吸汁性昆虫キジラミ-細菌間の多様な複合共生系
中鉢 淳1, 井上広光2, 広瀬 侑1 (1豊橋技科大, 2農業・食品産業技術総合研究機構)
私どものEIIRISプロジェクト研究では、昆虫共生細菌からの新規二次代謝産物の獲得と、その応用利用を目指した機能解析を進めている。これまでに、1) 他に類例のないオルガネラ様防衛共生体「Candidatus Profftella armatura （Gammaproteobacteria: Burkholderiales）」が重要害虫「ミカンキジラミ（Diaphorina citri, 半翅目: キジラミ上科）」やその近縁種に存在し、新規ポリケチド「ディアフォリン」を産生していること、2) ディアフォリンが抗腫瘍活性をはじめとする興味深い生物活性を持つこと、などを明らかにしている。
今回は、さらに新たな共生細菌を見出すべく、次世代シークエンサーを用いてキジラミ科に属する日本産キジラミ12種の共生細菌叢を解析した。
[BMC Microbiology 2022, 22(1):15]

各キジラミ種の雌雄各5頭より抽出したDNAを鋳型として
16S rRNA遺伝子のV3-V4領域をPCR増幅し、MiSeqによる塩基配列決定後、QIIME2で群集解析を行い、RAxMLを用いた最尤法により分子系統解析。

98 SV5 Anomoneura mori [Psyllidae: Psyllinae]
Anomoneura mori AEM98 [Psyllidae: Psyllinae] (AB013086)

0.1

Cacopsylla pyri AEM00 [Psyllidae: Psyllinae] (AF211131)
Cacopsylla myrthi AEM00 [Psyllidae: Psyllinae] (AF211129)
SV3 Cacopsylla burckhardti [Psyllidae: Psyllinae]
SV23 Cyamophila hexastigma [Psyllidae: Psyllinae]
SV15 Cacopsylla fatsiae [Psyllidae: Psyllinae]
SV6 Cacopsylla kiushuensis [Psyllidae: Psyllinae]

100

SV17 Cacopsylla satsumensis [Psyllidae: Psyllinae]
Cacopsylla pyricola IMB01 [Psyllidae: Psyllinae] (AF286118)
Cacopsylla brunneipennis AEM00 [Psyllidae: Psyllinae] (AF243138)
SV10 Cacopsylla jukyungi [Psyllidae: Psyllinae]
SV19 Cacopsylla peninsularis [Psyllidae: Psyllinae]
71
SV29 Cacopsylla peninsularis [Psyllidae: Psyllinae]
Cacopsylla peregrina AEM00 [Psyllidae: Psyllinae] (AF211130)

キジラミの種ごとに多様で新規な共生細菌叢を解明。とくに、- これまでアブラムシ（半翅目: アブラムシ上科）の共生体としてのみ認識されていたCa. Fukatsuia symbioticaとSerratia symbiotica（Gammaproteobacteria: Enterobacteriales）を、世界で初めてキジラミから検出した。

Relative abundance (%)

75

SV32 Psylla morimotoi [Psyllidae: Psyllinae]

Carsonella

97 SV42 Psylla morimotoi [Psyllidae: Psyllinae]

Pseudomonas

SV13 Cacopsylla coccinea [Psyllidae: Psyllinae]

Endosymbionts2

SV24 Cacopsylla biwa [Psyllidae: Psyllinae]

Heteropsylla texana MBE12 [Psyllidae: Ciriacreminae] (CP003544)

Arsenophonus

Spanioneura fonscolombii AEM00 [Psyllidae: Psyllinae] (AF211149)(39)

SodalisPsylla buxi AEM00 [Psyllidae: Psyllinae] (AF211146)
Arytaina genistae AEM00 [Psyllidae: Psyllinae] (AF243136)

Serratia

50

Psylla floccosa IMB01 [Psyllidae: Psyllinae] (AF286117)

Fukatsuia

SV18 Epiacizzia kuwayamai [Psyllidae: Macrocorsinae](47)

Ambiguous Enterobacteriaceae

Panisopelma sp. AEM00 [Psyllidae: Aphalaroidinae] (AF211145)

92
Panisopelma fulvescens AEM00 [Psyllidae: Aphalaroidinae] (AF211144)
53
Russelliana intermedia AEM00 [Psyllidae: Aphalaroidinae] (AF211148)

