štírci (Arachnida: Pseudoscorpiones) ‫كذرب‬ ЛОЖНОСКОРПИОНЫ Afterskorpione Šťúriky Pseudoscorpions Zaleszczotki Pseudoscorpioni False Scorpions Pseudoschorpioenen Pseudoescorpiones 擬蠍目 MOSSKORPIONER Ebaskorpionilised CHELONETHIDA Bogskorpioner Álskorpiók ‫דמויי‬-‫עקרב‬ Yalancıakrep app. 3380 species from 25 families http://www. http://museum.wa.gov.au/catalogues-beta/pseudoscorpions Shultz 2007 Haplocnemata Börner, 1904 Chelicerae – two-segmented pincers similar articulation in both groups rostrosoma empodium (= pulvillus, arolium) legs III and IV femur shorter than patella ( patello-tibial articulation) no sternum coxae in contact Alberti & Peretti 2002 Legg 1973 Papato et al. 2010 Dunlop et al. 2012 Sharma et al. 2014 Mol. Biol. Evol. Lozano-Fernandez et al. 2019 Balesteros & Sharma 2019 Ontano et al., 2021, Mol. Biol. Evol. doi: 10.1093/molbev/msab038 Flagellum Charakteristics chelicerae – two-segmented, chelate ventrolateral articulation Giribet & Sharma 2015 Charakteristics chelicerae – two-segmented, chelate ventrolateral articulation Charakteristics pedipalps – with venom glands (without Heterosphyronida) Cordylochernes Ideoroncus Neobisium Charakteristics legs Arolium Diplosphyronida (superfamily Neobisioidea a Garypoidea) Monosphyronida (superfamily Cheliferoidea) Heterosphyronida (superfamily Chthonioidea) Charakteristics carapax Charakteristics trachee opening on 3. and 4. opisthosomal segment Charakteristics Vascular system is opening (heart 4-2 pars of opening) extrection: excretory cells in stomach coxal glands Charakteristics nervous system concentrated eyes 0-2 pairs Charakteristics Feeding, digestion Charakteristics Cuticular sense organs Charakteristics Cuticular sense organs Reproduction pseudoscorpions reproduce sexually Reproduction Roncus NeobisiumChthonius Chthonius Neobisium Garypus Chernes Chelifer Chthoniidae Neobisiidae Cheiridiidae Olpiidae Reproduction fertilization is internal but indirect A) Spermatophors without females Chelifer cancroides Dinocheirus tumidus Atemnidae Chernetidae Cheliferidae ram`s-horn organ Reproduction fertilization is internal but indirect A) Spermatophors with females brood sac Reproduction 9-16 15-25 15-25 5 weeks 10 days 3 weeks 3-5 20-40 3 weeks Reproduction Life cycle nymphs (3 instars) protonymph deutonymph tritonymph adult 1 mm Life cycle longevity 2 – 4 years Abbundance up to 900 ex./m2 0% 10% 20% 30% 40% 50% 60% 70% 80% 90% 100% XII I II III IV V VI VII VIII IX X XI P D T samice samci Chthonius ischnocheles Gabbutt 1967Šťáhlavský 2000 Allochernes wideri Garypinus electri (Baltic amber) Phoresy Tridenchthoniidae 1 Lechytiidae 3 Syarinidae 1 Neobisiidae 1 Larcidae 1 Sternophoridae 1 Cheiridiidae 3 Cheliferidae 9 Atemnidae 7 Withiidae 3 Chernetidae 39 dispersal x theory of origin predation Chernes hahnii Opatová & Šťáhlavský 2008 Del-Claro K. & Tizo-Pedroso E. 2009: Ecological and evolutionary pathways of social behavior in Pseudoscorpions (Arachnida: Pseudoscorpiones). Acta Ethologica 12: 13-22. Paratemnoides nidificator Sociality Garypinus electri Baltic amber 35-40 mil. Pseudochthonius squamosus Dominican amber 30 mil. Dracochela deprehendor Mundochthonius styriacus Dunlop 2010 upraveno Harvey 1992 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. upraveno Harvey 1992 Benavides et al. 2019, Mol. Phyl. Evol. Beier M. Chamberlin J.C. Mahnert V. Judson M. Harvey M.S. Heurtault