Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2017-04-26
Page range: 1–33
Abstract views: 89
PDF downloaded: 51

A new subfamily of Feaellidae (Arachnida, Chelonethi, Feaelloidea) from Southeast Asia

Institut de Systématique, Évolution, Biodiversité (ISYEB), UMR 7205 CNRS, MNHN, UPMC, EPHE, Muséum national d’Histoire naturelle, Sorbonne Universités, 57 rue Cuvier, CP 53, 75005 Paris, France.
Arachnida Pseudoscorpiones systematics morphology biogeography karst amber palaeontology Cretaceous Germany Cambodia Myanmar Vietnam

Abstract

The first extant representatives of the pseudoscorpion family Feaellidae from Southeast Asia are described. Cybella n. gen. is proposed for Cybella deharvengi n. sp. (type species), collected from an isolated limestone hill in Hon Chong Province, Vietnam, and C. bedosae n. sp., found in a limestone cave in Kampuchea, Cambodia. Cybella species seem to be restricted to karst formations and are probably troglophilic. The type localities of the two known species are threatened by quarrying activities, these being particularly pressing in the case of C. deharvengi n. sp. Cybella shows important differences from other Feaellidae that require a modification of the familial diagnosis and justify the erection of a new subfamily, Cybellinae. The discovery of this group provides insights into the evolution of the unusual morphology of the family, notably concerning the pleural plates of Feaellinae, which are lacking in Cybellinae. The smaller sclerites of the pleura of Pseudogarypidae and Feaellidae are shown to be muscle apodemes, which provide an additional synapomorphy for Feaelloidea. Two types of coxal spines, termed primary and secondary, are distinguished in Feaelloidea, based on the presence of a lumen within the primary spines and its absence in secondary spines. The new morphological term atrial plate is proposed for a sclerotized plate of the male genitalia, extending between the lateral rods and the lateral apodemes. Claims that the internal genital setae of males of non-chthonioid pseudoscorpions are secretory are reviewed and found to lack support.

        Additional information concerning the fossil genus Protofeaella Henderickx, 2016 is provided, based on an adult male in amber from the Cretaceous (lowermost Cenomanian) of Myanmar. Protofeaella shares with Cybella the absence of pleural plates and the antiaxial position of the chemosensory setae of the movable chelal finger. However, it differs from both Cybellinae and Feaellinae in having relatively long chelal fingers that lack a tuberculate basal tooth, both of which are interpreted as symplesiomorphic states within Feaellidae. Protofeaella is therefore provisionally treated as a stem-group feaellid and not assigned to a subfamily.

        The existence of a Cretaceous member of the Pseudogarypidae is noted in the mid-Cretaceous (late Albian‒early Cenomanian) of Germany, representing the oldest record of this family.

 

References

  1. Andrade, R. de (2004) Primeiro registro de Feaellidae (Pseudoscorpiones) para a América do Sul. In: Machado, G. & Brescovit, A.D. (Eds.), Programma & Resumos. IV. Encontro de Aracnólogos do Cone Sul. São Pedro, São Paulo, Brazil, 7–12 December 2003, pp. 129.

    Anonymous (2016) Vietnam: government to sell Vicem stake. Global Cement Magazine, September 2016, 52. Available from: http://www.globalcement.com/pdf/eGCSept2016ns.pdf (accessed 1 September 2016)

    Arabi, J., Judson, M.L.I., Deharveng, L., Lourenço, W.R., Cruaud, C. & Hassanin, A. (2012) Nucleotide composition of CO1 sequences in Chelicerata (Arthropoda): detecting new mitogenomic rearrangements. Journal of Molecular Evolution, 74, 81–95.

    https://doi.org/10.1007/s00239-012-9490-7

    Beier, M. (1955) Pseudoscorpionidea, gesammelt während der schwedischen Expeditionen nach Östafrika 1937–1938 und 1948. Arkiv för Zoologi, Series 2, 7, 527–558.

    Benedict, E.M. (1978) False scorpions of the genus Apocheiridium Chamberlin from western North America (Pseudoscorpionida, Cheiridiidae). Journal of Arachnology, 5, 231‒241.

    Benedict, E.M. & Malcolm, D.R. (1978) The family Pseudogarypidae (Pseudoscorpionida) in North America with comments on the genus Neopseudogarypus Morris from Tasmania. Journal of Arachnology, 6, 81‒104.

