What will be your task? • Suggest and justify a phylogeny tree for particular arachnid groups based on the information mentioned in the lecture. • Correct use of terms and appropriate reasoning are important Terminology of cladistics • Taxon – group of organisms • monophyletic • paraphyletic (reptiles without birds) or polyphyletic (birds and bats) • Sister group – closest to the studied group • Outgroup – outside the group we study, for polarisation • Polarisation of the character – determination of the direction of evolution – searching for plesiomorphy • Phylogeny tree – presentation of the hyothesis about relationships • Nnode – represents speciation event • Plesiomorphic character - primitive • Apomorphic character – derived • Synpleziomorphy/apomorphy – common • Autapomorphy – exclusive • Homologic character – present in the ancestor of both taxa • Homoplasy • Konvergency – independent origin of apomorphy in branches without common ancestor (birds–insects) • Paralelism – common ancestors (fish–whales) Terminology of cladistics period: karbon – recent diversity: 1150 species, 9-10 families cephalon: eye tubercle, proboscis, hook-like chelicerae, palps, ovigers and 1 pair of walking legs 3 segments: 3 pairs of walking legs, „abdomen“ legs: 4-6 pairs, contains digestive tract and gonads Excretory and respiratory tract missing reproduction: gonopores on bases of legs class Pycnogonida (Pantopoda) Merostomata: Xiphosura period: Ordovic – recent diversity: 4 species, 3 genera, 1 family distribution: N America (Limulus polyphemus), SE Asia and Philippines (Tachypleus, Carcinoscorpius) body: up to 60 cm cuticle: special hyalinne layer prosoma: chelicerae+5 pairs of walking legs, last one for pushing forward, carapace with s genal thorns, heart lobe opisthosoma: 10 segments including 3segmented metasoma, dorsally fused – thoracetron with lateral thorns and telson, 6. pairs of plate appendages, 1. covering, rest with gills and a pair of chilaria – bifurcated appendages eyes: median and lateral (oculi) chelicerae: 3segmented legs: with gnathocoxae respiration: 5 pairs of lamellar gills excretion: 4 pairs of metanefridia food: solid – vorms, molluscs taxonomy: Synziphosurida (Silur-Devon), Xyphosurida (Karbon-recent) Merostomata: Chasmataspida period: Kambrium-Devon diversity: 6 genera prosoma: genal thorns on carapace opisthosoma: 13 segments, 3segmented mesosoma and 9segmented metasoma, telson, pairs of opercula period: Ordovic-Perm diversity: cca 300 species, over 60 genera, 22 families body: 10-200 cm – the largest arthropods ever prosoma: chelicerae + 5 pairs with gnathocoxae, 5. pair for swimming opisthosoma: telson, 5 pairs of appendages – opercula of gills, median appendix, flat metasoma, gill furrow on 1.-5. sternite eyes: median ocelli and a pair of lateral compound eyes chelicerae: 3 segments ecology: marine, fresh water, amphibious, predators Merostomata: Eurypterida Pterigotina Eurypteracea Transition to the terrestrian ecosystems: evolution of book lungs Ancestral stage: book lungs on 1-5 segment, in scorpions on 2-5 segnemt, in tetrapulmonates on 1-2 segment (Harvey 2002) Arachnida: 12-16 orders Gain aerial respiration anteriorly or anteroventrally directed mouth slit sensilla fluid feeding Lost carapacal pleural doublure cardiac lobe pedal gnathobases moveable endites scorpions: originally 19 segments rest: originally 18 segments prosoma: 6 segments opisthosoma: 12 segments Prosoma (solifugae, palpigrades, schizomida, harvestmen) Propeltidium = proterosoma First two pairs of legs mesopeltidium metapeltidium Opisthosoma (scorpions) mesosoma – tergite, sternite Metasoma - ringhisterosoma