Register      Login
Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography
RESEARCH ARTICLE

Supralittoral pseudoscorpions of the genus Garypus (Pseudoscorpiones : Garypidae) from the Indo-West Pacific region, with a review of the subfamily classification of Garypidae

Mark S. Harvey https://orcid.org/0000-0003-1482-0109 A B D , Mia J. Hillyer A , Jose I. Carvajal A and Joel A. Huey https://orcid.org/0000-0001-7108-0552 A B C
+ Author Affiliations
- Author Affiliations

A Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986, Australia.

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

C Adjunct, School of Natural Sciences, Edith Cowan University, Joondalup, WA 6027, Australia.

D Corresponding author. Email: mark.harvey@museum.wa.gov.au

Invertebrate Systematics 34(1) 34-87 https://doi.org/10.1071/IS19029
Submitted: 8 May 2019  Accepted: 10 July 2019   Published: 10 February 2020

Abstract

The pseudoscorpions of the genus Garypus L. Koch are restricted to seashore habitats where they occupy supralittoral and littoral zones primarily in tropical and subtropical areas. Few species have been recorded from the Indo-West Pacific region, and this project was devised to produce a review of the species found in museum collections and to test the relationships of the various garypid genera using a molecular analysis and an assessment of their morphology. A new subfamily classification is proposed with the subfamilies Garypinae, including Garypus and the new genus Anchigarypus Harvey (type species Garypus californicus Banks), and the Synsphyroninae for the other genera (Ammogarypus Beier, Anagarypus Chamberlin, Elattogarypus Beier, Eremogarypus Beier, Meiogarypus Beier, Neogarypus Vachon, Paragarypus Vachon, Neogarypus Vachon, Synsphyronus Chamberlin, and Thaumastogarypus Beier). The species-level revision of Garypus provides evidence for at least 14 species, most of which are known from only single localities. The following species are redescribed: G. insularis Tullgren from the Seychelles, G. krusadiensis Murthy & Ananthakrishnan from India and Sri Lanka, G. longidigitus Hoff from Queensland, Australia, G. maldivensis Pocock from the Maldives, G. nicobarensis Beier from the Nicobar Islands and G. ornatus Beier from the Marshall Islands. The holotype of G. insularis is a tritonymph, and not therefore readily identifiable. Nine new species are described: G. latens Harvey, sp. nov., G. malgaryungu Harvey, sp. nov., G. necopinus Harvey, sp. nov., G. postlei Harvey, sp. nov., G. ranalliorum Harvey, sp. nov. and G. weipa Harvey, sp. nov. from northern Australia, G. dissitus Harvey, sp. nov. from Cocos-Keeling Island, G. reong Harvey, sp. nov. and G. yeni Harvey, sp. nov. from Indonesia. A further possible new species from Queensland is described but not named, as it is represented by a single tritonymph. The subspecies of the Caribbean species G. bonairensis Beier are elevated to full species status: G. bonairensis, G. realini Hummelinck and G. withi Hoff. We supplement the descriptions with sequence data from five specimens from four species of Garypus and two species of Anchigarypus, and find COI divergence levels of 7–19% between Garypus species.

http://zoobank.org/References/16463E29-6F13-4392-9E41-46A4312C852B

Additional keywords: morphology, new species, taxonomy.


References

Álvarez-Padilla, F., and Hormiga, G. (2007). A protocol for digesting internal soft tissues and mounting spiders for scanning electron microscopy. The Journal of Arachnology 35, 538–542.
A protocol for digesting internal soft tissues and mounting spiders for scanning electron microscopy.Crossref | GoogleScholarGoogle Scholar |

Ashmole, N. P., and Ashmole, M. J. (1997). The land fauna of Ascension Island: new data from caves and lava flows, and a reconstruction of the prehistoric ecosystem. Journal of Biogeography 24, 549–589.
The land fauna of Ascension Island: new data from caves and lava flows, and a reconstruction of the prehistoric ecosystem.Crossref | GoogleScholarGoogle Scholar |

Beier, M. (1930). Die Pseudoskorpione des Wiener Naturhistorischen Museums. III. Annalen des Naturhistorischen Museums in Wien 44, 199–222.

