Presentation is loading. Please wait.

Presentation is loading. Please wait.

Lousy Phylogenies: Phthiraptera systematics and the antiquity of lice Division of Environmental and Evolutionary Biology Institute of Biomedical and Life.

Similar presentations


Presentation on theme: "Lousy Phylogenies: Phthiraptera systematics and the antiquity of lice Division of Environmental and Evolutionary Biology Institute of Biomedical and Life."— Presentation transcript:

1 Lousy Phylogenies: Phthiraptera systematics and the antiquity of lice Division of Environmental and Evolutionary Biology Institute of Biomedical and Life Sciences Graham Kerr Building University of Glasgow Scotland, UK Vincent S. Smith

2 Lice (Phthiraptera): biology i.Diversity 4,384 spp chewing lice (3,508 bird & 402 mammal spp) 543 spp sucking lice (812 spp mammal) iv.Cophylogeny, cospeciation & coevolution ii.Physical characteristics wingless, dorsoventrally flattened, 0.3-11mm, 3 nymphal instars feed on feathers, fur & skin debris (chewing lice) or blood adaptations against host grooming iii.Distribution mirror host distribution worldwide in virtually every habitat absent on Chiroptera, Cetacea, Microbiotheria, Monotremata, Notoryctemorphia, Pholidota & Sirenia.

3 - 1,350 spp in 6 families - Most primitive suborder - More vagile, less host specific - Widespread on birds - 2 fam. South American mammals & marsupials - 1 fam. on Australasian marsupials - 3,060 spp., # families ? - Widespread, ~90% spp. on birds - Avian groups poorly known - Highly host specific - Niche restricted - Mammal infesting species of veterinary significance - Just 3 species - On unrelated hosts - Acquired via shared use of watering holes? - Tiny mandibles present at end of a long rostrum pierce hosts thick skin - 550 spp. in 16 families - Absent from birds - Well studied - Greatest medical & veterinary significance - Vectors of several important diseases - Low prevalence on healthy hosts AmblyceraIschnoceraRhynchophthirinaAnoplura Lice (Phthiraptera): taxonomy Chewing Lice Mallophaga Sucking Lice (Anoplura)

4 Lice (Phthiraptera): phylogeny PsocopteraIschnoceraRhynchophthirinaAnopluraAmblycera Chewing Lice Mallophaga Sucking Lice (Anoplura) Kim & Ludwig, 1982 (Morphology) ( Parasitic Lice ( Phthiraptera) Lyal, 1985 (Morphology)

5 GenusFamilySuborder 3 532 3,060 1,344 Rhynchophthirina Amblycera Ischnocera Anoplura Mammals Birds Amblycera Ischnocera Rhyn… Anoplura Phthiraptera: Phylogenetic resolution, 1996

6 GenusFamilySuborder 3 532 3,060 1,350 Amblycera Ischnocera Anoplura Phthiraptera: Phylogenetic resolution, 2003 Rhyn… Mammals Birds

7 GenusFamilySuborder 3 532 3,060 1,344 Amblycera Ischnocera Rhyn… Anoplura 1996

8 Amblyceran Families 85 84 Grilicolinae Gyropinae Protogyropinae Trimenoponidae Ricinidae Laemobothriidae Boopiidae Menoponidae Liposcelidae (Psocoptera) Birds Worldwide & Australasian Marsupials South/Central American Rodents & Marsupials Gyropidae Analysed by Marshall, 2003 from Clay, 1970 (Morphology)

9 Laemobothriidae - Falconiforms Menacanthus/Menopon-cpx. - Birds worldwide Colpocephalum-complex - Birds worldwide Austromenopon-complex - Aquatic birds Dennyus-complex - Birds worldwide Boopidae - Australasian marsupials Ricinidae - Hummingbirds Avian Amblycera 64 57 87 73 1 1 61 1 75 61 3 100 99 9 93 85 3 100 12 100 15 96 88 100 98 7 8 89 76 4 3 5 90 74 2 1 1 62 2 98 96 1 1 1 1 1 1 52 3 2 80 57 4 92 75 5 51 1 3 1 2 1 2 1 2 2 1 1 2 Trochiloecetes rupununi Ricinus fringillae Trochiliphagus abdominalis Laemobothrion maximum Therodoxus oweni Boopia tarsata Paraheterodoxus insignis Latumcephalum lesouefi/macropus Paraboopia flava Rediella mirabilis Actornithophilus uniseriatus Plegadiphilus threskiornis Chapinia robusta Bonomiella columbae Pseudomenopon pilosum Ancistrona vagelli Dennyus hirundinis Myrsidea victrix Holomenopon brevithoracicum Eidmanniella pellucida Machaerilaemus laticorpus/latifrons Austromenopon crocatum Neomenopon pteroclurus Hohorstiella lata Osborniella crotophagae Eomenopon denticulatum Piagetiella bursaepelecani Ciconiphilus quadripustulatus Cuculiphilus fasciatus Ardeiphilus trochioxus Odoriphila clayae/phoeniculi Psittacomenopon poicephalus Colpocephalum zebra Comatomenopon elbeli/elongatum Gruimenopon longum Hoazineus armiferus Meromenopon meropis Trinoton anserinum Menacanthus stramineus Colimenopon urocolius Somaphantus lusius Amyrsidea ventralis Menopon gallinae Numidicola antennatus B L M B C D A Liposcelis bostrychophilus Marshall, 2003 44 genera, 147 char. (Morphology)