Appendages - chelicerae - pedipalps - 4 leg pairs chelate subchelate Respiratory system book lungs trachea Circulatory system Tubular heart in pericardium with pairs of ostia Digestive tract • esophagus – relatively straight and simple – lined with cuticle • sucking stomach – robust – lined with cuticle – Controlled with many muscles – Effective sucking pump Excretion entodermal Malpighian tubules – non-selective diffusion of salt solutions, reasorbtion of nutrients and water in the gut coxal glands = metanefridia depository excretion - guanin amblypygids uropygids palpigrades solifuges Nervous system: caudocranial fusion solifuge: 17 neuromeres as well as in palpigrades and ricinulei harvestman: 16 neuromeres scorpion: 19 neuromeres palpigrades: 17 neuromeressubesophageal g. – pedipalps, legs tick: 10 neuromeres tetrapulmonates and pseudoscorpions: 18 neuromeres front of opisthosoma vision: ocelli main (everse) – anterior median eyes, black, rod cells facing to the object, retina controlled by muscles, in Salticidae and Thomisidae large, absent in Dysderidae secondary (inverse) – rod cells immersed in retina, tapetum lucidum – layer of dark rexlexive pigment, no muscle control Cuticular sensory organs trichobothria - hearing slit sensilla – lyriform organs setae (=sensilla) - olfaction Reproduction gonochorists – gonopore on 8. body segment parthenogenesis: several spider species, Amblypygi, Palpigradi, Schizomida, scorpions, pseudoscorpions, mites spermatophores second. cop. organ penis ontogeny: ecdysis, direct Pseudoscorpiones period: Devonian – recent diversity: 3380 species, 439 genera, 25 families distribution: whole world cuticle: without hyalinne layer prosoma: carapace, rostrosoma – projections on palpal coxae in front of mouth, sternum covered by coxae opisthosoma: 12 segments, last one – circumanal ring pedicel: absent eyes: only lateral: 0-2 pairs legs: 6-7 segments, elongated patellae, 1-2 tarsal segments, 2 claws respiration: 2 pairs of trachea on 9.-10. segment (3.-4. abdominal) food: fluid reproduction: spermatophore, cocoon, protracted ovovivipary: embrya around gonopore chelicerae: 2 segments, chelate, spinning gland at the end of flexible finger – silken cells for copulation, ecdysis, overwintering pedipalps: 6 segments, with chela, venom gland, gnathocoxa Scorpiones period: Silur-recent diversity: cca 1900 species, 184 genera, 16 families body: 1–20 cm cuticle: special hyalinne layer prosoma: gnathosoma, small sternum between 3. and 4. leg pair, a pair of large epistomal arms running backwards toward endosternite mesosoma: 8 segments, 2: gonopore, 3: pektines, 4-7: book lung openings metasoma: 5 segments+telson (no true segment), paired venom gland pedicel: absent eyes: 1 pair of median, 0-5 pairs of lateral chelicerae: 3 segments, chelate pedipalps: 6 segnents, flexible finger legs: 7 segments, gnathocoxae on each pair, tarsi: 2 claws respiration: 4 pairs of book lungs food: fluid, hepatopancreas, stomotheca – space in front of mouth formed by coxapophyses of palps and 1. leg pair Toxicity - neurotoxin family Buthidae Tityus sp. – South America Centruroides exilicauda – Central Amerika Androctonus australis – North Afrika Leiurus quinquestriatus - Mediterranean Centruroides exilicauda Leiurus quinquestriatus Androctonus australis Pandinus imperator embrya develop in oviducts vivipary: eggs poor of yolk, nutrients absorbed through the wall of mother,s gut, juveniles are layd ovovivipary: eggs rich in yolk, only partly fed by mother, hatch shortly after laying Opening armSpecies specific spermatophore Spermia Ovarian diverticulum Reproduction Scorpiones – world