Beier, M. (1932). Pseudoscorpionidea I. Subord. Chthoniinea et Neobisiinea. Das Tierreich 57, i–xx, 1–258.

Beier, M. (1936). Zoologische Ergebnisse einer Reise nach Bonaire, Curaçao und Aruba im Jahre 1930. No. 21. Einige neue neotropische Pseudoscorpione. Zoologische Jahrbucher. Abteilung fur Systematik, Ökologie und Geographie der Tiere 67, 443–447.

Beier, M. (1952). Eine neue Garypus-Art (Pseudoscorp.) aus Japan. Zoologischer Anzeiger 149, 235–239.

Beier, M. (1957). Pseudoscorpionida. Insects of Micronesia 3, 1–64.

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

Beier, M. (1961). Pseudoscorpione von der Insel Ascension. Annals & Magazine of Natural History 3, 593–598.

Beier, M. (1963). Ordnung Pseudoscorpionidea (Afterskorpione). In ‘Bestimmungsbücher zur Bodenfauna Europas. Vol. 1’. (Akademie-Verlag: Berlin, Germany.)

Beier, M. (1973). Pseudoscorpionidea von Ceylon. Entomologica Scandinavica 4, 39–55.

Benavides, L. R., Cosgrove, J. G., Harvey, M. S., and Giribet, G. (2019). Phylogenomic interrogation resolves the backbone of the Pseudoscorpiones Tree of Life. Molecular Phylogenetics and Evolution. 139, 106509.
Phylogenomic interrogation resolves the backbone of the Pseudoscorpiones Tree of Life.Crossref | GoogleScholarGoogle Scholar | 31132522PubMed |

Berland, L. (1929). Arachnides et Crustacès. In ‘Faune de la France en Tableaux Synoptiques. Vol. 2’. (Ed. R. Perrier.) pp. 1–220. (Librairie Delagrave: Paris, France.)

Boissin, L. (1974). Étude ultrastructurale de la spermiogenèse de Garypus beauvoisi (Sav.) (Arachnides, Pseudoscorpions). Archives de Zoologie Expérimentale et Générale 115, 169–184.

Bouvier, E. A. (1896). Sur la ponte et le développement d’un Pseudoscorpionide, le Garypus saxicola, Waterhouse Bulletin de la Société Entomologique de France 1, 304–307, 342–343.

Brown, R. W. (1956). ‘Composition of Scientific Words’, rev. edn. (Smithsonian Institution Press: Washington, DC, USA.)

Callaini, G., and Dallai, R. (1984). Spermatozoïdes et phylogenèse chez les Garypides (Arachnida, Pseudoscorpions). Revue Arachnologique 4, 335–342.

Callaini, G., and Dallai, R. (1989). Les spermatozoïdes des Pseudoscorpions: étude comparative et considérations phylogénétiques. Revue Arachnologique 8, 85–97.

Castresana, J. (2000). Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17, 540–552.
Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis.Crossref | GoogleScholarGoogle Scholar | 10742046PubMed |

Chamberlin, J. C. (1924). Giant Garypus of the Gulf of California. Nature Magazine 2, 171–172, 175.

Chamberlin, J. C. (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 & Magazine of Natural History 5, 1–48–585–620.
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).Crossref | GoogleScholarGoogle Scholar |

Chamberlin, J. C. (1931). The arachnid order Chelonethida. Stanford University Publications, Biological Sciences 7(1), 1–284.

Chamberlin, J. C. (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.
The taxonomy of the false scorpion genus Synsphyronus with remarks on the sporadic loss of stability in generally constant morphological characters (Arachnida: Chelonethida).Crossref | GoogleScholarGoogle Scholar |

Choi, E.-H., and Hwang, U.-W. (2009). First record of maritime pseudoscorpion Garypus japonicus (Garypidae) from Korea. Korean Journal of Systematic Zoology 25, 261–264.

Duffey, E. (1964). The terrestrial ecology of Ascencion Island. Journal of Applied Ecology 1, 219–251.
The terrestrial ecology of Ascencion Island.Crossref | GoogleScholarGoogle Scholar |

Ellingsen, E. (1914). On the pseudoscorpions of the Indian Museum, Calcutta. Records of the Indian Museum 10, 1–14.