10 Psocoptera Menacanthus / Menopon -complex Ricinidae Laemobothriidae Boopiidae Austromenopon -complex Dennyus -complex Colpocephalum -complex Collected Eckfeld maar near Eifel, Germany Middle Eocene (Middle Lutetian, 44.3±0.4 Ma) First fossil louse Completes ordinal representation of fossil insects? Excellent preservation Phylogenetic affinities with modern feather lice Parasite of Anseriformes or Charadriiformes Crown group position Points to a long coevolutionary history with birds Suggests lice are very old Useful molecular calibration point First Fossil Louse (Amblycera: Menoponidae) Wappler, Smith & Dalgleish, In prep.

11 Ischnoceran Families GenusFamilySuborder 3 532 3,060 1,344 Mammals Birds Amblycera Ischnocera Rhyn… Anoplura Ischnocera Trichodectidae Heptapsogasteridae Philopteridae Hopkins & Clay, 1951 cf. Eicher, 1961 21 families Smith, 2000 Generic level phylogeny of Goniodidae & Heptapsogasteridae (Morphology) Heptapsogasteridae Goniodidae

12 Avian Ischnoceran Phylogeny Columbicola columbae Austrogoniodes waterstoni Harrisoniella hopkinsi Haffneria grandis Paraclisis diomedea Perineus nigrolimbatus Halipeurus pelagicus Philoceanus garrodiae Pseudonirmus gurlti Naubates fuliginosus Pelmatocerandra setosa Ardeicola smithersi Anaticola crassicornis Pectinopygus bassani Pectinopygus sulae Aquanirmus australis Ibidoecus plataleae Trabeculus schillingi Craspedonirmus colymbinus Quadraceps coenocoryphae Docophoroides brevis Saemundssonia desolata Saemundssonia haematopi Podargoecus strigoides Alcedoecus delphax Craspedorrhynchus platystomus Strigiphilus vapidus Sturnidoecus sturni Philopterus ornatus Vernoniella guimarãesi Acidoproctus rostratus Acidoproctus hilli Rallicola lugens Osculotes curtus Osculotes macropoda Coloceras damicorne Campanulotes bidentatus Goniocotes gallinae Goniodes pavonis Goniodes kéleri Chelopistes guttatus Strongylocotes angulocapitis Discocorpus c. cephalosus Splendoroffula ruwenzorornis Oxylipeurus dentatus Cuclotogaster madagascariensis Lipeurus caponis Degeeriella rufa Upupicola upupae Lagopoecus affinis Syrrhaptoecus falcatus Neopsittaconirmus borgiolii Paragoniocotes rotundus Brüelia semiannulata Falcolipeurus affulgeus Archolipeurus nandu Felicola (F.) viverriculae Geomydoecus (G.) heaneyi Trichodectes (T.) melis Bovicola limbatus Damalinia (D.) crenelata Smith, 2001 Generic level phylogeny Approx. 1/3 rd of all ischnoceran genera 51 genera, 138 characters (Morphology) Philoceanus -complex Goniodidae Degeeriella -complex Trichodectidae Lipeurus -complex Heptap. Saemundssonia -complex Philopterus -complex Multiple clades of lice on the same host

13 A. Circumfasciate Head B. Non-Circumfasciate Head N IN IIN IIIAdult C. Wing LiceD. Body LiceE. Head Lice Characters Defining Major Ischnoceran Clades

14 Philoceanus -complex Goniodidae Degeeriella -complex Trichodectidae Lipeurus -complex Heptap. Saemundssonia -complex Philopterus -complex Wing Lice Wing Lice CI: 0.5 RI: 0.95 Head Lice Body Lice Circumfasciate Non- circumfasciate CI: 1.0 RI: 1.0 Characters Defining Major Ischnoceran Clades Multiple clades occupy different host niches Incongruent with molecular data Morphological convergence Molecular data suggests many of these pairs are more closely related Character displacement? Sympatric speciation? Problems with Morphology! Smith et al, 2003