distribution family Buthidae: Buthus occitanus family Scorpionidae: Pandinus imperatorfamily Chactidae: Euscorpius tergestinus Europe: fam. Buthidae 9 species fam. Iuridae 3 species fam. Scorpionidae 3 species fam. Superstitionidae 1 species fam. Chactidae 13 species period: Silur-Perm, among the oldest terrestrial animals diversity: 50 species, 7 families body: 2-50 mm opisthosoma: tergites laterally divided to median and 1-2 pairs of lateral, 1. sternite absent, 1. tergite with a crest covering waist, 2-segmented pygidium waist: constricted eyes: median, lateral compound, later disappear chelicerae: 2segmented, subchelate, directing downwards, backwards-directing fang, „clasp-knife“ legs: of the same length, with short basitarsi respiration: 2 pairs of book lungs – 2. and 3. opisthosomal segment ecology: predators Trigonotarbida Thelyphonida (Uropygi) Period: Carbon - recent diversity: 108 species, 18 genera, 1 family body: up to 75 mm cuticle: hyalinne layer absent prosoma: carapace, 3 sternites opisthosoma: 12 segments, 10-12 seg.: pygidium, flagellum (telson) – direction of repugnatory gland secrete, paired openings of repugnatory glands on sides of the anus pedicel: present eyes: 1 pair of median, 3-5 pairs of lateral chelicerae: 2 segments, subchelate pedipalps: 6 segments, working horisontally, capturing prey, cx fused legs: 7 segments, 1. pair elongated, elongated patella, ta secondarily articulated, tarsi of the 2.-4. pair 3segmented respiration: 2 pairs of book lungs food: fluid, sucking oesophagus Mastigoproctus giganteus Reproduction • spermatophore is inserted by male pedipalps • retreats in soil • eggs attached to the gonopore • females carry prenymphs on their back Uropygi – world distribution Schizomida Period: Paleogene (Oligocene) - recent diversity: 258 species, 46 genera, 2 families distribution: humid tropics of the whole world, three introduced species in Europe body: up to 18 mm cuticle: without hyalinne layer prosoma: propeltidium, mesopeltidium, metapeltidium, pointing karapax, 2 sternites between 2. and 4. coxae opisthosoma: 12 segments, repugnatoric glands, mesosoma, metasoma – 3-4 segmented pygidium+ max. 4 segmented flagellum (telson) pedicel: present eyes: absent chelicerae: 2 segments, almost chelate pedipalps: 6 segments, movable in vertical direction, capturing prey, cx fused legs: 7 segments, 1. pair with elongated patella, tactile organ, ta 3 segments respiration: 1 pair of book lungs food: fluid Schizomus sp. Reproduction spermatophore eggs attached close to gonopore female guards eggs in retreat in soil Amblypygi period: Devonian - recent diversity: 158 species, 17 genera, 5 families body: flat, 10-45 mm cuticle: without hyalinne layer prosoma: carapace, wide sternum from 3 sternites opisthosoma: 12 segments, 3 small terminal segments, last one – pygidium, protruding vesicles pedicel: present eyes: 2 median, 6 lateral chelicerae: 2 segments, subchelate pedipalps: 6 segments, elongated, capturing prey, with thorns legs: 1. pair elongated, ta and ti subsegmented, 2-4 pair: 3segmented tarsi, 2 claws, pedal pulvilli respiration: 2 pairs of book lungs secretion: coxal glands on 3. leg pair food: fluid, sucking esophagus and proventriculus Reproduction • spermatophore • eggs attached to the gonopore • female carries praenymphs on her back Phrynus marginemaculatus Amblypygi – world distribution Mediterranean Charinus ionnaticus Sarax mediterraneus Musicodamon atlanteus Haptopoda period: Carboniferous diversity: 1 species body: flat, 12 mm prosoma: karapace pointed, with keels, sternum reduced - 2 sclerites opisthosoma: large genital plate waist: wide eyes: 2 median, 2 lateral eye tubercles chelicerae: 2 segments, clasp-knife pedipalps: pediform legs: 1. pair elongated, with 6 tarsal segments, 2.