proceedings of the 21st European Colloquium of Arachnology’, 4–9 August 2003, Saint Petersburg, Russian Federation. (Eds DV Logunov, D Penney) pp. 127–131. (KMK Scientific Press Ltd: Moscow, Russian Federation)

Judson MLI (2007) A new and endangered species of the pseudoscorpion genus Lagynochthonius from a cave in Vietnam, with notes on chelal morphology and the composition of the Tyrannochthoniini (Arachnida, Chelonethi, Chthoniidae). Zootaxa 1627, 53-68.
| Crossref | Google Scholar |

Judson MLI, Legg G (1996) Discovery of the pseudoscorpion Larca lata (Garypoidea, Larcidae) in Britain. Bulletin of the British Arachnological Society 10, 205-210.
| 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 |

Katoh K, Misawa K, Kuma K, Mityata 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 |

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 |

Koch L (1873) ‘Uebersichtliche Darstellung der Europäischen Chernetiden (Pseudoscorpione).’ (Bauer und Raspe: Nürnberg, German Empire) [In German]

Lee VF (1979) The maritime pseudoscorpions of Baja California, México (Arachnida: Pseudoscorpionida). Occasional Papers of the California Academy of Sciences 131, 1-38.
| Google Scholar |

Mahnert V (1974) Einige Pseudoskorpione aus Israel. Revue Suisse de Zoologie 81, 377-386 [In German].
| Crossref | Google Scholar |

Mahnert V (1976) Zwei neue Pseudoskorpion-Arten (Arachnida) aus griechischen Höhlen (Über griechische Pseudoskorpione VII). Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck 63, 177-183 [In German].
| Google Scholar |

Mahnert V (1981a) Sigles trichobothriaux chez les pseudoscorpions (Arachnida). Memorie, Atti della Società Toscana di Scienze Naturali Residente in Pisa B 88(supplemento), 185-192 [In French].
| Google Scholar |

Mahnert V (1981b) Die Pseudoskorpione (Arachnida) Kenyas. I. Neobisiidae und Ideoroncidae. Revue Suisse de Zoologie 88, 535-559 [In German].
| Crossref | Google Scholar |

Mahnert V (1982a) Die Pseudoskorpione (Arachnida) Kenyas II. Feaellidae; Cheiridiidae. Revue Suisse de Zoologie 89, 115-134 [In German].
| Google Scholar |

Mahnert V (1982b) Die Pseudoskorpion (Arachnida) Kenyas, IV. Garypidae. Annales Historico-Naturales Musei Nationalis Hungarici 74, 307-329 [In German].
| Google Scholar |

Mahnert V (1984a) Forschungen an der Somalilandküste. Am Strand und auf den Dünen bei Sar Uanle. 36. Pseudoscorpiones (Arachnida). Monitore Zoologico Italiano, n.s. 19(supplemento), 43-66 [In German].
| Google Scholar |

Mahnert V (1984b) Beitrag zu einer besseren Kenntnis der Ideoroncidae (Arachnida: Pseudoscorpiones), mit Beschreibung von sechs neuen Arten. Revue Suisse de Zoologie 91, 651-686 [In German].
| Google Scholar |

Mahnert V (1985) Weitere Pseudoskorpione (Arachnida) aus dem zentralen Amazonasgebiet (Brasilien). Amazoniana 9, 215-241 [In German].
| Google Scholar |

Mahnert V (1988) Zwei neue Garypininae-Arten (Pseudoscorpiones: Olpiidae) aus Afrika mit Bemerkungen zu den Gattungen Serianus Chamberlin und Paraserianus Beier. Stuttgarter Beiträge zur Naturkunde (A) 420, 1-11 [In German].
| Google Scholar |

Mahnert V (1991) Pseudoscorpions (Arachnida) from the Arabian Peninsula. Fauna of Saudi Arabia 12, 171-199.
| Google Scholar |

Mahnert V (1993) Pseudoskorpione (Arachnida: Pseudoscorpiones) von Inseln des Mittelmeers und des Atlantiks (Balearen, Kanarische Inseln, Madeira, Ascension), mit vorwiegend subterraner Lebensweise. Revue Suisse de Zoologie 100, 971-992 [In German].
| Crossref | Google Scholar |

Mahnert V (2007) Pseudoscorpions (Arachnida: Pseudoscorpiones) of the Socotra Archipelago, Yemen. Fauna of Arabia 23, 271-307.
| Google Scholar |

Mahnert V (2014) Pseudoscorpions (Arachnida: Pseudoscorpiones) from the Galapagos Islands (Ecuador). Revue Suisse de Zoologie 121, 135-210.
| Google Scholar |

Mahnert V, Schuster R (1981) Pachyolpium atlanticum n. sp., ein Pseudoskorpion aus der Gezeitenzone der Bermudas – Morphologie und Ökologie (Pseudoscorpiones: Olpiidae). Revue Suisse de Zoologie 88, 265-273 [In German].
| 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 |