J. Vachon M. Weygoldt P. Acarus cancroides Acarus scorpioides Stecker A. Verner P.H. Ducháč V. Červená et al. 2020 Arthropoda Selecta 29(2): 219–228 čel.: Chthoniidae (631/9) 1122 carapax broader in front part chelicerae large pedipalps without venom glands tergites undivided simple spermatophors ČR 3 genera: Mundochthonius Chthonius Ephippiochthonius E. tetrachelatus Preyssler, J. D. (1790): Verzeichniss Böhmischer Insekten. Prag 1-57. čel.: Neobisiidae (548/8) 2222 carapax of the same breadth chelicerae large tergites undivided ČR 3 genera: Neobisium Roncus Microbisium ■ čel.: Larcidae(15/1) 2222 2 pairs of eyes chelicerae small Tergites divided ČR: Larca lata čel.: Cheliferidae(275/3) 1111 chelicerae small Tergites divided ČR: Chelifer cancroides Mesochelifer ressli Dactylochelifer latreillei čel.: Chernetidae(652/13) 1111 Chelicerae small Tergites divided phoresy Chernetidae – Lamprochernes nodosus (phoresy) Chernes hahnii Dendrochernes cyrneus Anthrenochernes stellae ?? čel.: Cheiridiidae(71/2) 1111 Chelicerae small Tergites divided Body size small ČR: Cheiridium museorum Apocheiridium ferum 1mm 1mm under bark hollow trees Leaf litter, soil caves Kotrbová 2012 Garypinidae Chthoniidae Olpiidae Chernetidae Kotrbová 2012 Definition of species Biological species concept (reproductive or isolation concept) groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups (Mayr 1942) Mate-recognition species a group of sexually reproducing organisms that recognize one another as potential mates Evolutionary species an entity composed of organisms which maintains its identity from other such entities through time and over space, and which has its own independent evolutionary fate and historical tendencies (Simpson 1951) Typological species (morphospecies) a group of organisms in which individuals conform to certain fixed properties (a type) problems – variability, cryptic species, monophyly Parmakelis et al. 2013 Fet & Solegland 2002 Kovařík 1998 Rein 2021 1 genus/5 species 3 genera/90 species Euscorpius Fet & Solegland 2002 Euscorpius Štundlová et al. 2019, Mol. Phyl. Evol. Kovařík 1998 Rein 2021 1 genus/5 species 3 genera/90 species van Haarden et al. 2013 Moulds el al. 2007 ProtocheliferHorus Ohira et al. 2018 Mundochthonius Harms 2018 Pseudotyrannochthonius Muster et al. 2021, Ecol. Evol., doi: 10.1002/ece3.8088 „The German Barcode of Life campaign reveals high levels of undocumented diversity in European false scorpions“ - extinction Gene flow only between neighbouring populations. Extinction of some population cause very fast genetic separation. Cordylochernes scorpioides fast speciation Wilcox et al. 1997 fast speciation - Polyploidization or hybridization – frequently in plants Diplotemnus insolitus 2n=122Chernes hahnii 2n=49 Chernetidae Atemnidae - allopatric population splits into two geographically isolated populations vicariant new populations of similar size Pfeiler E, Bitler B.G., Castrezana S., Matzkin L.M. & Markow T.A. 2009: Genetic diversification and demographic history of the cactophilic pseudoscorpion Dinocheirus arizonensis from the Sonoran Desert. Molecular Phylogenetics and Evolution, 52: 133-141. Dinocheirus arizonensis phoretic dispersal on Odontoloxozus longicornis Plio-Pleistocene gradual speciation gradual speciation - allopatric peripatric separation of only small subpopulation Roncus alpinus Roncus binaghii Roncus tuberculatus