-4. pair with 4 tarsal segments Uraraneida period: Devon-Perm diversity: 2 species Attercopus, Permarachne Chelicerae: fang without setae, opening of the venom gland? Palps: femur with a field of spines (Fig. B) legs: comb-like tarsal claws, no fissure on the distal edge of patellae Opisthosoma: spigots along sternite edges (Fig. C) – production of silk, no spinnerets; postanal, at least 12 segmented flagellum relatives: sister group of spiders Idmonarachne brasieri – Carbon, no telson, no spinnerets Idmonarachne brasieri Period: Cretaceous Chelicerae: plagiognath, presence of venom gland unclear Palps: tarsus bifurcated – copulatory organ Legs: tarsus with three claws Eyes: on median and lateral tubercles Opisthosoma: pygidium with flagellem, multisegmented ALS and PLS shifted backwards, in position of AMS only spigots, PMS absent Chimerarachne yingi Araneae period: from Carbon (Palaeothele) diversity: over 50 thousand species, 110 families distribution: whole world body: 1-80 mm cuticle: without hyalinne layer prosoma: wide sternum, labium, apodema opisthosoma: originally 2 pairs of spinnerets on 4.-5. segment, pedicel=petiolus eyes: originally a pair of median, and a pair of lateral triads chelicerae: 2 segments, subchelate, venom gland pedipalps: 6 segments, tactile, in males secondary copulatory organs, gnathocoxae legs: 2-3 claws on tarsi respiration: 0-2 pairs of book lungs, 0-2 pairs of tracheae food: fluid, sucking aesophagus and proventriculus – stomodeal part of tract without prominent gnathocoxa tergites two pairs of book lungs eyes on median tubercle plagiognathchelicerae Distribution of Mesothelae Distribution of Mygalomorphae Ricinulei Period: Carbon - recent diversity: 58 species, 3 genera, 1 family body: 3-10 mm cuticle: without hyalinne layer prosoma: carapace, cucullus – grasping prey, sternum covered with coxae opisthosoma: 9 segments, 3segmented pygidium (anal conus), tergites laterally divided pedicel: present eyes: absent, only light sensitive spots legs: 7 segments, 2. pair elongated – tactile organ, no trichobotria, 2-4 tr and ta subsegmented, ta with 2 claws respiration: 1 pair of tracheae food: fluid chelicerae: 2 segments, chelate pedipalps: 6 segments, chelate, fused coxae form camarostom Reproduction Males transport spermatophore with secondary copulatory organs on ta and mt of the 3. leg pair • Female carries eggs under cucullus, juveniles on her back • larva with 3 leg pairs • three nymphal stages Ricinulei – world distribution tropic forests of Africa and America Phalangiotarbida period: Devonian-Permian diversity: 26 species, 3 families body: flat, 10-20 mm prosoma: sternum 1-2-2 opisthosoma: 10 segments+anal operculum, first 6 tergites short, hind ones fused?, sternites longitudinally divided into three plates, dorsal anal operculum waist: wide eyes: 3 pairs on median tubercle chelicerae: very small, 2segmented, subchelate?, directing forward, hidden in the space above cx I pedipalps: pediform, very small legs: coxae triangular, leg length increases from 1 to 4, article proportions are similar respiration: at least 2 pairs of opisthosomal spiracles – tracheae? ecology: predators, mimicry of lycophyte leaf? Opiliones period: Devonian-recent diversity: 6 000 species (in Czechia 37), 500 genera, 25 families distribution: whole world body: 2-22 mm cuticle: without hyalinne layer prosoma: karapace, pro-, meso-, metapeltidium, orificia of repugnatory glands above 2. cx, sternum absent, pair of large epistomal branches running backwards