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V. S. Smith & Kevin Johnson. The Diversification of Lice: Phylogeny
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The Diversification ofLice: Phylogeny
Vincent S. Smith& Kevin P. Johnson
Louse phylogeny four years on…
Book lice(Psocoptera)
Lice(Phthiraptera)
AmblyceraAmblycera IschnoceraIschnocera RhynchophthirinaRhynchophthirinaAnopluraAnoplura[ [ LiposcelisLiposcelis
sp.]sp.]
Chewing Lice“Mallophaga”
Sucking Lice(Anoplura)• 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• Some of the biggest studies unpublished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…
AmblyceraAmblycera IschnoceraIschnocera RhynchophthirinaRhynchophthirinaAnopluraAnoplura[ [ LiposcelisLiposcelis
sp.]sp.]
• 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• Some of the biggest studies unpublished
Amblycera
“Feather lice”
• No new deep-branch phylogenies!
- 1,350 spp. in 6 families- Widespread on birds- Few families on South American
mammals & marsupials- Boopiidae on Australasianmarsupials
• Mostly alpha-taxonomic work
Clay, 1970
• No extensive molecular trees
84
Grilicolinae
Gyropinae
Protogyropinae
Trimenoponidae
Ricinidae
Laemobothriidae
Boopiidae
Menoponidae
Liposcelidae (Psocoptera)
Gyropidae
85
Amblycera
“Feather lice”
• No new deep-branch phylogenies!
- 1,350 spp. in 6 families- Widespread on birds- Few families on South American
mammals & marsupials- Boopiidae on Australasianmarsupials
• Mostly alpha-taxonomic work
Clay, 1970
• No extensive molecular trees
84
Grilicolinae
Gyropinae
Protogyropinae
Trimenoponidae
Ricinidae
Laemobothriidae
Boopiidae
Menoponidae
Liposcelidae (Psocoptera)
Gyropidae
Birds Worldw
ide & Australasian M
arsupialsSouth/C
entral Am.
Rodents & M
arsupials
85
Amblycera
“Feather lice”
Laemobothriidae- Falconiforms
Menacanthus/Menopon-cpx.- Birds worldwide
Colpocephalum-complex- Birds worldwide
Austromenopon-complex- Aquatic birds
Dennyus-complex- Birds worldwide
Boopidae- Australasian marsupials
Ricinidae- Hummingbirds
6457
8773
1
1
611
7561
3100 999
93853
10010012
10010015
9688
100 98
7
8
8976
4
3
5
9074
2
1
1 622
9896
11
1
11
1
523
2 80574
92755 51
13
1
2
1
2
1
2
2
1
12
Trochiloecetes rupununiRicinus fringillaeTrochiliphagus abdominalisLaemobothrion maximumTherodoxus oweniBoopia tarsataParaheterodoxus insignisLatumcephalum lesouefi/macropusParaboopia flavaRediella mirabilisActornithophilus uniseriatusPlegadiphilus threskiornisChapinia robustaBonomiella columbaePseudomenopon pilosumAncistrona vagelliDennyus hirundinisMyrsidea victrixHolomenopon brevithoracicumEidmanniella pellucidaMachaerilaemus laticorpus/latifronsAustromenopon crocatumNeomenopon pteroclurusHohorstiella lata
Osborniella crotophagaeEomenopon denticulatumPiagetiella bursaepelecaniCiconiphilus quadripustulatusCuculiphilus fasciatusArdeiphilus trochioxusOdoriphila clayae/phoeniculiPsittacomenopon poicephalusColpocephalum zebraComatomenopon elbeli/elongatum
Gruimenopon longumHoazineus armiferusMeromenopon meropisTrinoton anserinumMenacanthus stramineusColimenopon urocoliusSomaphantus lusiusAmyrsidea ventralisMenopon gallinaeNumidicola antennatus
B
L
M B
C
D
A
Liposcelis bostrychophilus
Marshall, 200344 genera, 147 char.(Morphology)Parsimony, Strictconsensus
Ischnocera
“Feather lice”
- 3,104 spp. in an uncertain no. of families- Widespread on birds, 381 spp on mammals- Highly host specific (71% on a single host species)
Columbicola columbae
Austrogoniodes waterstoni
Harrisoniella hopkinsiHaffneria grandis