Diplorickettsia
Liberibacter

Acizzia uncatoides AEM00 [Psyllidae: Acizziinae] (AF211123)
Diaphorina cf. continua [Liviidae: Euphyllurinae] (TAAA01000002)
54
Diaphorina citri [Liviidae: Euphyllurinae] (CP003467)
99
Diaphorina citri Florida [Liviidae: Euphyllurinae] (EF450250)

Wolbachia

25- 世界の柑橘産業に致命的な打撃を与えているCa. Liberibacter
asiaticus （Alphaproteobacteria: Rhizobiales）と同属の潜在的植物病原体Ca. Liberibacter europaeusを、クワの害虫であるクワ
キジラミ(Anomoneura mori)から、世界で初めて検出した。

Rickettsia

97

othersDiaphorina lycii [Liviidae: Euphyllurinae] (TAAA01000003)
68
Diaphorina lycii Thao unpublished [Liviidae: Euphyllurinae] (AF280097)
Neotriozella hirsuta AEM00 [Triozidae] (AF211140)
Cecidotrioza sozanica IMB01 [Triozidae] (AF286120)
Trioza eugeniae AEM00 [Triozidae] (AF211151)

56

a

am

ai

oi
ot

ay

図2. Carsonellaの最尤系統樹。
太字: 本解析で得られた配列。

Trioza urticae AEM00 [Triozidae] (AF211152)w

Calophya schini AEM00 [Calophyidae: Calophyinae] (AF211132)

ku

sis

m
ig

zia

Aphalaroida inermis AEM00 [Psyllidae: Aphalaroidinae] (AF211125)

ciz

en

st
xa

m
or
im

he

a

a

yll

Glycaspis brimblecombei AEM00 [Aphalaridae: Spondyliaspidinae] (AF211137)

ia

hil

Ps

67

Ep

is

ris

ns

la
su

um

nin

ts

pe

sa
lla

sy

op
m

op
ac

ya

lla

lla
sy
op

ac
C

Trioza magnoliae MBE98 [Triozidae] (AF077604)

Bactericera cockerelli AEM00 [Triozidae] (AF211126)
70
Bactericera trigonica Katsir unpublished [Triozidae] (CP024798)

Blastopsylla occidentalis MBE98 [Aphalaridae: Spondyliaspidina] (AF077605)

C

e

gi
un

ue

ky

sh

kiu

ju

lla

lla

sy

sy

op

op
ac

C

sy

ac

ac

C

C

ea

ia
ts
fa

cc

lla
C

op

ti

in

rd

biw

ha

lla

rc
k

sy

bu

sy

lla

op

sy

ac

op

C

ra
eu

op

on

ac

om

C
C

ac

co

m
or
i

a

0

An

その他、様々な共生細菌を検出。
今後解析を進めることで、新規有用物質の獲得を目指す。

Heteropsylla cubana MBE12 [Psyllidae: Ciriacreminae] (CP003543)

82

Endosymbionts3

Cardiaspina maniformis Microbiome17 [Aphalaridae: Spondyliaspidinae] (KY427942)
Boreioglycaspis melaleucae AEM00 [Aphalaridae: Spondyliaspidinae] (AF211128)

58

Ctenarytaina eucalypti MBE12 [Aphalaridae: Spondyliaspidinae] (CP003541)