toward endosternite opisthosoma: 9-10 articles, only superficial waist: wide eyes: pair of median on ocularium chelicerae: 3 segments, chelate pedipalps: 6 segments, pedal, short, with or without claws, gnathocoxae legs: 6-7 segments, tarsi subsegmented: up to 50-100, 1-2 claws, 1. two pairs with gnathocoxae, autotomy respiration: openings of tracheae on 8. sternite food: solid, stomotheca – space formed by coxapophyses of palps and 1. leg pair Reproduction • sexual dimorphism •penis or ovipositor (with receptacula seminis in the apical part) under operculum genitale, fast copulation without courtship • eggs are laid to fissures penis ovipositor Opiliones Cyphophthalmi Phalangida Laniatores Palpatores Dispnoi: Ischyropsalidae (Nemastomatidae (Trogulidae Dicranolasmatidae)) Eupnoi: Sclerosomatidae Phalangiidae Sharma et Giribet 2014 Cyphophthalmi Repugnatoric glands on conic tubercle Sironidae - Siro carpaticus body: 1-6 mm eyes: 2 or completely blind opisthosoma: 10 segments, operculum genitale absent legs: 7 segments, tarsi are not secondarily articulated, ta 1.-2. with one claw ecology: under stones, in litter reproduction: males with 3 anal and 1 gland on 4. ta Phalangida: Laniatores Claws on legs I-II different from those on legs III-IV (dual or bifurcated) Family Trogulidae flat body with hood eye tubercle absent legs short, specialised on snails Family Ischyropsalidae Large chelicerae, longer than body – specialised on snails Dyspnoi cheliceral fingers with small regular teeth no additional stigma on ti ovipositor unsegmented Family Nemastomatidae Family Dicranolasmatidae Phalangida: Palpatores Family Sclerosomatidae Astrobunus, Gias, Leiobunum, Nelima Family Phalangiidae Elongated gnathocoxa of the 2. leg pair Eupnoi tibia of adults with additional stigmata ovipositor articulated Phalangida: Palpatores Palpigradi Period: Neogene (Pliocene) - recent diversity: 82 species, 6 genera, 2 families body: 3 mm cuticle: hyalinne layer absent prosoma: mesopeltidium small, 4-5 sternites: largest between palps and 1. leg pair, rostrum opisthosoma: 11 segments, metasoma: 3 segments, flagellum 14-15 segments (telson) waist: constricted eyes: absent chelicerae: 3 segments, chelate pedipalps: 6 segments, walking, with claws, gnathocoxae absent legs: 1. pair – tactile organ, paired claws, subsegmentation, gnathocoxae absent respiration: no organs reproduction: in some species parthenogenesis food: fluid Palpigradi – world distribution tropic, subtropic, in soil, in temperate zone in caves, under stones Slovakia: Eukoenenia spelaea Solifugae period: Carboniferous - recent diversity: 1100 species, 141 genera, 12 families body: 8-80 mm; cuticle: without hyalinne layer prosoma: pro-, meso-, metapeltidium, rostrosoma – projection on palpal coxae in front of mouth, sternum covered by coxae opisthosoma: 11 segments pedicel: absent; eyes: 2-6 median eyes chelicerae: 2 segments, chelate pedipalps: 6 segments, addhesive organ legs: 7 segments, elongated patellae, 1. leg pair - tactile organ, on tr and cx of the 4. pair 3-5 malleoli chemoreceptors, tr 2-4 subsegmented respiration: 3-5 pairs of tracheae; food: fluid Othoes saharae Reproduction • During courtship the male bites the female • Male uses chelicerae for transport of spermatophore (or directly from gonopore to gonopore) • Male chelicerae with flagellum • postembrya not able to walk, guarded in the nest by mother flagellum Solifugae – world distribution Evropa: 18 druhů (Řecko 10 druhů) arid parts of tropics and subtropics (excluding Australia) Europe: 18 species (Greece 10 species) Acari food: solid or fluid transfer of sperm: spermatophore, 3. leg