15 Rhynchophthirina: Haematomyzidae GenusFamilySuborder 3 532 3,060 1,344 Amblycera Ischnocera Anoplura Rhynchophthirina

16 Rhynchophthirina: Haematomyzidae Just 3 species in 1 family Present on elephants, wart hog & bush pig (perhaps also mammoths?) Monophyly & sister placement with Anoplura well supported Chewing lice (have outwardly opposing mandibles) Pool blood feeders (telmophages) cf vessel feeding Anoplura Spread via shared use of watering holes? Possible cryptic species? (morphologically identical species on African & Asian elephants) Specimens hard to come by! Haematomyzus elephantis ex Asian / African elephant Haematomyzus elephantis Haematomyzus hopkinsi Haematomyzus porci Part of Smith, in prep. 51 genera, 76 species 184 characters (Morphology)

17 3 532 3,060 1,344 Amblycera Ischnocera Rhyn… Anoplura Anopluran Families Anoplura At least 15 distinct families

18 Anoplura: familial phylogeny 541spp. in 15 families Patchily distributed across mammals Based on Kim, 1988 47 genera, 39 characters (Morphology) Monophyly of all 15 families supported 3 basal clades Many lice of host groups (including Primates) paraphyletic

19 Kims dataset re-analysed: Kim, 1988 reanalysed Strict Consensus Length=219 (CI=0.26, RI=0.62) OUTGROUP Polyplacidae 1 Polyplacidae 2 Polyplacidae 3 Hoplopleuridae Polyplacidae 4-12 Echinophthiriidae Enderleinellidae Linognathidae 1 Neolinognathidae Linognathidae 2 Pedicinidae Pthiridae Polyplacidae 13 Ratemiidae Hamophthiriidae Hybophthiridae Pediculidae Pecaroecidae Haematopinidae Microthoraciidae Polyplacidae 1 Polyplacidae 2 Polyplacidae 3 Polyplacidae 4-12 Linognathidae 1 Linognathidae 2 Polyplacidae 13 Paraphyly of 2 of the 5 families that were not monogenic Only 3 other nodes are compatible with Kims tree Just 1 node agrees with Kims original tree!

20 Haematomyzus elephantis Haematomyzus hopkinsi Pthirus gorillae Pthirus pubis Pediculus schaeffi Pediculus humanus Pediculus mjobergi Pediculus humanus var. americanus Scipio aulacodi Scipio tripedatus Neolinognathus sp. Neolinognathus elephantuli Solenopotes muntiacus Prolinognathus aethiopicus Prolinognathus ferrisi Linognathus setosus Linognathus aepycerus Linognathus africanus Latagophthirus rauschi Echinophthirius horridus Lepidophthirus macrorhini Antarctophthirus callorhini Antarctophthirus microchir Proechinophthirus fluctus Proechinophthirus zumpti Johnsonpthirus chlorotalpae Johnsonpthirus heliosciuri Lemurphthirus galagus Lemurphthirus stigmosus Linognathoides laeviusculus Sathrax durus Neohaematopinus callosciuri Neohaematopinus inornatus Abrocomaphthirus chilensis Ctenophthirus cercomydis Proenderleinellus calva Polyplax abyssinica Polyplax spinulosa Docophthirus acinetus Eulinognathus denticulatus Haemodipsus leporis Haemodipsus lyriocephalus Lagidiophthirus parvus Fahrenholzia ferrisi Fahrenholzia pinnata Galeophthirus caviae Phthirpediculus brygooi Phthirunculus sumatranus Atopophthirus emersoni Microphthirus uncinatus Enderleinellus osborni Werneckia nigeriensis Hoplopleura acanthopus Paradoxophthirus emarginata Hoplopleura pacifica Hoplopleura irritans Pterophthirus audax Pterophthirus splendida Ancistroplax crocidurae Haematopinoides squamosus Typhlomyophthirus bifoliatus Schizophthirus graphiuri Schizophthirus pleurophaeus Haematomyzus porci Microthoracius cameli Ratemia squamulata Hamophthirius galeopitheci Hybophthirus notophallus Haematopinus asini Haematopinus suis Pecaroecus javalii Pedicinus ferrisi Pedicinus patas Anoplura: phylogeny Haematomyzidae Linognathidae Polyplacidae 1 Neolinognathidae Enderleinellidae Hoplopleuridae Polyplacidae 2 Polyplacidae 3 Echinophthiriidae Microthoraciidae Ratemiidae Hamophthiriidae Hybophthiridae Haematopinidae Pecaroecidae Pedicinidae Pthiridae Pediculidae Smith, in prep. 51 genera, 76 species 184 characters (Morphology) Why are rodents so lousy? Why are other mammals relatively louse free cf. birds Good evidence for cophylogeny in some clades Comparative studies underway using these trees

21 Louse Molecular Studies Nuclear - EF1- (350-450bp) - 18S rRNA (1500+bp) - Wingless, PEPCK, DDC Hard to amplify Multiple copies RNA alignment issues Utility varies between genes & taxa Mitochondrial - COI (400-1500bp) - 12S rRNA (450bp) - Cyt. B (450bp) - ND4 (500bp) Easy to amplify Limited phylogenetic utility outside families Often restricted to use within closely related genera Need New Genes!