図1. キジラミ科12種の体内共生細菌叢。
縦軸: 各分類群に対応する配列の相対頻度。
0.2

Arsenophonus nasoniae [Nasonia vitripennis (wasp)] (CP038613)
Ca. Arsenophonus triatominarum [Triatoma rubrofasciata (assassin bug)] (DQ508185)
100 Arsenophonus endosymbiont [Cardiaspina tenuitela (Aphalaridae: Spondyliaspidinae)] (KY428657)=Secondary endosymbiont [Glycaspis brimblecombei (Aphalaridae: Spondyliaspidinae)] (EU043378)
99
SV27 Arsenophonus endosymbiont [Cacopsylla jukyungi (Psyllidae: Psyllinae)]
58
Morganella morganii (LT899955)
Salmonella enterica (MH155974)
Ca. Fukatsuia symbiotica [Maculolachnus submacula (aphid)] (FJ655539)
100 Ca. Fukatsuia symbiotica [Cinara confinis (aphid)] (LT600381)
53
68 100 SV31 Ca. Fukatsuia symbiotica [Psylla morimotoi (Psyllidae: Psyllinae)]= Ca. Fukatsuia symbiotica [Acyrthosiphon pisum (aphid)] (CP021659)
Ca. Hamiltonella defensa [Acyrthosiphon pisum (aphid)] (CP001277)
53
Ca. Regiella insecticola [Acyrthosiphon pisum (aphid)] (AY296734)
59
Serratia fonticola (CP054160)
Serratia plymuthica (CP015613)
90
SV37 Serratia symbiotica [Cacopsylla coccinea (Psyllidae: Psyllinae)]
95 SV38 Serratia symbiotica [Cacopsylla coccinea (Psyllidae: Psyllinae)]
SV35 Serratia symbiotica [Cacopsylla coccinea (Psyllidae: Psyllinae)]
97 90
SV39 Serratia symbiotica [Cacopsylla coccinea (Psyllidae: Psyllinae)]
SV40 Serratia symbiotica [Cacopsylla coccinea (Psyllidae: Psyllinae)]
Serratia symbiotica [Acyrthosiphon pisum (aphid)] (AB522706)
SV41 Serratia symbiotica [Cacopsylla coccinea (Psyllidae: Psyllinae)]
96
Escherichia coli (CP032667)
Pantoea ananatis (KT741004)
Pantoea agglomerans (AF130962)
90
Erwinia toletana (NR_104828)
Sodalis endosymbiont [Elymana sulphurella (leafhopper)] (MG835279)
Sodalis endosymbiont [Porphyrophora polonica (scale insect)] (MT232255)
54
Sodalis endosymbiont [Spalangia cameroni (wasp)] (KY985293)
73
Sodalis praecaptivus (CP006569)
Secondary symbiont [Coelostomidia pilosa (scale insect)] (KC447409)
SV14 Sodalis endosymbiont [Cacopsylla burckhardti (Psyllidae: Psyllinae)]
Ca. Sodalis pierantonius [Sitophilus oryzae (weevil)] (AF548137)
Sodalis endosymbiont [Microlynchia galapagoensis (louse fly)] (MF429884)
Sodalis glossinidius [Glossina morsitans (tsetse fly)] (AP008232)
98
Sodalis glossinidius [Glossina brevipalpis (tsetse fly)] (U64870)
Sodalis endosymbiont [Philaenus tesselatus (spittlebug)] (AB772197)
72
SV28 Sodalis endosymbiont [Cacopsylla kiushuensis (Psyllidae: Psyllinae)]
75
Secondary endosymbiont [Ctenarytaina eucalypti (Aphalaridae: Spondyliaspidinae)] (CP003546)
Sodalis endosymbiont [Cardiaspina maniformis (Aphalaridae: Spondyliaspidinae)] (KY428659)
SV34 Ambiguous Enterobacteriaceae symbiont [Psylla morimotoi (Psyllidae: Psyllinae)]
Secondary endosymbiont [Heteropsylla cubana (Psyllidae: Ciriacreminae)] (CP003547)
Ca. Moranella endobia [Planococcus citri (scale insect)] (CP002243)
100 Wigglesworthia glossinidia [Glossina brevipalpis (tsetse fly)] (BA000021)
Wigglesworthia glossinidia [Glossina morsitans (tsetse fly)] (L37339)
Ca. Schneideria nysicola [Orsillus depressus (seed bug)] (HE586114)
Ca. Kotejella greeniscae [Greenisca brachypodii (scale insect)] (KY558666)
Ca. Baumannia cicadellinicola [Homalodisca coagulata (leafhopper)] (CP000238)
99
Ca. Baumannia cicadellinicola [Graphocephala aurora (leafhopper)] (AF465797)
Ca. Blochmannia pennsylvanicus [Camponotus pennsylvanicus (ant)] (AY196850)
Ca. Blochmannia festinatus [Camponotus festinatus (ant)] (AY196851)
88
91 Ca. Blochmannia vafer [Camponotus vafer (ant)] (CP002189)
Secondary endosymbiont [Mycopsylla fici (Homotomidae)] (KT273275)
Secondary endosymbiont [Calophya schini (Calophyidae: Calophyinae)] (AF263560)
Endosymbiont [Protyora sterculiae (Carsidaridae)] (KY428660)
Primary endosymbiont [Pedicinus badii (louse)] (EU827263)
100 Ca. Puchtella pedicinophila [Pedicinus obtusus (louse)] (AB478979)
Secondary endosymbiont [Trioza magnoliae (Triozidae)] (AF077607)
Buchnera aphidicola [Acyrthosiphon pisum (aphid)] (BA000003)
99
Buchnera aphidicola [Schizaphis graminum (aphid)] (AE013218)
54 Buchnera aphidicola [Myzus persicae (aphid)] (CP002703)
Ca. Annandia adelgestsuga [Adelges tsugae (adelgid)] (KC331979)
Ca. Ishikawaella capsulata [Coptosoma parvipictum (stink bug)] (AB240161)
Ca. Rosenkranzia clausaccus [Elasmucha putoni (stink bug)] (AB368828)
Ca. Tachikawaea gelatinosa [Urostylis westwoodii (stink bug)] (AP014521)
69 Endosymbiont [Donacia porosicollis (leaf beetle)] (GQ480879)
61 88
Endosymbiont [Placobdelloides siamensis (leech)] (AB083058)
SV9 “Endosymbiots3” Enterobacteriaceae endosymbiont [Anomoneura mori (Psyllidae: Psyllinae)] = Y-symbiont [A. mori ] (AB013087)
SV2 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cyamophila hexastigma (Psyllidae: Psyllinae)]
Secondary endosymbiont [Cacopsylla myrthi (Psyllidae: Psyllinae)] (AF263559)“Arsenophonus” endosymbiont [Trichobius caecus (bat fly)] (DQ314768)
Secondary endosymbiont [Cacopsylla pyricola (Psyllidae: Psyllinae)] (AF286125)“Arsenophonus” endosymbiont [Cacopsylla pyricola (Psyllidae: Psyllinae) (KX077196)
SV25 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cacopsylla kiushuensis (Psyllidae: Psyllinae)]
57 SV16 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cacopsylla jukyungi (Psyllidae: Psyllinae)]
SV22 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cacopsylla fatsiae (Psyllidae: Psyllinae)]
100
SV11 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cacopsylla fatsiae (Psyllidae: Psyllinae)]
62
SV4 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cacopsylla coccinea (Psyllidae: Psyllinae)]
SV26 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cacopsylla peninsularis (Psyllidae: Psyllinae)]
SV1 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cacopsylla biwa (Psyllidae: Psyllinae)]
SV8 “Endosymbionts3” Enterobacteriaceae endosymbiont [Cacopsylla satsumensis (Psyllidae: Psyllinae)]
SV21 “Endosymbionts2” Enterobacteriaceae endosymbiont [Epiacizzia kuwayamai (Psyllidae: Macrocorsinae)]

