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Cryan Lab of Insect Phylogenetics

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Page 1: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cryan Lab of Insect Phylogenetics

Page 2: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Julie Urban, Ph.D., Ph.D.

• Urban, J.M., and J.R. Cryan. 2007. Evolution of the planthoppers (Insecta: Hemiptera: Fulgoroidea). Molecular Phylogenetics and Evolution 42: 556-572.

• Urban, J.M., and J.R. Cryan. In Press. Entomologically famous, yet evolutionarily unexplored: the first phylogeny of the Fulgoridae (Hemiptera: Fulgoridae). Molecular Phylogenetics and Evolution.

Systematics of FulgoroideaFulgoridaeBacterial endosymbionts

Page 3: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Gavin Svenson, Ph.D.

• Svenson, G.J., and M.F. Whiting. 2004. Phylogeny of Mantodea based on molecular data: evolution of a charismatic predator. Systematic Entomology 29: 359-370.

•Svenson, G.J., and M.F. Whiting. In review. Reconstructing the Origins of Praying Mantises (Dictyoptera, Mantodea): the roles of Gondwanan vicariance and morphological convergence. (Submitted to Cladistics Sept. 2008)

Systematics of MantodeaLarge-scale phylogeneticsAuditory physiology

Page 4: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Adam Bell (Ph.D. student)

Systematics of Cercopoidea

The “new guy”

Page 5: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

“leaf”-

& “treehoppers”(Membracoidea)

Page 6: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

“planthoppers”(Fulgoroidea)

Page 7: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

“spittlebugs”(Cercopoidea)

Page 8: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

BiogeographyWhere was the origin of these groups, and

what is the explanation for their current distributions?

Research QuestionsPhylogenetics

How are the higher taxonomic categoriesof Auchenorrhyncha related?

Morphological EvolutionAre the many unusual morphological and

physiological characters constrained?

Behavior / Ecology / Life HistoryWhat are the patterns of symbioses,

behavior, and host plant usage?

Page 9: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to
Page 10: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Membracidae

Cercopoidea

A?

Page 11: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Outgroups

Brachybelus cruralis

Callicentrus sp.

Centrodontus atlasCampylocentrus sp.

Eufairmairia fraterna

Gargara sp.Ischnocentrus niger

Ceraon vitta

Anchon sp.

Nessorhinus sp.

Platycentrus acuticornis

Pyrgauchenia sp.

Sextius virescens

Tylocentrus reticulatus

Deiroderes inermisLycoderes sp.

Microcentrus caryaeAntillotolania n.sp.

Cymbomorpha sp.Darnis latiorStictopelta sp.

Aconophora flavipes

Alchisme sp.

Cladonota apicalis

Calloconophora sp.

Enchenopa binotata

Guayaquila gracilicornis

Membracis sp.

Notocera sp.

Ochropepla moureiScalmophorus minutus

Undet. MembraciniTylopelta sp.

Umbonia crassicornis

Cyrtolobus arcuatus

Cyrtolobus fenestratus

Cyrtolobus fuliginosus

Cyrtolobus maculifrontis

Carynota mera

Cyrtolobus tuberosus

Ennya sp.

Glossonotus acuminatus

Harmonides sp.

Micrutalis calva

Metheisa lucillodes

Ophiderma definita

Ophiderma evelyna

Ophiderma flavicephalaOphiderma grisea

Ophiderma pubescens

Polyglypta sp.

Telamona monticolaTelamona unicolor

Xantholobus muticus

Acutalis tartareaCyphonia clavata

Stictocephalus brevitylusSpissistilus festinus

Stictocephala taurinaVestistilus variabilis

Antianthe sp.

