Abstract
A taxonomic revision of the subgenus Chthonius (Ephippiochthonius) in the Iberian Peninsula, Balearic Islands and Macaronesia is presented. New and previously insufficiently studied characteristics are shown to be taxonomically useful, such as the chelal lyrifissures patterns, chaetotaxy and condylar complex. Three taxa previously treated as subgenera of Chthonius are here raised to generic rank: Ephippiochthonius n. stat., Globochthonius n. stat., and Hesperochthonius n. stat. Two new genera are described: Cantabrochthonius n. gen. and Occidenchthonius n. gen. Thirty-five new species are described: Ephippiochthonius aini n. sp., E. andalucia n. sp., E. aurouxi n. sp., E. benimaquia n. sp., E. caceresi n. sp., E. castellonensis n. sp., E. comasi n. sp., E. fadriquei n. sp., E. galcerani n. sp., E. gonzalezi n. sp., E. henderickxi n. sp., E. ibiza n. sp., E. masoae n. sp., E. portugalensis n. sp., E. riberai n. sp., E. serengei n. sp., E. sevai n. sp., E. tarraconensis n. sp., E. vicenae n. sp., E. zaballosi n. sp., Occidenchthonius anae n. sp., O. beieri n. sp., O. ebo n. sp., O. felgueraorum n. sp., O. gardinii n. sp., O. hoerwegi n. sp., O. lencinai n. sp., O. mahnerti n. sp., O. mateui n. sp., O. montagudi n. sp., O. murcia n. sp., O. oromii n. sp., O. ortunoi n. sp., O. riopar n. sp. and O. serranoi n. sp. A neotype is designated for Ephippiochthonius catalonicus (Beier, 1939), n. comb. As result of the changes in generic rank, 45 new combinations for species are proposed.
References
Beier, M. (1930) Neue Höhlen-Pseudoscorpione der Gattung Chthonius. Eos, 6, 323–327.
Beier, M. (1931) Zur Kenntnis der Chthoniiden (Pseudoskorpione). Zoologischer Anzeiger, 93, 49–56.
Beier, M. (1932) Pseudoscorpionidea I. Subord. Chthoniinea et Neobisiinea. Tierreich, 57, i–xx, 1–258.
https://doi.org/10.1515/9783111435107.1Beier, M. (1934) Neue cavernicole und subterrane Pseudoscorpione. Mitteilungen über Höhlen- und Karstforschung, 1934, 53–59.
Beier, M. (1938) Vorläufige Mitteilung über neue Höhlenpseudoscorpione der Balkanhalbinsel. Studien aus dem Gebiete der Allgemeinen Karstforschung, der Wissenschaftlichen Höhlenkunde, der Eiszeitforschung und den Nachbargebieten, 3 (8), 5–8.
Beier, M. (1939) Die Pseudoscorpioniden-Fauna der iberischen Halbinsel. Zoologische Jahrbücher, Abteilung für Systematik Ökologie und Geographie der Tiere, 72, 157–202.
Beier, M. (1951) Die Pseudoscorpione Indochinas. Memoires du Museum National d'Histoire Naturelle, Paris, Nouvelle Serie, 1, 47–123.
Beier, M. (1953) Weiteres zur Kenntnis der iberischen Pseudoscorpioniden-Fauna. Eos, Madrid, 28, 293–302.
Beier, M. (1955) Neue Beiträge zur Kenntnis der iberischen Pseudoscorpioniden-Fauna. Eos, Madrid, 31, 87–122.
Beier, M. (1963) Ordnung Pseudoscorpionidea (Afterskorpione). Bestimmungsbücher zur Bodenfauna Europas. Vol. 1. Akademie-Verlag, Berlin, 313 pp.
Beier, M. (1965) Über Pseudoskorpione von den Kanaren. Annalen des Naturhistorischen Museums in Wien, 68, 375–381.
Beier, M. (1975a). Weitere bemerkenswerte Pseudoscorpione von Sizilien. Animalia, Catania, 2, 55–58.