pair, chelicerae, penis ovipositor larva with 3 pairs of legs diversity: so far known over 40 000 species, one tenth of real diversity? body: 0.1-30 mm cuticle: without hyalinne layer waist: wide eyes: lateral or absent chelicerae: originally chelate, 2-3 segments pedipalps: 6 segments, koxae fused Tagmata: gnathosoma, idiosoma legs: 6 (2-7) segments respiration: tracheae or no organs Parasitiformes (Anactinotrichida) Acariformes (Actinotrichida) Opilioacarida (=Notostigmata) Holothyrida (=Tetrastigmata) Ixodida (=Metastigmata) Gamasida (=Mesostigmata) Actinedida (=Prostigmata) Oribatida (=Cryptostigmata) Acaridida (=Astigmata) Cretaceous - recent Devonian - recent leg coxae loose (separate segments) Coxae fused with ventral body wall idiosoma undivided idiosoma transversally divided to propodosoma and histerosoma (between 2. and 3. leg pair) female gonopore transversal fissure female gonopore longitudinal fissure originally 18 segments originally 16 segments stigmas on the hind part of body stigmata are not on the hind part of body Setae without optically active chitin Setae with optically active chitin – birefringend – reflex polarised light Opilioacariformes (=Opilioacarida, Notostigmata) diverzity: 24 species, 9 genera, 1 family body: 1-2.5 mm, apparent border between pro- and opisthosoma opisthosoma: grooves – borders of segments, numerous lyrifissures on tergites – slit sensillae eyes: 2-3 (6) pairs chelicerae: 3 segments respiration: tergites with 4 pairs of stigmata reproduction: gonopore is not covered Holothyrida (=Tetrastigmata) diverzity: 3 families, 5 genera, 30 species body: 2-7 mm, sclerotised eyes: often absent respiration: 1 (2) pair of stigmata laterally from cx 1.-3. reproduction: gonopore covered with plates Ixodida (=Metastigmata) diverzity: 3 families, 22 genera, 860 species circulatory system: present chelicerae: hypostom pedipalps: 3 segments legs: ta of the 1. pair with Haller's organ – chemoreceptor (CO2), temperature and humidity cx separated respiration: stigmata on special plates in front or behind 4. leg pair diversity: 70 families, 10 thousands species circulatory system: present prosoma: tritosternum – branched appendage between cx I chelicerae: retractible respiration: 1 pair of stigmata laterally from cx II-IV reproduction: pedipalps and thick 2. leg pair – holding female during copulation Gamasida (=Mesostigmata) Phytoseiulus persimilis – predator of Tetranychus telarius Dermanyssus gallinae – čmelík kuří Varroa jacobsoni – kleštík včelí Uropoda – foresy of deutonymphae, the fiber is produced in anal area diversity: 120 families, 7 thousand species circulatory system: present respiration: stigmata close to bases of chelicerae or pedipalps Actinedida (=Prostigmata, Trombidiformes) Trombidium holosericeum – sametka rudá Neotrombicula autumnalis – sametka zarděnková Demodex foliculorum – trudník lidský Eriophyes vitis – vlnovník révový Acarapis woodi – roztočík včelíTetranychus telarius – sviluška chmelová Atax crassipes – vodule Oribatida (=Cryptostigmata) diversity: 3 families, 5 genera, 30 species opisthosoma: paired anal plate legs: coxae fused with ventral plate respiration: secondarily evolded traeae, stigmata are not apparent reproduction: paired genital plate Mesoplophora diversity: over 200 families tagmatisation: idiosoma divided by fissure between 2. and 3. leg pair to propodosoma and histerosoma respiration: tracheae absent reproduction: penis – true copulation Ingroup of oribatida Sarcoptiformes (=Acaridida, Astigmata) Sarcoptes scabiei – zákožka svrabová Acarus siro – sladkokaz moučný Dermatophagoides pteronyssinus – prachovka prachová – alergies