Morikawa K (1953a) [On a new pseudoscorpion of family Olpiidae.] Zoological Magazine, Tokyo 62, 327-328 [In Japanese].
| Google Scholar |

Morikawa K (1953b) Notes on Japanese Pseudoscorpiones. II. Family Cheiridiidae, Atemnidae and Chernetidae. Memoirs of Ehime University (2B) 1, 345-354.
| Google Scholar |

Morikawa K (1960) Systematic studies of Japanese pseudoscorpions. Memoirs of Ehime University (2B) 4, 85-172.
| Google Scholar |

Muchmore WB (1968) A new species of the pseudoscorpion genus Serianus (Arachnida, Chelonethida, Olpiidae) from North Carolina. Entomological News 79, 145-150.
| Google Scholar |

Muchmore WB (1974) Pseudoscorpions from Florida. 1. The genus Aldabrinus (Pseudoscorpionida: Olpiidae). The Florida Entomologist 57, 1-7.
| Crossref | Google Scholar |

Muchmore WB (1976) Pseudoscorpions from Florida and the Caribbean area. 5. Americhernes, a new genus based upon Chelifer oblongus Say (Chernetidae). The Florida Entomologist 59, 151-163.
| Crossref | Google Scholar |

Muchmore WB (1979) Pseudoscorpions from Florida and the Caribbean area. 7. Floridian diplosphyronids. The Florida Entomologist 62, 193-213.
| Crossref | Google Scholar |

Muchmore WB (1980) Three new olpiid pseudoscorpions from California (Pseudoscorpionida, Olpiidae). Pan-Pacific Entomologist 56, 161-169.
| Google Scholar |

Muchmore WB (1981a) Cavernicolous species of Larca, Archeolarca and Pseudogarypus with notes on the genera (Pseudoscorpionida, Garypidae and Pseudogarypidae). Journal of Arachnology 9, 47-60.
| Google Scholar |

Muchmore WB (1981b) The identity of Olpium minutum Banks (Pseudoscorpionida, Olpiidae). Journal of Arachnology 9, 337-338.
| Google Scholar |

Muchmore WB (1982) The genus Anagarypus (Pseudoscorpionida: Garypidae). Pacific Insects 24, 159-163.
| Google Scholar |

Muchmore WB (1992) Cavernicolous pseudoscorpions from Texas and New Mexico (Arachnida: Pseudoscorpionida). Texas Memorial Museum, Speleological Monographs 3, 127-153.
| Google Scholar |

Muchmore WB, Alteri CH (1974) The genus Parachernes (Pseudoscorpionida, Chernetidae) in the United States, with descriptions of new species. Transactions of the American Entomological Society 99, 477-506.
| 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 |

Murthy VA, Ananthakrishnan TN (1977) Indian Chelonethi. Oriental Insects Monograph 4, 1-210.
| Google Scholar |

Nassirkhani M, Doustaresharaf MM (2019) Description of a new pseudoscorpion species of the genus Amblyolpium Simon (Pseudoscorpiones: Garypinidae) from north-western Iran. Zoology in the Middle East 65, 268-273.
| Crossref | Google Scholar |

Nassirkhani M, Shoushtari RV, Abadi MR (2016) A new pseudoscorpion species in Amblyolpium (Pseudoscorpiones: Garypinidae) from a house in Kermanshah Province, Iran. Arachnologische Mitteilungen 52, 1-3.
| Crossref | Google Scholar |

Neethling JA, Haddad CR (2016) A systematic revision of the South African pseudoscorpions of the family Geogarypidae (Arachnida: Pseudoscorpiones). Indago 32, 1-80.
| Google Scholar |

Nelson SO Jr (1982) The external morphology and life history of the pseudoscorpion Microbisium confusum Hoff. Journal of Arachnology 10, 261-274.
| Google Scholar |

Novák J (2013) First records of Larca lata (Hansen, 1884) and Neobisium biharicum Beier, 1939 in Hungary. Opuscula Zoologica, Budapest 44, 161-166.
| Google Scholar |

Novák J, Harvey MS, Szabó M, Hammel JU, Harms D, Kotthoff U, Hörweg C, Brazidec M, Ösi A (2023) A new Mesozoic record of the pseudoscorpion family Garypinidae from Upper Cretaceous (Santonian) Ajkaite amber, Ajka area, Hungary. Cretaceous Research, 105709. [Published online 9 September 2023] 10.1016/j.cretres.2023.105709

Novák J, Harvey MS (2018) New species and records of the pseudoscorpion genus Geogarypus (Pseudoscorpiones: Geogarypidae) from India, Sri Lanka and New Guinea. Zootaxa 4394, 417-427.
| Crossref | Google Scholar |