population splits into two geographically isolated populations - sympatric formation of two or more descendant species from a single ancestral species all occupying the same geographic location Neobisium dolomiticum - parapatric between alopatric and sympatric speciation – gene flow still occurs Neobisium ruffoi Chernes hahnii Chernes similis under the bark leaf litter gradual speciation X0 neo XY Diplotene Diplotene 8 XX Y Chromosomal speciation “hybrid-sterility models” predicted that the recombination among rearranged chromosomes in heterokaryotypic hybrids generate unbalanced gametes and thus reduce fertility. Majority of closely related species differ in karyotypes (app. 90-98% speciation change karyotypes) Is it consequence or cause of speciation? Neobisium – Europe app. 200 species Neobisium carcinoides 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 X Y 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 X 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 22 23 2421 25 26 27 28 29 30 31 32 34 X Y33 Common European species Neobisium carcinoides is complex of several species. N. carcinoides I, 2n = 70 N. carcinoides II, 2n = 54 N. carcinoides III, 2n = 65 0,12 0,14 0,16 0,18 0,2 0,22 0,24 0,26 0,5 0,6 0,7 0,8 0,9 1 1,1 1,2 délka femuru pedipalpu šířkafemurupedipalpu Chernes hahnii Fig. 1: Results of Bayesian analysis ( GTR+I+Γ+covarion model). Bootstrap values above 0,95% are shown below branches. Values above branches correspond to bootstrap support obtained in Maximum Parsimony analysis (only values above 70% are reported). Lamprochernetinae Chernetinae * Neobisium erythrodactylum Neobisium polonicum Roncus montsenyensis Dactylochelifer latreillei Dendrochernes cyrneus (2n = 57) Chernes similis (2n = 63) Chernes hahnii (2n = 49) Chernes nigrimanus (2n = 59) Chernes vicinus Dinocheirus panzeri (2n = 77) Pselaphochernes anachoreta Pselaphochernes lacertosus (2n = 60) Allochernes wideri (2n = 55) Pselaphochernes scorpioides (2n = 41) Chernes cimicoides (2n = 61) Chernetidae protonymfa deutonymfa tritonymfa dospělec 1 mm collection pit fall traps... František Šťáhlavský Katedra zoologie PřF UK Viničná 7, Praha 2, 128 44 stahlf@natur.cuni.cz References: Beier, M. (1963) Ordnung Pseudoscorpionidea (Afterskorpione). Bestimmungsbücher zur Bodenfauna Europas. Lieferung 1. Akademie–Verlag, Berlin, 313 pp. Chamberlin, J.C. (1931) The arachnid order Chelonethida. Stanford Univ. Publ. Biol. Sci. 7, 1–284. Harvey, M.S. (1992) The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invert. Taxon. 6, 1373–1435. Harvey, M. S. (2009) Pseudoscorpions of the World, Version 1.2, Western Australian Museum, Perth. Available from: http://www.museum.wa.gov.au/arachnids/pseudoscorpions/ Harvey, M.S. (2011) Pseudoscorpions of the World, version 2.0. Western Australian Museum, Perth. http://www.museum.wa.gov.au/catalogues/pseudoscorpions Legg, G. & Jones, R. E. (1988) Pseudoscorpions (Arthropoda; Arachnida). Keys and notes for the identification of the species. In: Kermack, D. M. & Barnes, R. S. K. (Eds), Synopses of the British Fauna (New Series), No. 40. The Linnean Society of London and the Estuarine and Brackish–Water Sciences Association, Leiden–New York–København–Köln, pp. 1–159. 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. www.jorgenlissner.dk/pseudoscorpions.aspx www.museum.wa.gov.au/catalogues/pseudoscorpions/ www.arachnology.cz