Bellés, X. (2015).Joaquim Mateu (1921–2015), tota una vida dedicada a l’estudi dels insects. Museu de Ciències Naturals de Barcelona. El Blog del Museu. Available from: http://www.blogmuseuciencies.org/2015/04/joaquim-mateu-1921-2015-tota-una-vida-dedicada-a-lestudi-dels-insectes/ (accessed 30 December 2015)
Benedict, E.M. (1978) A new pseudoscorpion genus Malcolmochthonius n.g., with three new species from the western United States. Transactions of the American Microscopical Society, 97, 250–255.
https://doi.org/10.2307/3225602Benedict, E.M. & Malcolm, D.R. (1973) A new cavernicolous species of Apochthonius (Chelonethida: Chthoniidae) from the western United States with reference to troglobitic tendencies in the genus. Transactions of the American Microscopical Society, 92, 620–628.
https://doi.org/10.2307/3225272
Callaini, G. (1984) Osservazioni su alcune specie di Chthonius del sottogenere Ephippiochthonius Beier (Arachnida, Pseudoscorpionida, Chthoniidae). Notulae Chernetologicae XVII. Annali del Museo Civico di Storia Naturale di Genova, 85, 125–159.
Carabajal Márquez, E., García Carrillo, J. & Rodríguez Fernández, F. (2000) Descripción de dos nuevas especies de pseudoscorpiones cavernícolas de la provincia de Cádiz (Arachnida, Pseudoscorpionida, Chthoniidae, Neobisiidae). Graellsia, 58, 27–33.
https://doi.org/10.3989/graellsia.2000.v56.i0.307Carabajal Márquez, E., García Carrillo, J. & Rodríguez Fernández, F. (2001a) Nuevos pseudoscorpiones cavernícolas de la Sierra de Gádor (Almeria, España) (Arachnida, Pseudoscorpionida, Chthoniidae). Revista Ibérica de Aracnología, 3, 7–15.
Carabajal Márquez, E., García Carrillo, J. & Rodríguez Fernández, F. (2001b) Descripción de cuatro nuevos pseudoscorpiones cavernícolas de Andalucía, España (Arachnida, Pseudoscorpionida, Chthoniidae). Zoologica Baetica, 12, 169–184.
Carabajal Márquez, E., García Carrillo, J. & Rodríguez Fernández, F. (2011) Aportaciones al catálogo de los Pseudoscorpiones de Andalucía (España) (I) (Arachnida, Pseudoscorpiones). Boletín de la Sociedad Entomológica Aragonesa, 48, 115–128.
Carabajal Márquez, E., García Carrillo, J. & Rodríguez Fernández, F. (2012) Aportaciones al catálogo de los Pseudoscorpiones de Andalucía (España) (II) (Arachnida, Pseudoscorpiones, Chthoniidae) Descripción de tres nuevas especies de Chthonius C.L. Koch, 1843 de Cádiz, Málaga y Almería (España). Revista Ibérica de Aracnología, 21, 89–95.
Chamberlin, J.C. (1929a) A synoptic classification of the false scorpions or chela-spinners, with a report on a cosmopolitan collection of the same.—Part I. The Heterosphyronida (Chthoniidae) (Arachnida-Chelonethida). Annals and Magazine of Natural History, Series 10, 4, 50–80.
https://doi.org/10.1080/00222932908673028Chamberlin, J.C. (1929b) On some false scorpions of the suborder Heterosphyronida (Arachnida - Chelonethida). Canadian Entomologist, 61, 152–155.
https://doi.org/10.4039/Ent61152-7Chamberlin, J.C. (1931) The arachnid order Chelonethida. Stanford University Publications, Biological Sciences, 7 (1), 1–284.
Chamberlin, J.C. (1949) New and little-known false scorpions from various parts of the world (Arachnida, Chelonethida), with notes on structural abnormalities in two species. American Museum Novitates, 1430, 1–57.
Chamberlin, J.C. (1962) New and little-known false scorpions, principally from caves, belonging to the families Chthoniidae and Neobisiidae. Bulletin of the American Museum of Natural History, 123, 303–352.