Mem

bracidae

18S, 28S, EF-1α, Wg+ morphology

Stegaspidinae

Centrotinae +Centrodontinae

Smiliinae

Membracinae

Darninae

Page 12: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to
Page 13: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to
Page 14: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to
Page 15: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to
Page 16: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to
Page 17: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

4 mm

4 mm

Page 18: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to
Page 19: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

Cercopoidea Project

The Molecular Side

• Large‐scale phylogenetic investigation of neglected superfamily• Tests of classification at multiple taxonomic levels• Biogeography & Biology

Page 20: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

Taxon Sampling:• Target for NSF project is 500 taxa• 336 Total In‐group taxa currently included

• Machaerotidae• 6 of 31 genera

• Clastopteridae • 1 of 2 genera

• Aphrophoridae • 14 of 150 genera

• Epipygidae • 2 of ? genera

• Cercopidae • 33 of 142 genera

Page 21: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

Taxon Sampling:• Broad Diversity•

More to be added from unsorted material and newly   

collected specimens 

Page 22: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

Molecular Sampling:• Target for NSF project is 5 genes

• 18S rRNA

1,200 base pairs• 28S rRNA

1,900 base pairs 

• Cytochrome Oxidase I – 900 base pairs• Cytochrome Oxidase II – 630 base pairs• Histone 3 – 350 base pairs• Wingless – 300 base pairs

Page 23: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

New Gene Optimization:• NADH dehydrogenase subunit 4 –

1,338 base pairs

• New primers and PCR protocols• Optimized with Invitrogen

Platinum Taq• Highly informative gene in Mantodea

12S 16S COI COII ND4 TOTALS 18S 28S H3 Wingless TOTALS All Genes

Parsimony Gaps Missing Tree Length 6339 10928 32774 17504 32184 99729 1243 7395 2993 2350 13981 113710CI 0.104 0.084 0.054 0.065 0.068 0.526 0.285 0.092 0.156 0.089RI 0.476 0.467 0.427 0.444 0.418 0.753 0.717 0.66 0.705 0.483

Maximum Likelihood Tree Length 6379 11026 33001 17591 32388 100385 1225 7293 2943 2321 13782 114167CI 0.103 0.084 0.054 0.064 0.068 0.534 0.289 0.093 0.158 0.088RI 0.472 0.461 0.423 0.441 0.414 0.761 0.723 0.666 0.709 0.481

Bremer Value Sums 1844.9 2846.7 2233.6 -110.5 7074 13889 136.3 469.7 -1106 602.2 101.9 13989Percentage of Bremer Contribution by Gene 13% 20% 16% -1% 51% 99% 1% 3% -8% 4% 1% 1

Backbone nodes 53.3 83.9 -83.5 -78.5 290.8 266 -2.2 16.9 -87.2 24.4 -48.1 218percentage 24% 38% -38% -36% 133% 122% -1% 8% -40% 11% -22%

Major Clade Recovery Nodes 140.6 205.3 4.5 -152.4 282.4 480.4 89.8 3.7 -125.7 35.9 3.7 484percentage 29% 42% 1% -31% 58% 99% 19% 1% -26% 7% 1%

Minor Clade Recovery Nodes 110.4 140.6 151 -142.3 458.4 718.1 -8.1 14.6 -103.1 13.5 -83.1 635percentage 17% 22% 24% -22% 72% 113% -1% 2% -16% 2% -13%

Intra-Minor Clade Nodes 155.3 219.1 -49.4 -89.8 550.3 785.5 5.4 -1.2 -169.6 70.7 -94.7 691percentage 22% 32% -7% -13% 80% 114% 1% 0% -25% 10% -14%

Intra-Tribal Nodes 1385.3 2197.8 2211 352.5 5492.1 11639 51.4 435.7 -620.7 457.7 324.1 11961percentage 12% 18% 18% 3% 46% 97% 0% 4% -5% 4% 3%

NuclearMitochondrial

Page 24: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

New Gene Optimization:• Juvenile Hormone Binding Protein –

Targeted for its potential  as a more conserved gene 

to inform deep level  relationships. 

• PCR and cloning• degenerate primer design• RtPCR

may not be fruitful based on JH 

levels in adults, need nymphs, 

which may contain more JHBP to 

bind with JH•

could optimize in nymphs and 

then optimize primers for genomic 

gene amplification

Page 25: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

Current Molecular Coverage:• 18S rRNA

99 % of taxa

• 28S rRNA

99 % of taxa• Cytochrome Oxidase I – 87 % of taxa• Cytochrome Oxidase II – 54 % of taxa• Histone 3 – 95 % of taxa• Wingless – 93 % of taxa• ND4 – 55 % of taxa• JHBP – NA 

Page 26: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

The Preliminary Analysis:

• 117 Taxa to establish deep level relationships• Test familial relationships• Investigate deep level branching hypotheses