    Bird, T.L., Wharton, R.A. & Prendini, L. (2015) Cheliceral morphology in Solifugae (Arachnida): primary homology, terminology, and character survey. Bulletin of the American Museum of Natural History, 394, 1‒355.

    https://doi.org/10.1206/916.1

    Chamberlin, J.C. (1923) The genus Pseudogarypus Ellingsen (Pseudoscorpionida-Feaellidae). Entomological News, 34, 146–149, 161–166, pl. 5.

    Chamberlin, J.C. (1931) The arachnid order Chelonethida. Stanford University Publications, University Series: Biological Sciences, 7, 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.

    https://doi.org/10.1093/aesa/36.3.486

    Chamberlin, J.C. (1952) New and little-known false scorpions (Arachnida, Chelonethida) from Monterey County, California. Bulletin of the American Museum of Natural History, 99, 259–312.

    Dashdamirov, S. (2012) A new genus and species of false scorpion from northern Vietnam (Arachnida, Chelonethi, Neobisiidae). Tropical natural History, 12, 97–115.

    Deharveng, L. & Bedos, A. (2012) Diversity patterns in the tropics. In: White, W.B. & Culver D.C. (Eds), Encyclopedia of Caves. 2nd Edition. Academic Press, Waltham, Ma, pp. 238‒250.

    https://doi.org/10.1016/b978-0-12-383832-2.00032-3

    Deharveng, L. & Bedos, A. (2016) Microblattellus lecongmani. In: International Union for the Conservation of Nature, The IUCN Red List of Threatened Species, Version 2016-3. e.T89690813A89690824.

    https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T89690813A89690824.en

    Deharveng, L., Bedos, A., Le, C.K., Le, C.M. & Truong, Q.T. (2009) Endemic arthropods of the Hon Chong hills (Kien Giang), an unrivaled biodiversity heritage in Southeast Asia. In: Le, C.K., Truong, Q.T. & Ly, N.S. (Eds.), Beleaguered Hills: managing the biodiversity of the remaining karst hills of Kiên Giang, Vietnam. Nong Nghiep Publishing, Ho Chi Minh City, pp. 31–57.

    Deharveng, L., Le, C.K., Le, C.M. & Bedos, A. (2005) Hot issues in karst conservation: the biodiversity of Hon Chong Hills (southern Vietnam), with emphasis on invertebrate endemism. In: Batelaan, O., Dusar, M., Masschelein, J., Vu, T.T., Tran, T.V. & Nguyen, X.K. (Eds.), Trans-KARST-2004. Proceedings of the International Transdisciplinary Conference on Development and Conservation of Karst Regions. Hanoi, Vietnam, 13–18 September 2004, pp. 40. [abstract]

    Ellingsen, E. (1906) Report on the Pseudoscorpions of the Guinea Coast (Africa) collected by Leonardo Fea. Annali del Museo civico di Storia naturale di Genova, Series 3, 2, 243–265, pl. IV.

    Gao, Z.-Z. & Zhang, F. (2012) A new species of the genus Dactylochelifer Beier from Shanxi, China (Pseudoscorpiones: Cheliferidae). ISRN Zoology, Article 583830, 1‒4.
    https://doi.org/10.5402/2012/583830

    Gardini, G. (1991) Tre nuove specie di Roncus L. Koch, 1873 della Liguria occidentale (Pseudoscorpionida Neobisiidae). Memorie della Societ entomologica Italiana, 70, 313–334.

    Gardini, G. (1993) Roncus caprai n. sp. della Liguria orientale e note su R. gestroi Beier, 1930 (Pseudoscorpionida, Neobisiidae). Memorie della Societ entomologica Italiana, 71, 409–425.

    Grandjean, F. (1933) Observations sur les Acariens (5e série). Bulletin du Muséum national d’Histoire naturelle, Series 2, 5, 461‒468.

    Hammen, L. van der (1986) Comparative studies in Chelicerata IV. Apatellata, Arachnida, Scorpionida, Xiphosura. Zoologische Verhandelingen, 226, 1‒52.

    Harvey, M.S. (1985) The systematics of the family Sternophoridae (Pseudoscorpionida). Journal of Arachnology, 13, 141–209.

    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.
    https://doi.org/10.1071/ZO9860753

    Harvey, M.S. (1987) A revision of the genus Synsphyronus Chamberlin (Garypidae: Pseudoscorpionida: Arachnida). Australian Journal of Zoology, Supplement Series, 126, 1–99.