Ćurčić, B.P.M. (1973) Le sous-genre Globochthonius Beier 1931 dans la Méditerranée nord-occidentale: Chthonius (G.) globifer Simon 1879 (Chthoniidae, Pseudoscorpiones, Arachnida). Wissenschaftliche Mitteilungen des Bosnisch-herzegovinischen Landesmuseums, 3 (C), 77–84.
Ćurčić, B.P.M. (1974) The subgenus Globochthonius Beier, 1931 (Chthoniidae, Pseudoscorpiones): taxonomic considerations and biogeographic implications. Glasnik de l'Académie Serbe des Sciences et des Arts, Classe de Sciences Mathématiques et Naturelles, 36, 105–112.
Ćurčić, B.P.M., Rađa, T., Dimitrijević, R.N., Ćurčić, S.B., Ćurčić, N.B., Makarov, S.E. & Ilić, B.S. (2014) Microchthonius kasteli n. sp. (Chthoniidae, Pseudoscorpiones) – A new cave false scorpion from Croatia (Dalmatia). Archives of Biological Science, 66 (1), 437–443.
https://doi.org/10.2298/ABS1401437CĆurčić, B.P.M., Rađa, T., Dimitrijević, R.N., Vesović, N. & Ćurčić, S. (2015) On two new species of Microchthonius (Pseudoscorpiones, Chthoniidae) from Dalmatia, Croatia. Scientific Bulletin of the University of Shkodra “Luigj Gurakuqi”, 65, 80–93.
Edward, K.L. & Harvey, M.S. (2008) Short-range endemism in hypogean environments: the pseudoscorpion genera Tyrannochthonius and Lagynochthonius (Pseudoscorpiones: Chthoniidae) in the semiaird zone of Western Australia. Invertebrate Systematics, 22, 259–293.
https://doi.org/10.1071/IS07025
Fernández-Palacios, J.M. (2009) El relictualismo en islas oceánicas. El caso de la laurisilva macaronésica. In: Real, R. & Márquez, A.L. (Eds.), Biogeografia: Scientia Biodiversitatis. Universidad de Málaga, Málaga, pp. 13–24.
Fernández-Palacios, J.M. (2011) The islands of Macaronesia. In: Serrano, A.R.M., Borges, P.A.V., Boieiro, M. & Oromí, P. (Eds.), Terrestrial arthropods of Macaronesia. Biodiversity, ecology and evolution. Sociedad Portuguesa de Entomología, Lisbon, pp. 1–30.
Fernández-Palacios, J.M., Nascimento, L. de, Otto, R., Delgado, J.D., García-del-Rey, E., Arévalo, J.R. & Whittaker, R.J. (2011) A reconstruction of Palaeo-Macaronesia, with particular reference to the long-term biogeography of the Atlantic island laurel forests. Journal of Biogeography, 38, 226–246.
https://doi.org/10.1111/j.1365-2699.2010.02427.x
Gabbutt, P.D. & Vachon, M. (1963) The external morphology and life history of the pseudoscorpion Chthonius ischnocheles (Hermann). Proceedings of the Zoological Society of London, 140, 75–98.
https://doi.org/10.1111/j.1469-7998.1963.tb01855.x
Gardini, G. (1981) Pseudoscorpioni cavernicoli sardi. I. Chthoniidae (Pseudoscorpioni d’Italia, X). Revue arachnologique, 3, 101–114.
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.
Gardini, G. (1994) I generi Paraliochthonius Beier, 1956 e Spelyngochthonius Beier, 1955 in Italia (Pseudoscorpionida, Chthoniidae). Fragmenta Entomologica, Roma, 26, 1–10.
Gardini, G. (2000) Catalogo degli Pseudoscorpioni d´Italia. Fragmenta Entomologica, Roma, 32 (Supplement), 1–181.
Gardini, G. (2009) Neotype fixation and redescription of Chthonius tetrachelatus (Preyssler, 1790), type species of the subgenus Ephippiochthonius Beier, 1930 (Pseudoscorpiones: Chthoniidae). Klapalekiana, 45, 23–31.