• Parsimony: TNT new technology searches• Maximum Likelihood

• GARLI (stochastic genetic algorithm‐like approach)• TREEFINDER (partitioned likelihood searches) 

• Mixed Model Bayesian Analysis (MrBayes)• Not finished

Page 27: Cryan Lab of Insect Phylogeneticshydrodictyon.eeb.uconn.edu/projects/cicada/simon... · • JHBP –NA. Cercopoidea. Molecular Sampling. The Preliminary Analysis: • 117 Taxa to

Cercopoidea

Molecular Sampling

The Preliminary Results:S t r i c t c o n s e n s u s o f 4 t r e e s ( 0 t a x a e x c l u d e d )

T R O 0 0 1

L H 0 3 8G f o

F U L 0 5 6

D i n

D E L 1 5 5

C I C 0 0 9C I C 0 0 4

C E R 2 7 9 _ M a _ G r y p _ t u r

C E R 2 7 7 _ M a _ A p o m _ r e t

C E R 2 7 4 _ A p _ P o o p _ t e r rC E R 2 7 3 _ A p _ P o o p _ c o s

C E R 2 6 9 _ A p _ C l o v _ Z a m b

C E R 2 6 8 _ C e _ L o c r _ r u b

C E R 2 6 6 _ C e _ A u f i _ M a l a y

C E R 2 6 5 _ M a _ P e c t _ h y a

C E R 2 6 4 _ C e _ M a c h _ i n v

C E R 2 6 3 _ C e _ L o c r i s _ p u l

C E R 2 5 8 _ C e _ B a e t _ m a r

C E R 2 5 0 _ C e _ L o c r i s _ h a l

C E R 2 4 5 _ A p _ P t y e _ G h a n a

C E R 2 4 3 _ C e _ L o c r i s _ m a cC E R 2 4 2 _ C e _ L o c r i s _ a c t

C E R 2 4 0 _ C e _ B a e t _ c o m

C E R 2 3 8 _ C e _ S p h e _ p a r

C E R 2 3 6 _ A p _ L i o r _ T a i w a n

C E R 2 3 4 _ A p _ P e u c _ T a i w a n

C E R 2 3 1 _ A p _ P l i n _ M a l a y

C E R 2 2 6 _ C e _ F l o s s _ T a i w a n

C E R 2 2 5 _ C e _ C o s m _ u c h

C E R 2 2 4 _ M a _ T a i h _ g e i

C E R 2 2 1 _ C e _ M o n e _ P e r uC E R 2 1 8 _ C e _ P a c h _ b e l

C E R 2 1 7 _ C e _ M a h a _ t r i

C E R 2 1 6 _ C e _ N o t o _ e n t

C E R 2 1 3 _ C e _ A u f i _ P N G

C E R 2 1 2 _ A p _ E p i c _ P e r u

C E R 2 0 7 _ C e _ I p h i _ q u o

C E R 2 0 6 _ C e _ O c o a _ t u c

C E R 2 0 4 _ A p _ E p i c _ C o s t a R i c a

C E R 2 0 1 _ C e _ I s o z _ s p

C E R 1 9 8 _ M a _ E n d e _ b i s

C E R 1 9 5 _ A p _ A p h r _ p e r

C E R 1 9 4 _ C e _ M e r i _ F r G u i a n a

C E R 1 9 3 _ E p _ E i c i _ F r G u i a n a

C E R 1 9 0 _ C e _ N o t o _ G h a n a

C E R 1 8 9 _ C e _ N o t o _ e n t

C E R 1 8 8 _ A p _ P h i l _ T a i w a n

C E R 1 8 3 _ C e _ S u r a _ t r i

C E R 1 8 2 _ C e _ A r a c _ d i m

C E R 1 8 0 _ C e _ L e p t _ f u l v

C E R 1 7 9 _ C e _ C o s m _ a b d

C E R 1 7 8 _ C e _ C o s m _ b i s

C E R 1 7 6 _ C e _ A u f i _ M a l a y