Firstman, B. (1973). The relationship of the chelicerate arterial system to the evolution of the endosternite. The Journal of Arachnology 1, 1–54.

Folmer, O., Black, M., Hoeh, W., Lutz, R., and Vrijenhoek, R. C. (1994). DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3, 294–299.
| 7881515PubMed |

Gardiner, J. S. (1901). Introduction. Narrative and route of the expedition. In ‘The Fauna and Geography of the Maldive and Laccadive Archipelagoes: Being an Account of the Work Carried on and of the Collections Made by an Expedition during the Years 1899 and 1900. Vol. 1’. (Ed. J. S. Gardiner.) pp. 1–11. (Cambridge University Press: Cambridge, UK.)

Giribet, G., Carranza, S., Baguñà, J., Riutort, M., and Ribera, C. (1996). First molecular evidence for the existence of a Tardigrada + Arthropoda clade. Molecular Biology and Evolution 13, 76–84.
First molecular evidence for the existence of a Tardigrada + Arthropoda clade.Crossref | GoogleScholarGoogle Scholar | 8583909PubMed |

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

Harvey, M. S. (1987). A revision of the genus Synsphyronus Chamberlin (Garypidae: Pseudoscorpionida: Arachnida). Australian Journal of Zoology Supplementary Series 35, 1–99.
A revision of the genus Synsphyronus Chamberlin (Garypidae: Pseudoscorpionida: Arachnida).Crossref | GoogleScholarGoogle Scholar |

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

Harvey, M. S. (1992). The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy 6, 1373–1435.
The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida).Crossref | GoogleScholarGoogle Scholar |

Harvey, M. S. (2016). The systematics of the pseudoscorpion family Ideoroncidae (Pseudoscorpiones, Neobisioidea) in the Asian region. The Journal of Arachnology 44, 272–329.
The systematics of the pseudoscorpion family Ideoroncidae (Pseudoscorpiones, Neobisioidea) in the Asian region.Crossref | GoogleScholarGoogle Scholar |

Harvey, M. S., and Leng, M. C. (2008). The first troglomorphic pseudoscorpion of the family Olpiidae (Pseudoscorpiones), with remarks on the composition of the family. Records of the Western Australian Museum 24, 387–394.
The first troglomorphic pseudoscorpion of the family Olpiidae (Pseudoscorpiones), with remarks on the composition of the family.Crossref | GoogleScholarGoogle Scholar |

Harvey, M. S., and Muchmore, W. B. (2013). The systematics of the pseudoscorpion family Ideoroncidae (Pseudoscorpiones: Neobisioidea) in the New World. The Journal of Arachnology 41, 229–290.
The systematics of the pseudoscorpion family Ideoroncidae (Pseudoscorpiones: Neobisioidea) in the New World.Crossref | GoogleScholarGoogle Scholar |

Harvey, M. S., Ratnaweera, P. B., Udagama, P. V., and Wijesinghe, M. R. (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.
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.Crossref | GoogleScholarGoogle Scholar |

Harvey, M. S., Huey, J. A., Hillyer, M. J., McIntyre, E., and Giribet, G. (2016). 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.
The first troglobitic species of Gymnobisiidae (Pseudoscorpiones, Neobisioidea), from Table Mountain (Western Cape Province, South Africa) and its phylogenetic position.Crossref | GoogleScholarGoogle Scholar |

Heurtault, J., and Vannier, G. (1990). Thermorésistance chez deux pseudoscorpions (Garypidae), l’un du désert de Namibie, l’autre de la région de Gênes (Italie). Acta Zoologica Fennica 190, 165–171.