Gardini, G. (2010) Gli Chthonius (Globochthonius) d’Italia e di Slovenia (Pseudoscorpiones Chthoniidae). Gortania, 31, 53–64.
Gardini, G. (2013) A revision of the species of the pseudoscorpion subgenus Chthonius (Ephippiochthonius) (Arachnida, Pseudoscorpiones, Chthoniidae) from Italy and neighbouring areas. Zootaxa, 3655 (1), 1–151.
https://doi.org/10.11646/zootaxa.3655.1.1Gardini, G. (2014) The species of the Chthonius heterodactylus group (Arachnida, Pseudoscorpiones, Chthoniidae) from the eastern Alps and the Carpathians. Zootaxa, 3887 (2), 101–137.
https://doi.org/10.11646/zootaxa.3887.2.1Gardini, G. & Benelli, R. (1991) The external morphology of the pseudoscorpion Roncus andreinii. Journal of Zoology, London, 223, 27–48.
Geldmacher, J., Hoernle, K., van den Bogaard, P., Duggen, S. & Werner, R. (2005) New 40K/39Ar age and geochemical data from seamounts in the Canary and Madeira volcanic provinces: support for the mantle plume hypothesis. Earth and Planetary Science Letters, 237, 85–101.
https://doi.org/10.1016/j.epsl.2005.04.037Hadži, J. (1930) [Contribution à la connaissance des Pseudoscorpions cavernicoles]. Glasnik Académie Royale Serbe, Belgrade, 67, 115–148. [in Serbian, with French summary]
Hadži, J. (1933). Prinos poznavanju pseudoskorpijske faune Primorja. Prirodoslovna Istraživanja Kraljevine Jugoslavije, 18, 125‒192.
Hansen, H.J. (1893‒1894) Organs and characters in different orders of arachnids. Entomologiske Meddelelser, 4, 137–251.
Harvey, M.S. (1991) Catalogue of the Pseudoscorpionida. Manchester University Press, Manchester, vi + 726 pp.
Harvey, M.S. (1992) The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy, 6, 1373–1435.
https://doi.org/10.1071/IT9921373
Harvey, M.S. (2013) Pseudoscorpions of the World, version 3.0. Western Australian Museum, Perth. Available from: http://www.museum.wa.gov.au/catalogues-beta/pseudoscorpions (accessed 30 December 2015)
Harvey, M.S., Barba Díaz, R., Muchmore, W.B. & Peréz González, A. (2007) Pseudalbiorix, a new genus of Ideoroncidae (Pseudoscorpiones, Neobisioidea) from central America. Journal of Arachnology, 34, 610–626.
https://doi.org/10.1636/S05-28.1
Harvey, M.S. & Du Preez, G. (2014) A new troglobitic ideoroncid pseudoscorpion (Pseudoscorpiones: Ideoroncidae) from southern Africa. The Journal of Arachnology, 42, 105–110.
https://doi.org/10.1636/K13-55.1
Harvey, M.S. & Mould, L.G. (2006) A new troglomorphic species of Austrochthonius (Pseudoscorpiones: Chthoniidae) from Australia, with remarks on Chthonius caecus. Records of the Western Australian Museum, 23, 205–211.
https://doi.org/10.18195/issn.0312-3162.23(2).2006.205-211Harvey, M.S. & Muchmore, W.B. (2013) The systematics of the pseudoscorpion family Ideoroncidae (Pseudoscorpiones: Neobisioidea) in the New World. Journal of Arachnology, 41, 229–290.
https://doi.org/10.1636/K13-42.1
Heurtault, J. (1972) Chthonius (C.) petrochilosi (Arachnide, Pseudoscorpion, Chthoniidae), nouvelle espèce cavernicole de Grèce. Biologia Gallo-Hellenica, 4, 19–25.
Heurtault, J. (1975) Deux nouvelles espèces de pseudoscorpions Chthoniidae (Arachnides) cavernicoles de Corse: Chthonius (E.) remyi et Chthonius (E.) siscoensis. Annales de Spéléologie, 30, 313–318.