    Harvey, M.S. (1989) A new species of Feaella Ellingsen from north-western Australia (Pseudoscorpionida: Feaellidae). Bulletin of the British arachnological Society, 8, 41–44.

    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. (1996a) Small arachnids and their value in Gondwanan biogeographic studies. In: Hopper, S.D., Chappill, J.A., Harvey, M.S. & George, A.S. (Eds.), Gondwanan heritage: past and future of the Western Australian biota. Surrey Beatty & Sons, Chipping Norton, pp. 155–162.

    Harvey, M.S. (1996b) The biogeography of Gondwanan pseudoscorpions (Arachnida). Revue suisse de Zoologie, Hors Série (1), 255–264.

    Harvey, M.S. (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.

    Harvey, M.S., Andrade, R. & Pinto-da-Rocha, R. (2016) The first New World species of the pseudoscorpion family Feaellidae (Pseudoscorpiones: Feaelloidea) from the Brazilian Atlantic Forest. Journal of Arachnology, 44, 227‒234.
    https://doi.org/10.1636/JoA-S-16-007

    Harvey, M.S. & Edward K.L. (2007) A review of the pseudoscorpion genus Ideoblothrus (Pseudoscorpiones, Syarinidae) from western and northern Australia. Journal of natural History, 41, 445–472.

    https://doi.org/10.1080/00222930701219123

    Harvey, M.S. & 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.
    https://doi.org/10.18195/issn.0312-3162.24(4).2008.387-394

    Harvey, M.S. & Muchmore W.B. (1990) The systematics of the family Menthidae (Pseudoscorpionida). Invertebrate Taxonomy, 3, 941–964.

    https://doi.org/10.1071/IT9890941

    Harvey, M.S., Ratnaweera, P.B., Udagama, P.V. & 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.

    https://doi.org/10.1080/00222933.2012.707251

    Harvey, M.S. & 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. [2009]

    https://doi.org/10.1636/A09-53.1

    Harvey, M.S. & Volschenk, E.S. (2007) Systematics of the Gondwanan pseudoscorpion family Hyidae (Pseudoscorpiones: Neobisioidea): new data and a revised phylogenetic hypothesis. Invertebrate Systematics, 21, 365–406.

    https://doi.org/10.1071/IS05030

    Henderickx, H. (2009) A new Feaella species (Pseudoscorpiones: Feaellidae) from Kenya. Phegea, 37, 41–47.

    Henderickx, H. & Boone, M. (2014) The first fossil Feaella Ellingsen, 1906, representing an unexpected pseudoscorpion family in Baltic amber (Pseudoscorpiones, Feaellidae). Entomo-info, 25, 5–11.

    Henderickx, H. & Boone, M. (2016) The basal pseudoscorpion family Feaellidae Ellingsen, 1906 walks the earth for 98.000.000 years: an new fossil genus has been found in Cretaceous Burmese amber (Pseudoscorpiones: Feaellidae). Entomo-info, 27, 7–13.

    Hennig, W. (1966) Phylogenetic Systematics. University of Illinois Press, Urbana, xiii + 263 pp.

    Heurtault, J. (1983) Pseudoscorpions de Côte d’Ivoire. Revue arachnologique, 5, 1–27.

    Heurtault-Rossi, J. & Jézéquel, J.-F. (1965) Observations sur Feaella mirabilis Ell. (Arachnide, Pseudoscorpion). Les chélicères et les pattes-mâchoires des nymphes et des adultes. Description de l’appareil reproducteur. Bulletin du Muséum national d’Histoire naturelle, Series 2, 37, 450–461.

    International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, XXIV + 306 pp.

    IUCN (2012) Planning for biodiversity action in Vietnam. International Union for the Conservation of Nature, Gland, Switzerland. Available from: https://www.iucn.org/fr/node/12528 (accessed 1 September 2016)

    Judson, M.L.I. (1990) 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. (1998) A sternophorid pseudoscorpion (Chelonethi) in Dominican amber, with remarks on the family. Journal of Arachnology, 26, 419–428.