Hoff, C. C. (1947). New species of diplosphyronid pseudoscorpions from Australia. Psyche, Cambridge 54, 36–56.
New species of diplosphyronid pseudoscorpions from Australia.Crossref | GoogleScholarGoogle Scholar |

Huelsenbeck, J. P., and Ronquist, F. (2001). MRBAYES: Bayesian inference of phylogeny. Bioinformatics 17, 754–755.
MRBAYES: Bayesian inference of phylogeny.Crossref | GoogleScholarGoogle Scholar | 11524383PubMed |

Hummelinck, P. W. (1948). Studies on the fauna of Curaçao, Aruba, Bonaire and the Venezuelan Islands: No. 13. Pseudoscorpions of the genera Garypus, Pseudochthonius, Tyrannochthonius and Pachychitra. Natuurwetenschappelijke Studiekring voor Suriname en Curaçao 5, 29–77.

Judson, M. L. I. (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.
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).Crossref | GoogleScholarGoogle Scholar |

Katoh, K., and Standley, D. M. (2013). MAFFT multiple sequence alignment software Version 7: improvements in performance and usability. Molecular Biology and Evolution 30, 772–780.
MAFFT multiple sequence alignment software Version 7: improvements in performance and usability.Crossref | GoogleScholarGoogle Scholar | 23329690PubMed |

Kearse, M., Moir, R., Wilson, A. C., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P., and Drummond, A. J. (2012). Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28, 1647–1649.
Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data.Crossref | GoogleScholarGoogle Scholar | 22543367PubMed |

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

Kumar, S., Stecher, G., and Tamura, K. (2016). MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33, 1870–1874.
MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets.Crossref | GoogleScholarGoogle Scholar | 27004904PubMed |

Lanfear, R., Calcott, B., Ho, S. Y. W., and Guindon, S. (2012). PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29, 1695–1701.
PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses.Crossref | GoogleScholarGoogle Scholar | 22319168PubMed |

Lanfear, R., Calcott, B., Kainer, D., Mayer, C., and Stamatakis, A. (2014). Selecting optimal partitioning schemes for phylogenomic datasets. BMC Evolutionary Biology 14, 82.
Selecting optimal partitioning schemes for phylogenomic datasets.Crossref | GoogleScholarGoogle Scholar | 24742000PubMed |

Lee, V. F. (1979). The maritime pseudoscorpions of Baja California, México (Arachnida: Pseudoscorpionida). Occasional Papers of the California Academy of Sciences 131, i–iv, 1–38.

Lighton, J. R. B., and Joos, B. (2002a). Discontinuous gas exchange in a tracheate arthropod, the pseudoscorpion Garypus californicus: occurrence, characteristics and temperature dependence. Journal of Insect Science 2, 1–4.
Discontinuous gas exchange in a tracheate arthropod, the pseudoscorpion Garypus californicus: occurrence, characteristics and temperature dependence.Crossref | GoogleScholarGoogle Scholar |

Lighton, J. R. B., and Joos, B. (2002b). Discontinuous gas exchange in the pseudoscorpion Garypus californicus is regulated by hypoxia, not hypercapnia. Physiological and Biochemical Zoology 75, 345–349.
Discontinuous gas exchange in the pseudoscorpion Garypus californicus is regulated by hypoxia, not hypercapnia.Crossref | GoogleScholarGoogle Scholar |

Mahnert, V. (1979). Pseudoskorpione (Arachnida) aus dem Amazonas-Gebiet (Brasilien). Revue Suisse de Zoologie 86, 719–810.
Pseudoskorpione (Arachnida) aus dem Amazonas-Gebiet (Brasilien).Crossref | GoogleScholarGoogle Scholar |

Mahnert, V. (1981). Die Pseudoskorpione (Arachnida) Kenyas. I. Neobisiidae und Ideoroncidae. Revue Suisse de Zoologie 88, 535–559.
Die Pseudoskorpione (Arachnida) Kenyas. I. Neobisiidae und Ideoroncidae.Crossref | GoogleScholarGoogle Scholar |

Mahnert, V. (1982). Die Pseudoskorpione (Arachnida) Kenyas, IV. Garypidae. Annales Historico-Naturales Musei Nationalis Hungarici 74, 307–329.