Paraclisis diomedeaPerineus nigrolimbatusHalipeurus pelagicusPhiloceanus garrodiaePseudonirmus gurltiNaubates fuliginosusPelmatocerandra setosaArdeicola smithersiAnaticola crassicornisPectinopygus bassaniPectinopygus sulaeAquanirmus australisIbidoecus plataleaeTrabeculus schillingiCraspedonirmus colymbinusQuadraceps coenocoryphaeDocophoroides brevisSaemundssonia desolataSaemundssonia haematopiPodargoecus strigoidesAlcedoecus delphaxCraspedorrhynchus platystomusStrigiphilus vapidusSturnidoecus sturniPhilopterus ornatusVernoniella guimarãesiAcidoproctus rostratusAcidoproctus hilliRallicola lugensOsculotes curtusOsculotes macropodaColoceras damicorneCampanulotes bidentatusGoniocotes gallinaeGoniodes pavonisGoniodes kéleriChelopistes guttatus
Strongylocotes angulocapitisDiscocorpus c. cephalosus
Splendoroffula ruwenzorornisOxylipeurus dentatusCuclotogaster madagascariensisLipeurus caponisDegeeriella rufaUpupicola upupaeLagopoecus affinisSyrrhaptoecus falcatusNeopsittaconirmus borgioliiParagoniocotes rotundusBrüelia semiannulataFalcolipeurus affulgeusArcholipeurus nandu
Felicola (F.) viverriculaeGeomydoecus (G.) heaneyi
Trichodectes (T.) melisBovicola limbatusDamalinia (D.) crenelata
Smith, 200141 genera, 138 char.
(Morphology, parsimony,strict consensus)
Ischnocera
“Feather lice”
- 3,104 spp. in an uncertain no. of families- Widespread on birds, 381 spp on mammals- Highly host specific (71% on a single host species)
Columbicola columbae
Austrogoniodes waterstoni
Harrisoniella hopkinsiHaffneria grandis
Paraclisis diomedeaPerineus nigrolimbatusHalipeurus pelagicusPhiloceanus garrodiaePseudonirmus gurltiNaubates fuliginosusPelmatocerandra setosaArdeicola smithersiAnaticola crassicornisPectinopygus bassaniPectinopygus sulaeAquanirmus australisIbidoecus plataleaeTrabeculus schillingiCraspedonirmus colymbinusQuadraceps coenocoryphaeDocophoroides brevisSaemundssonia desolataSaemundssonia haematopiPodargoecus strigoidesAlcedoecus delphaxCraspedorrhynchus platystomusStrigiphilus vapidusSturnidoecus sturniPhilopterus ornatusVernoniella guimarãesiAcidoproctus rostratusAcidoproctus hilliRallicola lugensOsculotes curtusOsculotes macropodaColoceras damicorneCampanulotes bidentatusGoniocotes gallinaeGoniodes pavonisGoniodes kéleriChelopistes guttatus
Strongylocotes angulocapitisDiscocorpus c. cephalosus
Splendoroffula ruwenzorornisOxylipeurus dentatusCuclotogaster madagascariensisLipeurus caponisDegeeriella rufaUpupicola upupaeLagopoecus affinisSyrrhaptoecus falcatusNeopsittaconirmus borgioliiParagoniocotes rotundusBrüelia semiannulataFalcolipeurus affulgeusArcholipeurus nandu
Felicola (F.) viverriculaeGeomydoecus (G.) heaneyi
Trichodectes (T.) melisBovicola limbatusDamalinia (D.) crenelata
Philoceanus-complex
Goniodidae
Degeeriella-complex
Trichodectidae
Lipeurus-complex
Heptap.
Saemundssonia-complex
Philopterus-complex
• Historically the least studied
• Now comparatively well known
• Extensive molecular & morphological work Smith, 2001
41 genera, 138 char.(Morphology, parsimony,
strict consensus)
Ischnocera
“Feather lice”
• Morphological phylogeny defined by:- shape of the head- body form (wing, head or body louse)
A. Circumfasciate
B. Non-Circumfasciate
N I N II N III Adult
C. Wing D. Body E. Head
Ischnocera
“Feather lice”
• Morphological phylogeny defined by:- shape of the head- body form (wing, head or body louse)
Wing
CI: 0.5RI: 0.95
Head
Body
Wing
Circumfasciate
Non-circumfasciate
CI: 1.0RI: 1.0
Ischnocera
“Feather lice”
• Morphological phylogeny defined by:- shape of the head- body form (wing, head or body louse)- multiple clades of lice on the same host
Wing
CI: 0.5RI: 0.95
Head
Body
Wing
Circumfasciate
Non-circumfasciate
CI: 1.0RI: 1.0
Ischnocera
“Feather lice”
• Morphological phylogeny defined by:- shape of the head- body form (wing, head or body louse)- multiple clades of lice on the same host
Wing
CI: 0.5RI: 0.95
Head
Body
Wing
Circumfasciate
Non-circumfasciate
CI: 1.0RI: 1.0
• Major clades incongruent with the molecular data!