    Judson, M.L.I. (2007a) 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. (2007b) First fossil record of the pseudoscorpion family Pseudochiridiidae (Arachnida, Chelonethi, Cheiridioidea) from Dominican amber. Zootaxa, 1393 (1), 45–51.
    https://doi.org/10.11646/zootaxa.1393.1.5

    Judson, M.L.I. (2012) Reinterpretation of Dracochela deprehendor (Arachnida: Pseudoscorpiones) as a stem-group pseudoscorpion. Palaeontology, 55, 261‒283.

    https://doi.org/10.1111/j.1475-4983.2012.01134.x

    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, 57‒79.

    https://doi.org/10.5281/zenodo.50595

    Kew, H.W. (1912) On the pairing of pseudoscorpions. Proceedings of the zoological Society of London, 25, 376‒390.

    Kiernan, K. (2010) Human impacts on geodiversity and associated natural values of bedrock hills in the Mekong Delta. Geoheritage, 2, 101–122.

    https://doi.org/10.1007/s12371-010-0015-8

    LafargeHolcim (2016) LafargeHolcim to divest LafargeHolcim Vietnam. Media Release, 4 August 2016, LafargeHolcim, Zurich. Available from: http://www.lafargeholcim.com/lafargeholcim-to-divest-lafargeholcim-vietnam (accessed 1 September 2016)

    Legg, G. (1973) Spermatophore formation in the pseudoscorpion Chthonius ischnocheles (Chthoniidae). Journal of Zoology, London, 170, 367–394.

    https://doi.org/10.1111/j.1469-7998.1973.tb01384.x

    Legg, G. (1974) A generalised account of the male genitalia and associated glands of pseudoscorpions. Bulletin of the British arachnological Society, 3 (3), 66‒74.

    Legg, G. (1975a) The genitalia and associated glands of five British species belonging to the family Neobisiidae (Pseudoscorpiones: Arachnida). Journal of Zoology, 177, 123–151.

    https://doi.org/10.1111/j.1469-7998.1975.tb05974.x

    Legg, G. (1975b) Spermatophore formation in pseudoscorpions. In: van Helsdingen, P.J. (Ed.), Proceedings of the 6th International Arachnological Congress, Amsterdam, 22–26 April 1974, pp. 141–144.

    Lossau, N. (2002) Pseudoskorpione aus der Kreidezeit in rheinischem Kalksteinbruch entdeckt. Die Welt, 22 Mai 2002, 29.

    Mahnert, V. (1982a) Die Pseudoskorpione (Arachnida) Kenyas II. Feaellidae; Cheiridiidae. Revue Suisse de Zoologie, 89, 115–134.

    https://doi.org/10.5962/bhl.part.82432

    Mahnert, V. (1982b) Sigles trichobothriaux chez les pseudoscorpions (Arachnida). Atti della Societ Toscana di Scienze naturali, Memorie (B), 88 (Supplement), 185–192. [1981]

    Mahnert, V. (2011) 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. (2014) Pseudoscorpions (Arachnida: Pseudoscorpiones) from the Galapagos Islands (Ecuador). Revue suisse de Zoologie, 121, 135‒210.

    Morris, J.C.H. (1948) A new genus of pseudogarypin pseudoscorpions possessing pleural plates. Papers and Proceedings of the Royal Society of Tasmania, 1947, 43–47.

    Muchmore, W.B. (1973a) The genus Chitrella in America (Pseudoscorpionida, Syarinidae). Journal of the New York entomological Society, 81, 183–192.

    Muchmore, W.B. (1973b) The pseudoscorpion genus Mexobisium in Middle America (Arachnida: Pseudoscorpionida). Association for Mexican Cave Studies, 5, 63–72.

    Muchmore, W.B. (1981) Cavernicolous species of Larca, Archeolarca and Pseudogarypus with notes on the genera (Pseudoscorpionida, Garypidae and Pseudogarypidae). Journal of Arachnology, 9, 47–60.

    Muchmore, W.B. (1982a) Some new species of pseudoscorpions from caves in Mexico (Arachnida, Pseudoscorpionida). Texas Memorial Museum, Bulletin, 28, 63–78.

    Muchmore, W.B. (1982b) The genera Ideobisium and Ideoblothrus, with remarks on the family Syarinidae (Pseudoscorpionida). Journal of Arachnology, 10, 193–221.

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

    https://doi.org/10.1016/j.ympev.2008.06.002

    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.

    https://doi.org/10.5431/aramit5001

    Nassirkhani, M., Doustar Sharaf, M.M. & Azimi, N. (2016) Redescription of a well-known pseudoscorpion species, Dinocheirus panzeri (C.L. Koch) (Pseudoscorpiones: Chernetidae). Journal of Zoology Studies, 2 (6), 6‒12.