Mahnert, V. (1984). Forschungen an der Somalilandküste. Am Strand und auf den Dünen bei Sar Uanle. 36. Pseudoscorpiones (Arachnida). Monitore Zoologico Italiano, n.s. Supplemento 19, 43–66.
Forschungen an der Somalilandküste. Am Strand und auf den Dünen bei Sar Uanle. 36. Pseudoscorpiones (Arachnida).Crossref | GoogleScholarGoogle Scholar |

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

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

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

Mahnert, V., and 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.
Pachyolpium atlanticum n. sp., ein Pseudoskorpion aus der Gezeitenzone der Bermudas – Morphologie und Ökologie (Pseudoscorpiones: Olpiidae).Crossref | GoogleScholarGoogle Scholar |

Makioka, T. (1968). Morphological and histochemical studies on embryos and ovaries during the embryo-breeding of the pseudoscorpion, Garypus japonicus. Science Report of the Tokyo Kyoiku Daigaku (B) 13, 207–227.

Makioka, T. (1969). A temporary gonopodium in a pseudoscorpion, Garypus japonicus. Science Report of the Tokyo Kyoiku Daigaku (B) 14, 113–120.

Makioka, T. (1976). Alternative occurrence of two ovarian functions in the adult pseudoscorpion, Garypus japonicus Beier. Acta Arachnologica 27, 8–15.
Alternative occurrence of two ovarian functions in the adult pseudoscorpion, Garypus japonicus Beier.Crossref | GoogleScholarGoogle Scholar |

Makioka, T. (1979). Structures of the adult ovaries in different functional phases of the pseudoscorpion, Garypus japonicus Beier. I. The ovary in the resting phase. Acta Arachnologica 28, 71–81.
Structures of the adult ovaries in different functional phases of the pseudoscorpion, Garypus japonicus Beier. I. The ovary in the resting phase.Crossref | GoogleScholarGoogle Scholar |

Muchmore, W. B. (1991). Pseudoscorpionida. In ‘Diversidad Biológica en la Reserva de la Biosfera de Sian Ka’an Quintana Roo, México’. (Eds D. Navarro L., and J. G. Robinson.) pp. 155–173. (Centro de Investigaciones de Quintana Roo, México: Chetumal, Mexico.)

Murienne, J., Harvey, M. S., and Giribet, G. (2008). First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata). Molecular Phylogenetics and Evolution 49, 170–184.
First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata).Crossref | GoogleScholarGoogle Scholar | 18603009PubMed |

Murthy, V. A., and Ananthakrishnan, T. N. (1977). Indian Chelonethi. Oriental Insects Monograph 4, 1–210.

Nassirkhani, M. (2015). Notes on Olpiidae (Arachnida: Pseudoscorpiones) from Iran: description of Cardiolpium bisetosum sp. nov. and redescription of Olpium omanense. Arachnologische Mitteilungen 50, 1–10.
Notes on Olpiidae (Arachnida: Pseudoscorpiones) from Iran: description of Cardiolpium bisetosum sp. nov. and redescription of Olpium omanense.Crossref | GoogleScholarGoogle Scholar |

Pocock, R. I. (1904). Arachnida. In ‘The Fauna and Geography of the Maldive and Laccadive Archipelagoes: Being an Account of the Work Carried on and of the Collections Made by an Expedition During the Years 1899 and 1900. Vol. 2’. (Ed. J. S. Gardiner.) pp. 797–805. (Cambridge University Press: Cambridge, UK.)

Poinar, G. O., Thomas, G. M., and Lee, V. F. (1985). Laboratory infection of Garypus californicus (Pseudoscorpionida, Garypidae) with neoaplectanid and heterorhabditid nematodes (Rhabditoidea). The Journal of Arachnology 13, 400–401.

Redikorzev, V. (1926). Pseudoscorpion nouveaux du Caucase. Entomologicheskoe obozrenie (Revue Russe d’Entomologie) 20, 1–4.