Piedra-Jiménez DF, Alvarez-Padilla F, González Santillán E (2019) Two new species of pseudoscorpions (Arachnida: Pseudoscorpiones) from a Mexican oak forest near Pico de Orizaba National Park. The Journal of Arachnology 47, 95-109.
| Crossref | Google Scholar |

Redikorzev V (1938) Les pseudoscorpions de l’Indochine française recueillis par M. C. Dawydoff. Mémoires du Muséum National d’Histoire Naturelle, Paris 10, 69-116 [In French].
| Google Scholar |

Roewer CF (1936) Chelonethi oder Pseudoskorpione. In ‘Bronn’s Klassen und Ordnungen des Tierreichs. Vol. 5(IV)(6)(1)’. (Ed. HG Bronns) pp. 1–160. (Akademische Verlagsgesellschaft M.B.H.: Leipzig, German Republic) [In German]

Sakayori H (1989) Postembryonic development of a neotenic pseudoscorpion, Microbisium pygmaeum (Ellingsen, 1907). Acta Arachnologica 38, 55-62.
| Crossref | Google Scholar |

Schawaller W (1994) Pseudoskorpione aus Thailand (Arachnida: Pseudoscorpiones). Revue Suisse de Zoologie 101, 725-759 [In German].
| Crossref | Google Scholar |

Simon E (1898) Étude sur les Arachnides de la région des Maures (Var). Feuille des Jeunes Naturalistes (3)(29), 2-4 [In French].
| Google Scholar |

Simon E (1900) Arachnida. In ‘Fauna Hawaiiensis. Vol. 2’. (Ed D Sharp) pp. 443–519. (Cambridge University Press)

Stanczak N, Harvey MS, Harms D, Hammel J, Kotthoff U, Loria SF. () Discovery of an extinct Eocene pseudoscorpion genus (Garypinoidea: Garypinidae) supports extinction and range contraction in Europe. PeerJ , in press.
| Google Scholar |

Tömösváry Ö (1884) Adatok az álskorpiók ismeretéhez (Data ad cognitionem Pseudoscorpionum). Természetrajzi Füzetek 8, 16-27 [In Hungarian].
| Google Scholar |

Tooren D (2002a) Pseudoscorpions of the genera Pachyolpium, Novohorus and Amblyolpium (Pseudoscorpiones: Olpiidae) from St Eustatius (Statia), St Martin (Sint Maarten) and Anguilla (Lesser Antilles, Leeward group). Zoologische Mededelingen 76, 451-472.
| Google Scholar |

Tooren D (2002b) Pseudoscorpions (Pseudoscorpiones: Olpiidae) of the genus Apolpium from Venezuela, and the genera Pachyolpium, Leptolpium gen. nov. and Serianus from Curaçao, Aruba and Bonaire. Zoologische Mededelingen 76, 141-192.
| 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 |

Tullgren A (1907) Zur Kenntnis aussereuropäischer Chelonethiden des Naturhistorischen Museums in Hamburg. Mitteilungen aus dem Naturhistorischen Museum in Hamburg 24, 21-75 [In German].
| Google Scholar |

Vachon M (1964) Sur l'établissement de formules précisant l’ordre d’apparition des trichobothries au cours du développement post-embryonnaire chez les Pseudoscorpions (Arachnides). Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences, Paris 258, 4839-4842 [In French].
| Google Scholar |

Vitali-di Castri V (1962) La familia Cheiridiidae (Pseudoscorpionida) en Chile. Investigaciones Zoológicas Chilenas 8, 119-142 [In Spanish].
| Google Scholar |

Vitali-di Castri V (1965) Cheiridium danconai n. sp. (Pseudoscorpionida) con consideraciones sobre su desarrollo postembrionario. Investigaciones Zoológicas Chilenas 12, 67-92 [In Spanish].
| Google Scholar |

With CJ (1905) X.—On Chclonethi, chiefly from the Australian region, in the collection of the British Museum, with observations on the “coxal sac” and on some cases of abnormal segmentation. Annals and Magazine of Natural History 15(85), 94-143.
| Crossref | Google Scholar |

With CJ (1906) The Danish expedition to Siam 1899–1900. III. Chelonethi. An account of the Indian false-scorpions together with studies on the anatomy and classification of the order. Oversigt over det Konigelige Danske Videnskabernes Selskabs Forhandlinger (7) 3, 1-214.
| Google Scholar |

With CJ (1907) On some New Species of Cheliferidæ, Hans., and Garypidæ, Hans., in the British Museum. Journal of the Linnean Society of London, Zoology 30, 49-85.
| Crossref | Google Scholar |

Zaragoza JA (2005) Two new cave-dwelling Larca species from the south-east of Spain (Arachnida, Pseudoscorpiones, Larcidae). Revue Suisse de Zoologie 112, 195-213.
| Crossref | Google Scholar |