図3. 腸内細菌科共生細菌の最尤系統樹。
赤字: キジラミの共生細菌, 青字: それ以外の動物の共生細菌。
世界で初めてキジラミからCa. Fukatsuia symbioticaとSerratia symbioticaを検出。

Ctenarytaina longicauda AEM00 [Aphalaridae: Spondyliaspidinae] (AF211134)
Tainarys sordida AEM00 [Aphalaridae: Rhinocolinae] (AF211150)
100 Mycopsylla fici Fromont unpublished [Homotomidae: Macrohomotominae] (KT273280)
Mycopsylla proxima Fromont unpublished [Homotomidae: Macrohomotominae] (KT273279)
Aphalara longicaudata AEM00 [Aphalaridae: Aphalarinae] (AF243137)
Protyora sterculiae Microbiome17 [Carsidaridae] (KY427941)
100

Pachypsylla pallida AEM00 [Aphalaridae: Pachypsyllinae] (AF211142)

80

Pachypsylla celtidismamma IMB01 [Aphalaridae: Pachypsyllinae] (AF286122)

89

Pachypsylla venusta Science06 [Aphalaridae: Pachypsyllinae] (AP009180)
Ca. Portiera aleyrodidarum [Bemisia tabaci (whitefly)] (AF400453)
Gammaproteobacterium symbiont [Metapocyrtus yonagunianus (weevil)] (LC014980)

SV12 Pseudomonas symbiont [Psylla morimotoi (Psyllidae: Psyllinae)]

0.2

0.5

95

SV36 Diplorickettsia sp. [Psylla morimotoi (Psyllidae: Psyllinae)]

Pseudomonas rhizosphaerae DSM 16299 (CP009533)
65

Diplorickettsia massiliensis 20B [Ixodes ricinus (tick)] (NR_117407)

Pseudomonas graminis DSM 11363 (Y11150)

50

Pseudomonas lutea OK2 (AY364537)

62

62
Diplorickettsia sp. NS15 (JN606082)

SV30 Pseudomonas symbiont [Psylla morimotoi (Psyllidae: Psyllinae)]
85

Diplorickettsia sp. [Diaphorina cf. continua (Liviidae: Euphyllurinae)] (TAAA01000010)