Rix, M. G., Harvey, M. S., and Roberts, J. D. (2010). A revision of the textricellin spider genus Raveniella (Araneae: Araneoidea: Micropholcommatidae): exploring patterns of phylogeny and biogeography in an Australian biodiversity hotspot. Invertebrate Systematics 24, 209–237.
A revision of the textricellin spider genus Raveniella (Araneae: Araneoidea: Micropholcommatidae): exploring patterns of phylogeny and biogeography in an Australian biodiversity hotspot.Crossref | GoogleScholarGoogle Scholar |

Rix, M. G., Cooper, S. J. B., Meusemann, K., Klopfstein, S., Harrison, S. E., Harvey, M. S., and Austin, A. D. (2017). Post-Eocene climate change across continental Australia and the diversification of Australasian spiny trapdoor spiders (Idiopidae: Arbanitinae) Molecular Phylogenetics and Evolution 109, 302–320.
Post-Eocene climate change across continental Australia and the diversification of Australasian spiny trapdoor spiders (Idiopidae: Arbanitinae)Crossref | GoogleScholarGoogle Scholar | 28126515PubMed |

Sato, H. (1978). [Notes on the Japanese maritime pseudoscorpions, Garypus japonicus and Nipponogarypus enoshimaensis]. Atypus 71, 45–48.

Sato, H. (1980). [Influence of humidity on three pseudoscorpions Microcreagris japonica, Garypus japonicus and Haplochernes boncicus]. Memoirs of the Education Institute for Private Schools in Japan 72, 57–63.

Scherzer, K.v. (1861). ‘Reise der osterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair.’ (Kaiserlich-koniglichen Hof- und Staatsdruckerei: Vienna, Austria.)

Schulte, G. (1976). Litoralzonierung von Pseudoskorpionen an der nordamerikanischen Pazifikküste (Arachnida: Pseudoscorpiones: Neobisiidae, Garypidae). Entomologica Germanica 3, 119–124.

Simon, E. (1879). Les Ordres des Chernetes, Scorpiones et Opiliones. In ‘Les Arachnides de France. Vol. 7’. pp. 1–332. (Librairie Encyclopédique de Roret: Paris, France.)

Stamatakis, A. (2006). RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22, 2688–2690.
RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models.Crossref | GoogleScholarGoogle Scholar | 16928733PubMed |

Talavera, G., and Castresana, J. (2007). Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments. Systematic Biology 56, 564–577.
Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments.Crossref | GoogleScholarGoogle Scholar | 17654362PubMed |

Tullgren, A. (1907). Zur Kenntnis aussereuropäischer Chelonethiden des Naturhistorischen Museums in Hamburg. Mitteilungen aus dem Naturhistorischen Museum in Hamburg 24, 21–75.

Vachon, M. (1936). Sur le développement postembryonnaire des Pseudoscorpions (quatrième note). Les formules chaetotaxiques des pattes-mâchoires. Bulletin du Muséum National d’Histoire Naturelle 8, 77–83.

Vachon, M. (1937). Pseudoscorpions nouveaux des collections du Muséum National d’Histoire Naturelle de Paris. (3e note). Bulletin de la Société Entomologique de France 42, 188–190.

Vachon, M. (1938). Recherches anatomiques et biologiques sur la réproduction et le développement des Pseudoscorpions. Annales des Sciences Naturelles, Zoologie 1, 1–207.

Weygoldt, P. (1970). Vergleichende Untersuchungen zur Fortpflanzungsbiologie der Pseudoscorpione. II. Zeitschrift für die Zoologische Systematik und Evolutionforschung 8, 241–259.
Vergleichende Untersuchungen zur Fortpflanzungsbiologie der Pseudoscorpione. II.Crossref | GoogleScholarGoogle Scholar |

Weygoldt, P. (1971). Vergleichend-embryologische Untersuchungen an Pseudoscorpionen. V. Das Embryonalstadium mit seinem Pumporgan bei verschiedenen Arten und sein Wert als taxonomisches Merkmal. Zeitschrift für die Zoologische Systematik und Evolutionforschung 9, 3–29.
Vergleichend-embryologische Untersuchungen an Pseudoscorpionen. V. Das Embryonalstadium mit seinem Pumporgan bei verschiedenen Arten und sein Wert als taxonomisches Merkmal.Crossref | GoogleScholarGoogle Scholar |

Whiting, M. F., Carpenter, J. M., Wheeler, Q. D., and Wheeler, W. C. (1997). The Strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology. Systematic Biology 46, 1–68.
The Strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology.Crossref | GoogleScholarGoogle Scholar | 11975347PubMed |