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Elephant, warthog & bush pig lice. Haematomyzus elephantis. 3 spp. in a single family ... (elephants, warthog & bush pig) Spread via shared use of watering ...
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The Diversification of Lice: Phylogeny Vincent S. Smith & Kevin P. Johnson
Book lice (Psocoptera) Lice (Phthiraptera) Chewing Lice“Mallophaga” Sucking Lice(Anoplura) [ Liposcelis sp.] Amblycera Ischnocera Rhynchophthirina Anoplura Louse phylogeny four years on… • 40+ phylogenies! • Now cover 80% of louse genera • Mix of molecules & morphology • Beginning to reconcile differences • Still a long way to go! • Some of the biggest studies unpublished
Louse phylogeny four years on… • 40+ phylogenies! • Now cover 80% of louse genera • Mix of molecules & morphology [ Liposcelis sp.] • Beginning to reconcile differences Amblycera Ischnocera Rhynchophthirina Anoplura • Still a long way to go! • Some of the biggest studies unpublished
Menoponidae Birds Worldwide & Australasian Marsupials Boopiidae • Few families on South American mammals & marsupials Laemobothriidae Ricinidae Trimenoponidae Protogyropinae South/Central Am. Rodents & Marsupials Gyropidae Gyropinae Grilicolinae Liposcelidae (Psocoptera) Amblycera “Feather lice” • 1,350 spp. in 6 families • Widespread on birds 85 • Boopiidae on Australasian marsupials • No new deep-branch phylogenies! • Mostly alpha-taxonomic work 84 • No extensive molecular trees Clay, 1970
Ricinidae 100 100 93 - Hummingbirds 85 12 Laemobothriidae 3 - Falconiforms 100 100 98 100 8 61 75 15 61 1 100 96 99 3 88 9 7 89 Boopidae Dennyus-complex Austromenopon-complex Colpocephalum-complex 76 90 4 - Australasian marsupials - Birds worldwide - Aquatic birds - Birds worldwide 74 5 1 62 2 2 98 96 3 1 1 1 1 1 1 52 1 3 1 1 80 2 57 92 4 75 1 5 51 87 73 1 3 2 1 Menacanthus/Menopon-cpx. 64 2 57 - Birds worldwide 1 1 2 2 1 2 Amblycera “Feather lice” Liposcelis bostrychophilus Trochiloecetes rupununi Ricinus fringillae Trochiliphagus abdominalis Laemobothrion maximum L Therodoxus oweni Boopia tarsata Paraheterodoxus insignis B Latumcephalum lesouefi/macropus Paraboopia flava Rediella mirabilis Actornithophilus uniseriatus Plegadiphilus threskiornis Chapinia robusta Bonomiella columbae Pseudomenopon pilosum Ancistrona vagelli Dennyus hirundinis A Myrsidea victrix Holomenopon brevithoracicum Eidmanniella pellucida Machaerilaemus laticorpus/latifrons Austromenopon crocatum Neomenopon pteroclurus B M Hohorstiella lata Osborniella crotophagae Eomenopon denticulatum Piagetiella bursaepelecani Ciconiphilus quadripustulatus Cuculiphilus fasciatus Ardeiphilus trochioxus Odoriphila clayae/phoeniculi Psittacomenopon poicephalus C Colpocephalum zebra Comatomenopon elbeli/elongatum Gruimenopon longum Marshall, 2003 Hoazineus armiferus Meromenopon meropis 44 genera, 147 char. (Morphology) Trinoton anserinum Menacanthus stramineus Colimenopon urocolius Parsimony, Strict consensus Somaphantus lusius Amyrsidea ventralis D Menopon gallinae Numidicola antennatus
Philoceanus -complex Saemundssonia -complex Philopterus -complex • Historically the least studied Goniodidae • Now comparatively well known Heptap. Lipeurus -complex • Extensive molecular & morphological work Degeeriella -complex Smith, 2001 41 genera, 138 char. (Morphology, parsimony, strict consensus) Trichodectidae Ischnocera “Feather lice” Haffneria grandis Harrisoniella hopkinsi Paraclisis diomedea Perineus nigrolimbatus • 3,104 spp. in an uncertain no. of families Halipeurus pelagicus Philoceanus garrodiae Pseudonirmus gurlti Naubates fuliginosus • Widespread on birds, 381 spp on mammals Pelmatocerandra setosa Ardeicola smithersi Anaticola crassicornis Pectinopygus bassani • Highly host specific (71% on a single host species) 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 Austrogoniodes waterstoni Strongylocotes angulocapitis Discocorpus c. cephalosus Columbicola columbae 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 Geomydoecus (G.) heaneyi Felicola (F.) viverriculae Trichodectes (T.) melis Bovicola limbatus Damalinia (D.) crenelata
Wing Non-circumfasciate - multiple clades of lice on the same host Head A. Circumfasciate B. Non-Circumfasciate Body Circumfasciate N I N II N III Adult Wing C. Wing D. Body E. Head CI: 0.5 RI: 0.95 CI: 1.0 RI: 1.0 Ischnocera “Feather lice” • Morphological phylogeny defined by: - shape of the head - body form (wing, head or body louse) • Major clades incongruent with the molecular data!