Ischnocera
“Feather lice”
Bovicola limbatusDamalinia (Damalinia) crenelata
Discocorpus c. cephalosusStrongylocotes angulocapitisStrongylocotes complanatusTrichodopeostusHeptapsogaster temporalis
Kelloggia conicepsLamprocorpus hirsutus
Megaginus sordidusMegapeostus asymmetricus
Nothocotus subsimilisOrnicholax robustus
Pectenosoma verrucosaPterocotes aberrans
Rhopaloceras rudimentariusCuclotocephalus extraneus
Physconella kelloggiChelopistes guttatus
Chelopistes meleagridisPachyskelotes orthopleurus
Goniodes pavonisGoniodes kÈleri
Passonomedea hopkinsiColoceras damicorne
Campanulotes comparAuricotes affinis
Goniocotes gallinaePhysconelloides cubanusLabicotes guttatusKodocephalon latumAustrogoniodes waterstoniTrichophilopterus babakotophilusPsittoecus vanzoliniiParagoniocotes rotundusNeopsittaconirmus borgioliiHarrisoniella hopkinsiHaffneria grandisHarrisoniella copeiPerineus nigrolimbatusEpisbates pederiformisBedfordiella unicaHalipeurus pelagicusNaubates fuliginosusPhiloceanus garrodiaePseudonirmus gurltiPelmatocerandra setosaParaclisis diomedeaArdeicola smithersiArdeicola elongata
Fulicoffula longipilaFurnariphilus pageiAnaticola crassicornis
Pseudolipeurus similisPseudophilopterus hirsutus
Psittaconirmus (Psittaconirmus) zinkiEmersoniella bracteata
Hopkinsiella clavigeraRotundiceps cordatus
Trogoniella aequatorialeCuclotogaster madagascariensis
Lipeurus caponisOxylipeurus dentatus
Splendoroffula ruwenzorornisColumbicola columbaeTurturicola salimaliiColilipeurus colius
Colinicola mearnsiSyrrhaptoecus falcatus
Degeeriella rufaLagopoecus affinis
Cotingacola rupicolaeCuculicola atopus
Megapodiella nakataeUpupicola upupae
Buceroemersonia clarkeiAcutifrons chimango
Picicola snodgrassiBuceronirmus deignani
Buceronirmus
Trogoninirmus strigilatusBucorvellus docophorus
CapraiellaCapraiella sabzak
OtidoecusParagoniocotes venezolanus
Paroncophorus javanicusTinamotaecola
Centropodiella borneoensisRhynonirmus scolopacis
Pectinopygus bassaniPectinopygus sulae
Wilsoniella absitaTrabeculus schillingiDocophoroides brevis
Craspedonirmus colymbinusCarduiceps cingulatus
Quadraceps coenocoryphaeSaemundssonia desolataSaemundssonia haematopi
Incidifrons fulicaeMeropoecus meropis
Anatoecus icterodesAlcedoffula alcyonae
Formicaricola willisiAquanirmus australisCraspedorrhynchus platystomus
Strigiphilus vapidusPodargoecus strigoidesAlcedoecus delphaxAegypoecus brevicollisForficuloecus emersoniSturnidoecus sturniCuculoecus latifronsEchinophilopterus protrususPhilopterus ornatusNyctibicola longirostrisPenenirmus auritusCummingsiella ambiguaArdeiphagus cochleariusBizarrifrons magusBuerelius longicepsIncidifrons transpositusCirrophthirius testudinariusLunaceps numeniiFormicaphagus pittasomaeTheresiella geminaMulcticolaTurnicola angustissimusIbidoecus plataleae
Neophilopterus heteropygusAustrophilopterus spinosus
Bucerocophorus pachycnemisVernoniella guimar„esi
Osculotes curtusOsculotes macropoda
Rallicola lugensBr¸elia semiannulata
Brueelia spBBrueelia ornatissima
Pseudocophorus perijanusAcidoproctus rostratusAcidoproctus hilli
Ornithobius goniopleurusBothriometopus macrocnemisEsthiopterum giganteumFalcolipeurus affulgeusArcholipeurus nanduStruthiolipeurus struthionisMeinertzhageniella lataFalcolius elbeliDahlemhornia asymmetricaNesiotinus demersusTrichodectes (Trichodectes) melisFelicola (Felicola) viverriculae
Geomydoecus (Geomydoecus) heaneyi
• Trust me, the same pattern holds!