22 Lice all sites pos. 1 pos. 2 pos. 3 all sites pos. 1 pos. 2 pos. 3 all sites pos. 1 pos. 2 pos. 3 1 1 0.12 129.9 1 1 0.15 169.78 1 1 0.0001 44.79 0.018 0.013 0.016 0.41 0.016 0.039 0.017 0.8 0.25 0.20 0.0001 2.49 55.6 76.9 7.5 316.8 61.5 25.6 8.8 212.2 4.1 5.0 1.0 18.0 Physconelloidinae Columbicola aphids Data Set Sites Ratio (COI:EF1 COI EF1- Rate Parameter Relative Substitution Rate (Mit. COI vs nuc. EF1- )

23 Louse 12S Secondary Structure mit. 12S rRNA Page et al, 2002

24 0 20 40 60 80 100 120 Length 35404550556065707580 0 1 2 3 4 5 6 7 8 9 10 Length 0 20 40 60 80 100 120 140 160 180 200 Length 152025303540455055606570758085 0 2 4 6 8 10 12 14 Length Segment 3: 38'-39' Segment 5: 34'-33' 0 20 40 60 80 100 120 140 160 Length 51015202530354045 0 2 4 6 8 10 12 Length Segment 2: 36-38Segment 1: 33-33' 0 20 40 60 80 100 120 140 160 Length insects 210220230240250260270280290300310320 0 1 2 3 4 5 6 7 Length lice 12S Length Variation Insects vs. Lice Page et al, 2002

25 cox1cox2 atp8atp6 cox3 nad3 nad5nad4 nad4Lnad6 cobnad2rrnLrrnSnad2 L2L2 KDGHARNS 1 EFTPVS2S2 L1L1 IQMWCY cox1cox3nad1 cob nad4 nad4L nad2 nad3 nad5cox2rrnLrrnS atp6atp8nad6 S2S2 L2L2 KNAYQVPTDIFS 1 CMWERIIGL 1 Gene arrangement of Heterodoxus macropus (Amblycera) Inferred gene arrangement of ancestral insect Adapted from Shao, Campbell & Barker, 2001 Louse Mitochondrial Genome

26 Louse Molecular Studies Nuclear - EF1- (350-450bp) - 18S rRNA (1500+bp) - Wingless, PEPCK, DDC Hard to amplify Multiple copies RNA alignment issues Utility varies between genes & taxa Mitochondrial - COI (400-1500bp) - 12S rRNA (450bp) - Cyt. B (450bp) - ND4 (500bp) Easy to amplify Limited phylogenetic utility outside families Often restricted to use within closely related genera Need New Genes!

27 ISCHNOCERA AMBLYCERA ANOPLURA RHYN… Louse Nuclear Genes Cruickshank et al, 2001 350 bp single copy EF1- Limited phylogenetic utility above families What Not to do!

28 Phylogenetic Utility 18S rRNA Smith et al, 2003

29 Much recent progress on basal phthirapteran phylogeny Monophyly of suborders well supported Complex pattern of host-louse relationships Distinctive molecular evolution in lice Incongruence between molecules & morphology Ongoing Questions… To what extent do lice cospeciate with their hosts? Why are some hosts more lousy than others? How old are the major louse clades? Are Phthiraptera truly paraphyletic? Conclusions…

30 Acknowledgements Collaboration Stephen Barker Dale Clayton Tom Ford Kevin Johnson Rod Page Isabel Marshall David Reed Funding Wellcome Trust, UK Natural Environment Research Council, UK

31 0.60.50.40.30.20.10 0 100 200 300 ML Corrected Divergences (EF1- 3 rd Positions) ML Corrected Divergences (COI 3 rd Positions) Slope approximates relative rate Around 1000:1 !!! Pairwise ML Corrected Distances for 3 rd Sites (Nuclear EF1- against Mit. COI) Johnson et al, 2003

32 18S – not a panacea for basal louse Phylogeny! Uncorrected NJ Bayesian 0.1 Anoplura Ischnocera Amblycera Liposcelis sp. Trogiomorph Psocoptera


Download ppt "Lousy Phylogenies: Phthiraptera systematics and the antiquity of lice Division of Environmental and Evolutionary Biology Institute of Biomedical and Life."

Similar presentations


Ads by Google