Heurtault-Rossi, J. (1968) Quelques remarques sur deux espèces cavernicoles de Chthonius des départements des Bouches-du Rhône et du Gard: Chthonius (C.) cephalotes (Simon, 1875) et Chthonius mayi sp. nov. (Pseudoscorpions, Chthoniidae). Bulletin du Muséum National d'Histoire Naturelle, Paris, Series 2, 39, 912–922.
Hoff, C.C. (1949) The pseudoscorpions of Illinois. Bulletin of the Illinois Natural History Survey, 24, 407–498.
Judson, M.L.I. (1990a) On the presence of Chthonius (C.) halberti Kew and Chthonius (C.) ressli Beier in France with remarks on the status of Kewochthonius Chamberlin and Neochthonius Chamberlin (Arachnida, Chelonethida, Chthoniidae). Bulletin du Muséum national d’Histoire naturelle, Paris, Series 4, 11, 593–603.
Judson, M.L.I. (1990b) Observations on the form and function of the coxal spines of some chthonioid pseudoscorpions from Cameroon (Arachnida, Chelonethida). Acta Zoologica Fennica, 190, 195–198.
Judson, M.L.I. (1992) The pseudoscorpion subgenus Chthonius (Sigmodactylus) Hadži, 1930: type fixation and consequent synonymy with Pseudochthonius Balzan, 1892 (Chelonethi, Chthoniidae). Bulletin of the British Arachnological Society, 9, 64.
Judson, M.L.I. (2001) Synonymy of Cecoditha (Cecodithinae) with Austrochthonius (Chthoniinae) (Chelonethi, Chthoniidae). Journal of Arachnology, 29, 141–145.
https://doi.org/10.1636/0161-8202(2001)029[0141:SOCCWA]2.0.CO;2Judson, M.L.I. (2007) A new and endangered pseudoscorpion of the genus Lagynochthonius (Arachnida, Chelonethi, Chthoniidae) from a cave in Vietnam, with notes on chelal morphology and the composition of the Tyrannochthoniini. Zootaxa, 1627, 53–68.
Judson, M.L.I. (2016) Pseudoscorpions (Arachnida, Chelonethi) in Mexican amber, with a list of extant species associated with mangrove and Hymenaea trees in Chiapas. Boletín de la Sociedad Geológica Mexicana, 68 (1), 57–79. Available from: http://boletinsgm.igeolcu.unam.mx/bsgm/index.php/volumenes-volumes/cuarta-epoca/295-volumen-volume-68-2016/1414-volumen-68-numero-1-2016 (accessed 24 June 2016)
https://doi.org/10.5281/zenodo.50595
Judson, M.L.I. & Legg, G. (1996) Discovery of the pseudoscorpion Larca lata (Garypoidea, Larcidae) in Britain. Bulletin of the British Arachnological Society, 10 (6), 205–210.
Lagar, A. (1972a) Contribución al conocimiento de los Pseudoescorpiones de España. I. Miscelanea Zoologica, 3, 17–21.
Lagar, A. (1972b) Contribución al conocimiento de los Pseudoescorpiones de España. II. Speleon, 19, 45–52.
Leclerc, P. (1981) Nouveaux Chthoniidae cavernicoles de la bordure orientale des Cévennes (France) (Arachnides, Pseudoscorpions). Revue Arachnologique, 3, 115–131.
Legg, G. (1975) The genitalia and associated glands of five British species belonging to the family Chthoniidae (Pseudoscorpiones: Arachnida). Journal of Zoology, London, 177, 99–121.
https://doi.org/10.1111/j.1469-7998.1975.tb05973.x
Mahnert, V. (1977) Spanische Höhlenpseudoskorpione. Miscelanea Zoologica, 4, 61–104.
Mahnert, V. (1978). Pseudoskorpione (ausgenommen Olpiidae, Garypidae) aus Congo-Brazzaville (Arachnida, Pseudoscorpiones). Folia Entomologica Hungarica, 31, 69–133.