Pseudomonas viridiflava ATCC 13223 (NR_114482)

Pseudomonas syringae ATCC 19310 (AF094749)

Diplorickettsia sp. MSebKT1 [Macrosteles sexnotatus (leafhopper)] (AB795342)

Ca. Pseudomonas adelgestsugas [Adelges tsugae (Hemiptera: Adelgidae)](CP026512)
90

Rickettsiella endosymbiont Ixotasmani1 [Ixodes tasmani (tick)] (KP994858)

Pseudomonas fluorescens ATCC 13525 (AF094725)
65

66

Pseudomonas putida ATCC 12633 (AF094736)

Rickettsiella grylli [Gryllus bimaculatus (cricket)] (U97547)

67
Pseudomonas protegens CHA0T (AJ278812)
Ca. Rickettsiella viridis Ap-RA04 [Acyrthosiphon pisum (aphid)] (AP018005)

55
Pseudomonas entomophila L48 (AY907566)

62

SV5 Ca. Carsonella ruddii [Anomoneura mori (Psyllidae: Psyllinae)]

Ctenarytaina spatulata MBE12 [Aphalaridae: Spondyliaspidinae] (CP003542)

76

52

77
Rickettsiella symbiont [Asellus aquaticus (waterlouse)] (AY44704)

Pseudomonas stutzeri ATCC 17588 (AF094748)
97
Pseudomonas resinovorans ATCC 14235 (AB021373)

Rickettsiella melolonthae BBA1806/LAM6-D/2004 [Melolontha spp. (beetle)] (EF408231)
Pseudomonas aeruginosa DSM 50071 (CP012001)
90

Rickettsiella agriotidis JKI_E1959/09D [Agriotes sp. (beetle)] (HQ640943)

Pseudomonas endosymbiont [Paederus fuscipes (Copleoptera: Staphylinidae)](KJ081228)
Pseudomonas alcaligenes ATCC 14909 (AF094721)

Coxiella burnetii Heizberg [Ixodid tick] (CP014561)

SV5 Ca. Carsonella ruddii [Anomoneura mori (Psyllidae: Psyllinae)]
SV5 Ca. Carsonella ruddii [Anomoneura mori (Psyllidae: Psyllinae)]

図4. Pseudomonasの最尤系統樹。
赤字: キジラミの共生細菌, 青字: それ以外の動物の共生細菌。

図5. Diplorickettsiaの最尤系統樹。
赤字: キジラミの共生細菌, 青字: それ以外の動物の共生細菌。

Rickettsia endosymbiont [Cerobasis guestfalica (booklouse)](DQ652596)

0.02
0.05

Wolbachia <A> [Drosophila melanogaster (fly)] (EU096232)

SV20 Rickettsia endosymbiont[Cacopsylla peninsularis (Psyllidae: Psyllinae)]

84

Wolbachia <A> [Drosophila simulans (fly)] (CP001391)
SV43 Wolbachia [Psylla morimotoi (Psyllidae: Psyllinae)]= Wolbachia <B> [Diaphorina citri/lycii (Liviidae: Euphyllurinae)] (TAAA01000005)

Rickettsia endosymbiont [Cacopsylla melanoneura (Psyllidae: Psyllinae)](LR800105)

72

Wolbachia <B> [Diaphorina cf. continua (Liviidae: Euphyllurinae)] (TAAA01000008)

Rickettsia endosymbiont [Chamaepsylla hartigii (Psyllidae: Psyllinae)](LR800074)

57

Wolbachia <B> [Bactericera cockerelli (Triozidae)] (EF372596)

54

Wolbachia <B> [Drosophila simulans (fly)] (CP003883)

SV33 Ca. Liberibacter europaeus [Anomoneura mori (Psyllidae: Psyllinae)]

0.1
96

Ca. Rickettsia torix [Bemisia tabaci (whitefly)] (MG063879)

Wolbachia <B> [Tetranychus urticae (mite)] (EU499319)
87

Ca. Liberibacter europaeus (TAAA01000007) [Diaphorina cf. continua (Liviidae: Euphyllurinae)]= Ca. Liberibacter europaeus NR-01 (FN678792) [Cacopsylla pyri (Psyllidae: Psyllinae)]

99

79

Rickettsia symbiont [Nephotettix cincticeps (planthopper)] (KU586121)
Wolbachia <B> [Diaphorina lycii (Liviidae: Euphyllurinae)] (TAAA01000011)

87

Wolbachia <B> [Nasonia vitripennis (parasitoid wasp)] (M84686)