C E R 1 7 5 _ C e _ S p h e _ r u b

C E R 1 7 2 _ A p _ A p h r _ s a r

C E R 1 7 1 _ A p _ P h i l _ b i l

C E R 1 6 7 _ A p _ C l o v _ M a l a y

C E R 1 6 6 _ C e _ S p h e _ p r o

C E R 1 6 5 _ C e _ H o m a _ c o c

C E R 1 6 4 _ C e _ C o s m _ d i m

C E R 1 6 0 _ C e _ I s o z _ a s t

C E R 1 5 9 _ A p _ C l o v _ s p

C E R 1 5 6 _ M a _ M a c h _ t a k

C E R 1 5 2 _ C e _ L a c c _ p r a

C E R 1 5 1 _ C e _ M a x a _ q u a

C E R 1 4 9 _ A p _ E s c r _ s pC E R 1 4 7 _ A p _ S p h o _ g i g

C E R 1 4 5 _ C e _ L a c c _ g r a

C E R 1 4 3 _ C e _ C a t r _ s e m

C E R 1 4 2 _ C e _ M a x a _ d i v

C E R 1 3 8 _ C l _ C l a s _ l a e

C E R 1 3 6 _ C l _ C l a s _ a g r

C E R 1 3 5 _ C e _ P r o s _ p l a

C E R 1 3 1 _ C e _ C a t r _ i n s

C E R 1 2 9 _ M a _ P e c t _ s t a

C E R 1 2 8 _ M a _ M a c h _ p u gC E R 1 2 5 _ M a _ H i n d _ b i p

C E R 1 2 4 _ C e _ A u f i _ s p

C E R 1 2 3 _ M a _ P o l y _ s e rC E R 1 2 0 _ M a _ C h a e _ v i c

C E R 1 0 8 _ C e _ H u a i _ i n c

C E R 1 0 7 _ C e _ T o m a _ b o l

C E R 1 0 6 _ C e _ A e n e _ a l b

C E R 1 0 5 _ C e _ I p h i _ p e r

C E R 1 0 4 _ C e _ M i c r _ s a r

C E R 0 9 5 _ A p _ C e p h _ s i c

C E R 0 9 4 _ C e _ M a h a _ c o s

C E R 0 9 1 _ E p _ E v e x _ e m pC E R 0 9 0 _ E p _ E v e x _ r a k

C E R 0 8 9 _ E p _ E p i p _ a f a

C E R 0 8 7 _ C e _ O c o a _ o r n

C E R 0 7 0 _ C e _ S p h e _ l a t

C E R 0 6 9 _ C e _ A e n e _ v a r

C E R 0 6 7 _ C e _ P r o s _ s i m

C E R 0 6 6 _ C e _ S p h e _ d i s

C E R 0 6 4 C e A e n e c o n

C E R 0 6 2 _ C e _ A e n e _ r e d

C E R 0 6 0 _ C e _ Z u l i _ c a r

C E R 0 3 3 _ A p _ P h i l _ m a gC E R 0 2 3 _ A p _ P h i l _ t e s

C E R 0 2 2 _ C e _ C a l l _ s p

C E R 0 2 1 _ C l _ C l a s _ x a n

C E R 0 2 0 _ C l _ C l a s _ t e s

C E R 0 1 9 _ C e _ Z u l i _ v i l

C E R 0 1 6 _ C l _ C l a s _ b r u

C E R 0 1 5 _ C e _ P r o s _ b i c

C E R 0 0 9 _ A p _ L e p y _ q u a

C E R 0 0 8 _ A p _ A p h r _ q u a

C E R 0 0 7 _ A p _ A p h r _ c r i

C E R 0 0 6 _ C l _ C l a s _ o b t

C E R 0 0 5 _ A p _ A p h r _ a l n

C E R 0 0 4 _ A p _ L e p y _ c o l

C E R 0 0 3 _ C l _ C l a s _ p r o

C E R 0 0 2 _ A p _ N e o p _ l i n

C E R 0 0 1 _ A p _ P h i l _ s p u

A H 0 0 9 _ M a _ P e c t _ r e t

Parsimony: TNT new technology searches

0.2

CER021_Cl_Clas_xan

CER204_Ap_Epic_CostaRica

CER277_Ma_Apom_ret

CER234_Ap_Peuc_Taiwan