Mahnert, V. (1982) Neue höhlenbewohnende Pseudoskorpione aus Spanien, Malta und Griechenland (Arachnida, Pseudoscorpiones). Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 55, 297–304.
Mahnert, V. (1989) Les pseudoscorpions (Pseudoscorpiones, Arachnida) récoltés pendant la campagne biospéologique 1987 à Minorque. Endins, 14/15, 85–87.
Mahnert, V. (1993) Pseudoskorpione (Arachnida: Pseudoscorpiones) von Inseln des Mittelmeers und des Atlantiks (Balearen, Kanarische Inseln, Madeira, Ascensión), mit Vorwiegend subterraner Lebensweise. Revue suisse de Zoologie, 100 (4), 971–992.
https://doi.org/10.5962/bhl.part.79896
Mahnert, V. (1997) New species and records of pseudoscorpions (Arachnida, Pseudoscorpiones) from the Canary Islands. Revue suisse de Zoologie, 104 (3), 559–585.
https://doi.org/10.5962/bhl.part.80011
Mahnert, V. (2011a) A nature’s treasury: Pseudoscorpion diversity of the Canary Islands, with the description of nine new species (Pseudoscorpiones, Chthoniidae, Cheiridiidae) and new records. Revista Ibérica de Aracnología, 19, 27–45.
Mahnert, V. (2011b) New records of pseudoscorpions from the Juan Fernandez Islands (Chile), with the description of a new genus and three new species of Chernetidae (Arachnida: Pseudoscorpiones). Revue suisse de Zoologie, 118, 17‒29.
https://doi.org/10.5962/bhl.part.117795Mahnert, V. (2013) Nachträge zur Pseudoskorpionfauna (Arachnida: Pseudoscorpiones) der Höhlen der Insel Santorin (Thera) (Kykladen, Griechenland). Revue suisse de Zoologie, 120 (1), 93–100.
Mahnert, V. (2014) Pseudoscorpions (Arachnida: Pseudoscorpiones) from the Galapagos Islands (Ecuador). Revue suisse de Zoologie, 121 (2), 135–210.
Muchmore, W.B. (1965) North American cave pseudoscorpions of the genus Kleptochthonius, subgenus Chamberlinochthonius (Chelonethida, Chthoniidae). American Museum Novitates, 2234, 1–27.
Muchmore, W.B. (1968) Two new species of chthoniid pseudoscorpions from the western United States (Arachnida: Chelonethida: Chthoniidae). Pan-Pacific Entomologist, 44, 51–57.
Muchmore, W.B. (1969) The pseudoscorpion genus Neochthonius Chamberlin (Arachnida, Chelonethida, Chthoniidae) with description of a cavernicolous species. American Midland Naturalist, 81, 387‒394.
https://doi.org/10.2307/2423978Muchmore, W.B. (1974) New cavernicolous species of Kleptochthonius from Virginia and West Virginia (Pseudoscorpionida, Chthoniidae). Entomological News, 85, 81–84.
Muchmore, W.B. (1976) New cavernicolous species of Kleptochthonius, and recognition of a new species group within the genus (Pseudoscorpionida: Chthoniidae). Entomological News, 87, 211–217.
Muchmore, W.B. (1991) Pseudoscorpions from Florida and the Caribbean area. 14. New species of Tyrannochthonius and Lagynochthonius from caves in Jamacia [sic], with discussion of the genera (Chthoniidae). Florida Entomologist, 74, 110–121.
https://doi.org/10.2307/3495247
Muchmore, W.B. (1994) Three unusual new epigean species of Kleptochthonius (Pseudoscorpionida: Chthoniidae). Jeffersoniana, 6, 1–13.
Muchmore, W.B. (2000) New species and records of Kleptochthonius from Indiana (Pseudoscorpionida, Chthoniidae). Journal of Arachnology, 28, 293–299.
https://doi.org/10.1636/0161-8202(2000)028[0293:NSAROK]2.0.CO;2Pedroche Fernández, A. & Mendoza López, D. (1994) Estudio descriptivo de cavidades kársticas de la Manga de Villaluenga (Cádiz). AEQUA, Grupo Andaluz para el Estudio del Cuaternario, Jerez, 80 pp.