    Nassirkhani, M. & Hamidi, K. (2015) Withius nanus Mahnert, 1988 (Pseudoscorpiones: Withiidae); a new record of phoresy on Goodwin’s brush-tailed mouse from Iran. Journal of Zoology Studies, 2, 13‒21.

    Nassirkhani, M. & Harvey, M.S. (2013) Description of the postlarval stages of Dactylochelifer gracilis Beier, Pseudoscorpiones: Cheliferidae. International Journal of Zoology, Article 782638, 1‒15.

    https://doi.org/10.1155/2013/782638

    Nassirkhani, M. & Takalloo zade, H.M. (2013) The first report of Dactylochelifer spasskyi Redikorzev (Pseudoscorpiones: Cheliferidae) from Iran. Journal of entomological Research, 4, 335‒340.

    Nassirkhani, M. & Vafai Shoushtari, R. (2014) Redescription and remarks on the species Minniza persica (Pseudoscorpiones: Olpiidae) from Iran. Zoology in the Middle East, 60, 272‒277.

    https://doi.org/10.1080/09397140.2014.939814

    Nassirkhani, M., Vafai Shoushtari, R. & Rahmat Abadi, M. (2016) A new pseudoscorpion species in Amblyolpium (Pseudoscorpiones: Garypinidae) from a house in Kermanshah Province, Iran. Arachnologische Mitteilungen, 52, 1–3.

    https://doi.org/10.5431/aramit5201

    Petrunkevitch, A. (1925a) Arachnida from Panama. Transactions of the Connecticut Academy of Arts and Sciences, 27, 51‒248.

    Petrunkevitch, A. (1925b) Descriptions of new or inadequately known American spiders. Annals of the entomological Society of America, 18, 313‒324.

    https://doi.org/10.1093/aesa/18.3.313

    Proctor, H.C. (1993) Mating biology resolves trichotomy for cheliferoid pseudoscorpions (Pseudoscorpionida, Cheliferoidea). Journal of Arachnology, 21, 156–158.

    Shi, G., Grimaldi, D.A., Harlow, G.E., Wang, J., Wang, J., Yang, M., Lei, W., Li, Q. & Li, X. (2012) Age constraint on Burmese amber based on U–Pb dating of zircons. Cretaceous Research, 37, 155–163.

    https://doi.org/10.1016/j.cretres.2012.03.014

    Vachon, M. (1938a) Recherches anatomiques et biologiques sur la reproduction et le développement des Pseudoscorpions. Annales des Sciences naturelles, Zoologie, Series 11, 1, 1–207.

    Vachon, M. (1938b) Remarques sur la famille des Cheiridiidae, Chamberlin, propos d’un nouveau genre et d’une nouvelle espèce: Paracheiridium decaryi (Arachnida Pseudoscorpionidae). Bulletin de la Sociét entomologique de France, 43, 235–241.

    Viehofen, A., Hartkopf-Fröder, C. & Friis, E.M. (2008) Inflorescences and flowers of Mauldinia angustiloba sp. nov. (Lauraceae) from middle Cretaceous karst infillings in the Rhenish Massif, Germany. International Journal of Plant Sciences, 169, 871–889.

    https://doi.org/10.1086/589975

    Villegas-Guzmán, G.A. & Francke, O.F. (2009) Description of the male of the genus Typhloroncus Muchmore, 1979 (Pseudoscorpiones: Ideoroncidae). In: Cokendolpher, J.C. & Reddell, J.R. (Eds.), Studies on the cave and endogean fauna of North America V. Texas Memorial Museum Speleological Monographs, 7, pp. 63‒66.

    Weygoldt, P. (1969) The biology of pseudoscorpions. Harvard University Press, Cambridge Mass., xiv + 145 pp.

    Weygoldt, P. (1970) Vergleichende Untersuchungen zur Fortpflanzungsbiologie der Pseudoscorpione II. Zeitschrift für Zoologische Systematik und Evolutionsforschung, 8, 241–259.
    https://doi.org/10.1111/j.1439-0469.1970.tb00878.x

    With, C.J. (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. Det Konigelige Danske Videnskabernes Selskabs Skrifter, Series 7, 3, 1–214, 1 map, pls I–IV.