CER033_Ap_Phil_mag

CER218_Ce_Pach_bel

CER165_Ce_Homa_coc

CER183_Ce_Sura_tri

CER147_Ap_Spho_gig

CER175_Ce_Sphe_rub

CER004_Ap_Lepy_col

CER002_Ap_Neop_lin

CER190_Ce_Noto_Ghana

CER212_Ap_Epic_Peru

CER224_Ma_Taih_gei

CER008_Ap_Aphr_qua

CER268_Ce_Locr_rub

CER273_Ap_Poop_cos

CER142_Ce_Maxa_div

CER264_Ce_Mach_inv

CER062_Ce_Aene_red

CER245_Ap_Ptye_GhanaCER188_Ap_Phil_Taiwan

CER176_Ce_Aufi_Malay

CER136_Cl_Clas_agr

CER149_Ap_Escr_sp

CER180_Ce_Lept_fulv

CER194_Ce_Meri_FrGuiana

CER060_Ce_Zuli_car

CER123_Ma_Poly_ser

CER152_Ce_Lacc_pra

CER001_Ap_Phil_spu

CER172_Ap_Aphr_sar

CER206_Ce_Ocoa_tuc

CER242_Ce_Locris_act

CER263_Ce_Locris_pul

CER094_Ce_Maha_cos

CER015_Ce_Pros_bic

CER007_Ap_Aphr_cri

CER107_Ce_Toma_bol

CER087_Ce_Ocoa_orn

CER019_Ce_Zuli_vil

CER167_Ap_Clov_Malay

CER225_Ce_Cosm_uch

CER067_Ce_Pros_sim

CER009_Ap_Lepy_qua

CER131_Ce_Catr_ins

CER217_Ce_Maha_tri

TRO001

Din

CER105_Ce_Iphi_per

CER095_Ap_Ceph_sic

CER064_Ce_Aene_con

CER240_Ce_Baet_com

DEL155

CER090_Ep_Evex_rak

CIC004

CER258_Ce_Baet_mar

CER120_Ma_Chae_vic

CER016_Cl_Clas_bru

CER193_Ep_Eici_FrGuiana

CER178_Ce_Cosm_bis

LH038

CER195_Ap_Aphr_per

CER135_Ce_Pros_pla

CER124_Ce_Aufi_sp

CER106_Ce_Aene_alb

FUL056

CER171_Ap_Phil_bil

CER151_Ce_Maxa_qua

CER129_Ma_Pect_sta

CER023_Ap_Phil_tes

CER022_Ce_Call_spCER213_Ce_Aufi_PNG

CER238_Ce_Sphe_par

AH009_Ma_Pect_ret

CER108_Ce_Huai_inc

CER198_Ma_Ende_bis

CER069_Ce_Aene_var

CER005_Ap_Aphr_aln

CER182_Ce_Arac_dim

CER231_Ap_Plin_Malay

CER266_Ce_Aufi_Malay

CER091_Ep_Evex_emp

CER070_Ce_Sphe_lat

CER274_Ap_Poop_terr

CIC009

CER003_Cl_Clas_pro

CER201_Ce_Isoz_sp

CER156_Ma_Mach_tak

CER145_Ce_Lacc_gra

CER179_Ce_Cosm_abd

CER160_Ce_Isoz_ast

Gfo

CER104_Ce_Micr_sar

CER236_Ap_Lior_Taiwan

CER066_Ce_Sphe_dis

CER128_Ma_Mach_pug

CER006_Cl_Clas_obt

CER020_Cl_Clas_tes

CER250_Ce_Locris_hal

CER159_Ap_Clov_sp

CER164_Ce_Cosm_dim

CER207_Ce_Iphi_quo

CER189_Ce_Noto_ent

CER143_Ce_Catr_sem

CER166_Ce_Sphe_pro

CER269_Ap_Clov_Zamb

CER125_Ma_Hind_bip

CER279_Ma_Gryp_tur

CER138_Cl_Clas_lae

CER226_Ce_Floss_Taiwan

CER243_Ce_Locris_mac

CER265_Ma_Pect_hya

CER221_Ce_Mone_Peru

CER089_Ep_Epip_afa

CER216_Ce_Noto_ent

Maximum Likelihood: GARLI

Machaerotidae

Clastopteridae

Aphrophoridae

Epipygidae

Old WorldCercopidae

New WorldCercopidae

Epipygidae

Machaerotidae

Clastopteridae

Aphrophoridae

Old WorldCercopidae

New WorldCercopidae