Pérez, T. & Zaragoza, J.A. (2010) Sobre Chthonius (Eppiochthonius) cazorlensis Carabajal Márquez, García Carrillo & Rodríguez Fernández, 2001, stat. nov. (Arácnida: Pseudoscorpiones: Chthoniidae), endemismo de la Cueva Secreta del Sagreo, La Iruela, Jaén, España. Monografías Bioespeleológicas, 5, 17–22.
Reboleira, A.S.P.S., Zaragoza, J.A., Gonçalves, F. & Oromí, P. (2010) Titanobochica, surprising discovery of a new cave-dwelling genus from southern Portugal (Arachnida: Pseudoscorpiones: Bochicidae). Zootaxa, 2681, 1–19.
Reboleira, A.S.P.S., Zaragoza, J.A., Gonçalves, F. & Oromí, P. (2012) Lusoblothrus, a new syarinid pseudoscorpion genus (Arachnida) from Portugal, occupying an isolated position within the Holarctic fauna. Zootaxa, 3544, 52–62.
Reboleira, A.S.P.S., Zaragoza, J.A., Gonçalves, F. & Oromí, P. (2013) On hypogean Roncocreagris (Arachnida: Pseudoscorpiones: Neobisiidae) from Portugal, with descriptions of three new species. Zootaxa, 3670 (2), 283–299.
https://doi.org/10.11646/zootaxa.3670.2.11
Schuster, R.O. (1962) New species of Kewochthonius Chamberlin from California (Arachnida: Chelonethida). Proceedings of the Biological Society of Washington, 75, 223–226.
Šťáhlavský, F. & Král, J. (2004) Karyotype analysis and achiasmatic meiosis in pseudoscorpions of the family Chthoniidae (Arachnida: Pseudoscorpiones). Hereditas, 140, 49–60.
https://doi.org/10.1111/j.1601-5223.2004.01783.xVachon, M. (1940) Éléments de la faune portugaise des Pseudoscorpions (Arachnides) avec description de quatre espèces nouvelles. Anais da Faculdade de Ciencias do Porto Academia Polytechnica do Porto, 25, 7–30.
Vachon, M. (1941a) Chthonius tetrachelatus P. (Pseudoscorpions) et ses formes immatures (1re note). Bulletin du Muséum National d'Histoire Naturelle, Paris, Series 2, 13, 442–449.
Vachon, M. (1941b) Chthonius tetrachelatus P. (Pseudoscorpions) et ses formes immatures (2e note). Bulletin du Muséum National d'Histoire Naturelle, Paris, Series 2, 13, 540–547.
Vachon, M. & Heurtault-Rossi, J. (1964) Une nouvelle espèce française de Pseudoscorpion cavernicole: Spelyngochthonius provincialis (Chthoniidae) du départment de l'Hérault. Bulletin du Muséum National d'Histoire Naturelle, Paris, Series 2, 36, 80–85.
Weygoldt, P. (1968) Vergleichend-embryologische Untersuchungen an Pseudoscorpionen. IV. Die Entwicklung von Chthonius tetrachelatus Preyssl., Chthonius ischnocheles (Hermann) (Chthoniinea, Chthoniidae) und Verrucaditha spinosa Banks (Chthoniinea, Tridenchthoniidae). Zeitschrift für Morphologie und Ökologie der Tiere, 63, 111–154.
https://doi.org/10.1007/BF00401032Zaragoza, J.A. (1984) Un nuevo Chthonius cavernícola de la Provincia de Alicante (Arachnida: Pseudoscorpionidea, Chthoniidae). Mediterranea, Serie de Estudios Biologicos, 7, 49–54.
https://doi.org/10.14198/MDTRRA1984.7.03Zaragoza, J.A. (1985a) Nuevos o interesantes Chthoniidae cavernícolas del País Valenciano (Arachnida, Pseudoscorpiones). Miscelánea Zoológica, 9, 145–158.