Smith & Johnson, in prep. All ischnceran genera, Adult & Nymphal Morph. Strict Consensus • Trust me, the same pattern holds! Ischnocera “Feather lice”
Haematomyzus elephantis Haematomyzus hopkinsi Haematomyzus porci • Phylogeny ? • Only 1 species available for molecular work Rhynchophthirina Elephant, warthog & bush pig lice • 3 spp. in a single family • Confined to relatively unrelated hosts (elephants, warthog & bush pig) • Spread via shared use of watering holes? • Still chewing lice (telmophages) Haematomyzus elephantis ex Asian / African elephant
51 genera, 76 species • 184 morphological characters • Well supported • Consistent with limited mol. data • Unpublished Anoplura Sucking lice Rhynchophthirina • 540 species in 15 families Hybophthiridae Haematopinidae Ratimidae • Confined to mammals (834 host species) Linognathidae Neolinognathidae Microthoracidae • 50% confined to a single host Echinophthiridae • Blood vessel feeders (solenophages) Hamophthiridae Pecaroecidae Pedicinudae Pthiridae Pediculidae Polyplacidae Enderleinellidae Smith, Strict Consensus In Prep. Hoplopleuridae
All 15 families monophyletic except Polyplacidae • Other Anoplura grade into • Other Anoplura grade into the rodent lice the rodent lice • Strong correlation with host taxonomy • Major clades are: - primate lice - seal lice - ungulate lice • Many monogenic families Typhlomyophthirus Anoplura Sucking lice • Rodent lice monophyletic Rhynchophthirina Hybophthiridae Haematopinidae Ratimidae Linognathidae Neolinognathidae Microthoracidae Echinophthiridae Hamophthiridae Pecaroecidae Pedicinudae Pthiridae Pediculidae Polyplacidae Enderleinellidae Hoplopleuridae
Pediculidae Pthiridae Pedicinidae Reed, Smith et al, 2004 Anoplura Sucking lice • Focus on Primate lice Rhynchophthirina Hybophthiridae Haematopinidae Ratimidae Linognathidae Neolinognathidae Microthoracidae Echinophthiridae Hamophthiridae Pecaroecidae Pedicinudae Pthiridae Pediculidae Polyplacidae Enderleinellidae Hoplopleuridae
Pediculus humanus Human head & body louse morphology Head Lice Body Lice Pediculus capitis De Geer, 1767 Pediculus humanus Linnaeus, 1758 Larger & lighter Smaller & darker Antennae longer & slender Antennae shorter & thicker Middle leg tibial length shorter Middle leg tibial length longer Weak indentations between abdominal segments Below neck. Common in the seams of clothing (especially females), particularly those worn next to the skin (shirts, vests etc.). Males more commonly over garments surface. Restricted to the hair on the scalp, particularly on the nape and in the area behind the ears. Not on the eyebrows or eyelashes. Does not “naturally” vector disease Vectors relapsing fever, trench fever, & epidemic typhus Traditionally considered indistinguishable! Killed no one Killed millions Vectors a category B bioterrorism agent Does not vector a bioterrorism agent
HdWdth PtWdth LgTibLgth TBL TBL: Total Body Length HdWdth: Postantennal Head Width PtWdth: Posterior Pterothoracic Width LgTibLgth: 2nd Leg Tibial Length Pediculus humanus Four discriminating characters
Females Males 6 6 4 4 1 1 2 2 Function 2 0 0 3 Group Centroids 2 Ungrouped Cases -2 -2 3 2 3 Pubic Lice -4 -4 2 Head Lice 1 Body Lice -6 -6 -10 -5 0 5 10 15 -10 -5 0 5 10 15 Function 1 Pediculus humanus Discriminant function analysis Reed, Smith et al, 2004 We can morphologically separate head & body lice I want your pill-box specimens!!!
Females Males 6 6 4 4 1 1 2 2 Function 2 0 0 3 Group Centroids 2 Ungrouped Cases -2 -2 3 2 3 Pubic Lice -4 -4 2 Head Lice 1 Body Lice -6 -6 -10 -5 0 5 10 15 -10 -5 0 5 10 15 Function 1 Pediculus humanus Discriminant function analysis Reed, Smith et al, 2004 Are these differences present in the molecular data?