Smith & Johnson, in prep.All ischnceran genera,
Adult & Nymphal Morph.Strict Consensus
RhynchophthirinaElephant, warthog & bush pig lice
Haematomyzus elephantis
- 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)
ex Asian / African elephantex Asian / African elephant
• Phylogeny
• Only 1 species available for molecular work
Haematomyzus elephantisHaematomyzus hopkinsiHaematomyzus porci
?
AnopluraSucking lice
RhynchophthirinaHybophthiridaeHaematopinidaeRatimidae
Linognathidae
NeolinognathidaeMicrothoracidae
Echinophthiridae
HamophthiridaePecaroecidaePedicinudaePthiridaePediculidae
Polyplacidae
Enderleinellidae
Hoplopleuridae
• 51 genera, 76 species
• 184 morphological characters
• Well supported
• Consistent with limited mol. data
• UnpublishedSmith, StrictConsensus
In Prep.
- 540 species in 15 families- Confined to mammals
(834 host species)- 50% confined to a single host
- Blood vessel feeders (solenophages)
• Other Anoplura grade intothe rodent lice
- ungulate lice- seal lice
• Major clades are:- primate lice
AnopluraSucking lice
• Many monogenic families
RhynchophthirinaHybophthiridaeHaematopinidaeRatimidae
Linognathidae
NeolinognathidaeMicrothoracidae
Echinophthiridae
HamophthiridaePecaroecidaePedicinudaePthiridaePediculidae
Polyplacidae
Enderleinellidae
Hoplopleuridae
• Rodent lice monophyletic
• All 15 families monophyleticexcept Polyplacidae
Typhlomyophthirus
• Strong correlation with hosttaxonomy
• Other Anoplura grade intothe rodent lice
AnopluraSucking lice
RhynchophthirinaHybophthiridaeHaematopinidaeRatimidae
Linognathidae
NeolinognathidaeMicrothoracidae
Echinophthiridae
HamophthiridaePecaroecidaePedicinudaePthiridaePediculidae
Polyplacidae
Enderleinellidae
Hoplopleuridae
• Focus on Primate lice
Pediculidae Pthiridae Pedicinidae
Reed, Smithet al, 2004
Pediculus humanusHuman head & body louse morphology
Head Lice Body LicePediculus humanus Linnaeus, 1758Pediculus capitis De Geer, 1767
Smaller & darker Larger & lighter
Antennae shorter & thicker Antennae longer & slender
Middle leg tibial length shorter Middle leg tibial length longer
Weak indentations betweenabdominal segments
Restricted to the hair on thescalp, particularly on the nape
and in the area behind the ears.Not on the eyebrows or
eyelashes.
Below neck. Common in theseams of clothing (especiallyfemales), particularly thoseworn next to the skin (shirts,
vests etc.). Males morecommonly over garments
surface.
Vectors relapsing fever, trenchfever, & epidemic typhus
Does not “naturally” vectordisease
Killed millionsKilled no one
Vectors a category Bbioterrorism agent
Does not vector a bioterrorismagent
Traditionallyconsidered
indistinguishable!
Pediculus humanusFour discriminating characters
TBLTBL
TBL: Total Body Length
LgTibLgthLgTibLgth
LgTibLgth: 2nd Leg Tibial Length
HdWdthHdWdth
HdWdth: Postantennal Head Width
PtWdthPtWdth
PtWdth: Posterior Pterothoracic Width
Pediculus humanusDiscriminant function analysis
Function 1
Females Males
Func
tion
2
Group Centroids
Ungrouped Cases
1
2 3
1
2
3
3 Pubic Lice
2 Head Lice
1 Body Lice-10 -5 0 5 10 15 -10 -5 0 5 10 15
6
4
2
0
-2
-4
-6
6
4
2
0
-2
-4
-6
Reed, Smithet al, 2004
I want your pill-box specimens!!!We can morphologically separate head & body lice
Pediculus humanus
Function 1
Females Males
Func
tion
2
Group Centroids
Ungrouped Cases
1
2 3
1
2
3
3 Pubic Lice
2 Head Lice
1 Body Lice-10 -5 0 5 10 15 -10 -5 0 5 10 15
6
4
2
0
-2
-4
-6
6
4
2
0
-2
-4
-6
Reed, Smithet al, 2004
Are these differences present in the molecular data?
Discriminant function analysis
Questions?