Zaragoza, J.A. (1985b) Chthonius (Ephippiochthonius) verai, nueva especie nueva especie cavernícola del sureste español (Arachnida, Pseudoscorpiones, Chthoniidae). Mediterránea, Serie de Estudios Biologicos, 8, 5–15.
Zaragoza, J.A. (1990) Escorpiones, pseudoescorpiones y solífugos de la provincia de Alicante. In: Ayudas a la investigación 1986–1987. Vol. 1. Instituto de Cultura Juan Gil-Albert, Diputación Provincial de Alicante, Alicante, pp. 45–70.
Zaragoza, J.A. (2001) Minniza iberica n. sp., primera cita del género para la Peninsula Ibérica y Redescripción de Minniza algerica Beier, 1931 (Arachnida, Pseudoscorpionida, Olpiidae). Revista Ibérica de Aracnología, 3, 69–78.
Zaragoza, J.A. (2005) Two new cave-dwelling Larca species from the South-East of Spain (Arachnida, Pseudoscorpiones, Larcidae). Revue suisse de Zoologie, 112 (1), 195–213.
https://doi.org/10.5962/bhl.part.80295
Zaragoza, J.A. (2007) Catálogo de los pseudoescorpiones de la Península Ibérica e Islas Baleares (Arachnida: Pseudoscorpiones). Revista Ibérica Aracnología, 13, 3–91.
Zaragoza, J.A. (2010) Arcanobisium, a remarkable new genus, representing a new subfamily with a relictual distribution from eastern Spain (Arachnida: Pseudoscorpiones: Syarinidae). Zootaxa, 2491, 41–60.
Zaragoza, J.A. (2012) Chthonius (Ephippiochthonius) cardosoi, a new hypogean species from Portugal (Pseudoescorpiones: Chthoniidae). Revista Ibérica de Aracnología, 20, 25–30.
Zaragoza, J.A. (2013a) Los pseudoscorpiones (Arachnida) subterráneos de Jaén. In: Grupo de Espeleología de Villacarrillo (G.E.V.) (Ed.), Los invertebrados de hábitats subterráneos de Jaén, 2013, pp. 65–71. [Pérez Fernández, T. y Pérez Ruiz, A. (Coord.)]
Zaragoza, J.A. (2013b) Ord. Pseudoscorpiones de Geer, 1778. Fauna Ibérica, Museo Nacional de Ciencias Naturales, Madrid. Available from: http://www.faunaiberica.es/faunaib/arthropoda/arach/pseudoscorpiones_04.php (accessed 15 June 2016)
Zaragoza, J.A. & Harvey, M.S. (2005) The first record of the genus Mundochthonius Chamberlin (Pseudoscorpiones: Chthoniidae) from Spain: Mundochthonius gallaecicus sp. nov. Revista Ibérica de Aracnología, 12, 17–23.
Zaragoza, J.A. & Henderickx, H. (2009) Roncus elbulli (Arachnida, Pseudoscorpiones), a new species from Cap de Creus Nature Park (Catalonia, Spain), with a key to the Spanish species of the genus Roncus. Zookeys, 8, 19–34.
https://doi.org/10.3897/zookeys.8.110
Zaragoza, J.A. & Pérez, T. (2013) Hypogean pseudoscorpions (Arachnida) from Jaén province (Andalusia, Spain), with descriptions of four new species and a new synonymy. Zootaxa, 3700 (2), 201–225.
https://doi.org/10.11646/zootaxa.3700.2.1Zaragoza, J.A. & Šťáhlavský, F. (2008) New Roncus species (Pseudoscorpiones: Neobisiidae) from Montseny Natural Park (Catalonia, Spain), with remarks on karyology. Zootaxa, 1693, 27–40.
Zaragoza, J.A. & Vadell, M. (2013) Chthonius (Chthonius) campaneti, a new pseudoscorpion species from Coves del Campanet, Mallorca (Spain). Revista Ibérica de Aracnología, 23, 33–41.