Ca. Liberibacter europaeus Psy6/BrS (JX244258/JX244259) [Arytainilla spartiophila (Psyllidae: Psyllinae)]

96

97

Mycetome symbiont [Stegobium paniceum (beetle)] (JQ805029)

Wolbachia <B> [Diaphorina citri (Liviidae: Euphyllurinae)] (EF433793)

82

SV7 Wolbachia [Cacopsylla biwa, Cyamophila hexastigma (Psyllidae: Psyllinae),
Epiacizzia kuwayamai (Psyllidae: Macrocorsinae)]= Wolbachia <B> [Diaphorina citri (Liviidae: Euphyllurinae)] (TAAA01000013)

Ca. Liberibacter americanus Sao Paulo (NC_022793) [Diaphorina citri (Liviidae: Euphyllurinae)]

99

Rickettsia endosymbiont [Deronectes platynotus (beetle)] (FM177875)
98

Wolbachia <B> [Armadillidium vulgare (woodlouse)] (AJ133196)

67

Ca. Liberibacter solanacearum NZ083338 (EU935004) [Bactericera cockerelli (Triozidae)]

94

Rickettsia endosymbiont [Chrysotropia ciliata (lacewing)] (MF156626)

Wolbachia <C> [Dirofilaria immitis (nematode)] (Z49261)

Ca. Rickettsia mendelii [Ixodes ricinus (tick)] (KJ882317)
Wolbachia <C> [Onchocerca volvulus (nematode)] (HG810405)

63
Wolbachia <J> [Dipetalonema gracile (nematode)] (AJ548802)

Rickettsia endosymbiont [Liophloeus gibbus (weevil)] (MN621120)

67

Ca. Liberibacter caribbeanus (KP012551) [Diaphorina citri (Liviidae: Euphyllurinae)]
Wolbachia <D> [Brugia malayi (nematode)] (NR_074571)

Rickettsia rickettsii Iowa [Ixodid tick] (CP000766)

99

74

Wolbachia <D> [Wuchereria bancrofti (nematode)] (AF093510)

Ca. Liberibacter asiaticus_Ishi-1 (AP014595) [Diaphorina citri (Liviidae: Euphyllurinae)]

Wolbachia <F> [Nasutitermes nigriceps (termite)] (DQ837204)

88
79

Ca. Liberibacter africanus PTSAPSY (NZ_CP004021) [Trioza erytreae (Triozidae)]

Rickettsia japonica YH [Haemaphysalis hystricis (tick)] (NR_074459)

Wolbachia <F> [Coptotermes acinaciformis (termite)] (DQ837197)

Rickettsia typhi Wilmington [Xenopsylla cheopis (flea)] (NR_036948)
Wolbachia <O> [Bemisia tabaci (whitefly)] (KF454771)

80

Rickettsia prowazekii Brein1 [Pediculus humanus (louse)] (NR_044656)

Wolbachia <K> [Bryobia sp. (mite)] (EU499316)

Ca. Liberibacter brunswickensis Asol15 (KY077741) [Acizzia solanicola (Psyllidae: Acizziinae)]

Wolbachia <I> [Orchopeas leucopus (flea)] (AY335924)

Liberibacter crescens BT-1 (NC_019907)

Orientia chuto Dubai [Trombiculid mite] (NR_117903)

99

Wolbachia <I> [Ctenocephalides felis (flea)] (AY335923)
Wolbachia <L> [Radopholus similis (nematode)] (EU833482)

Wolbachia pipientis A (EU096232) [Drosophila melanogaster]

69
95

Orientia tsutsugamushi Karp [Leptotrombidium pallidum (mite)] (NR_025860)

Wolbachia <E> [Folsomia candida (springtail)] (EU831094)
94

Occidentia massiliensis Os18 [Ornithodoros sonrai (tick)] (NR_149220)

Wolbachia <E> [Mesaphorura italica (springtail)] (AJ575104)
Ca. Liberibacter asiaticus Ishi-1 (AP014595)

図6. Liberibacterの最尤系統樹。
太字: 世界で初めてクワキジラミ(Anomoneura mori)からLiberibacterを検出。

図7. Wolbachiaの最尤系統樹。
赤字: キジラミの共生細菌, 青字: それ以外の動物の共生細菌。

Wolbachia pipientis A [Drosophila melanogaster (fly)] (EU096232)

図8. Rickettsiaの最尤系統樹。
赤字: キジラミの共生細菌, 青字: それ以外の動物の共生細菌。