Cover Page The handle http://hdl.handle.net/1887/33207 holds various files of this Leiden University dissertation.

The handle http://hdl.handle.net/1887/33207 holds various files of this Leiden University dissertation.

Author: Meij, Sancia Esmeralda Theonilla van der

Title: Evolutionary diversification of coral-dwelling gall crabs (Cryptochiridae) Issue Date: 2015-06-03

Host relations and DNA reveal a cryptic gall crab species (Crustacea: Decapoda: Cryptochiridae) associated with mushroom corals (Scleractinia: Fungiidae)

Sancia E.T. van der Meij

Abstract

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VSHFLHVFORVHO\UHODWHGWRFungicola fagei7KLVQHZVSHFLHVGHVFULEHGKHUHDIWHUDVFungicola syzygia VSQRYLV

SUHGRPLQDQWO\DVVRFLDWHGZLWKWKHPXVKURRPFRUDOJHQHUDCycloseris and PleuractisZKHUHDVLWVVLEOLQJVSHFLHV

F. fagei LVRQO\NQRZQWREHDVVRFLDWHGZLWKWKHKRVWJHQHUDPodabacia and Sandalolitha%DVHGRQPRUSKRORJ\

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Introduction

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Material and methods

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Results

Molecular analyses

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F. fagei F. syzygia VSQRY F. utinomi

Fungicola fagei Q    

F. syzygia VSQRYQ    

F. utinomi Q    

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Fungicola fagei Q   

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54222 on Lithophyllon repanda

54237 on Podabacia crustacea 54238 on Podabacia crustacea 54239 on Pleuractis granulosa 53231 on Lithophyllon repanda

53711 on Lithophyllon repanda

54246 on Halomitra pileus

54017 on Fungia fungites

53233 on Sandalolitha robusta 54056 on Lithophyllon repanda

54236 on Cycloseris somervillei

53220 on Pleuractis granulosa

54228 on Podabacia crustacea 53688 on Lithophyllon scabra

56110 on Pleuractis paumotensis 54044 on Pleuractis paumotensis

54183 on Podabacia motuporensis 53773 on Lithophyllon repanda

54230 on Cycloseris tenuis

54218 on Cycloseris costulata

54240 on Pleuractis moluccensis

53232 on Podabacia crustacea 54016 on Pleuractis paumotensis

53227 on Pleuractis paumotensis

54216 on Cycloseris sinensis 53230 on Lithophyllon repanda

54235 on Cycloseris costulata

54020 on Pleuractis gravis 53237 Pseudocryptochirus viridis

54245 on Sandalolitha robusta 54006 Utinomiella dimorpha

54234 on Lithophyllon ranjithi

54220 on Cycloseris sinensis

54243 on Podabacia crustacea 54217 on Cycloseris fragilis

54244 on Lithophyllon repanda 53774 on Lithophyllon repanda

53712 on Lithophyllon repanda

54219 on Cycloseris costulata

54231 on Podabacia crustacea 53224 on Pleuractis granulosa 53229 on Lithophyllon repanda

53217 on Fungia fungites

53234 on Sandalolitha robusta 54232 on Podabacia crustacea 54233 on Lithophyllon ranjithi 54224 on Danafungia horrida

54226 on Lithophyllon repanda

54178 on Podabacia motuporensis 54221 on Pleuractis paumotensis

1

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outgroups

F. syzygia sp. nov.

F. fagei F. utinomi

0.03



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Host specificity

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repanda.

Systematic account

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Type species. Troglocarcinus utinomi )L]HDQG6HUqQH

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Remarks.)L]HDQG6HUqQHGHVFULEHGWKHVXEJHQXVFungicola LQWKHJHQXVTroglocar- cinus EDVHG RQ WKH IROORZLQJ FKDUDFWHULVWLFV  ÁDW FDUDSDFH RQ WKH GRUVRYHQWUDO VLGH  ÁDW

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Fungicola utinomi )L]HDQG6HUqQH )LJV&,$+

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RYLJIRQFungia > Lithophyllon@repanda01+1,8ZDV(JRRGFRQ- GLWLRQPRQFungia > Lithophyllon@repanda. Natural History Museum (LondonUK):

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Fig. 4. Fungicola fagei 501+&UXV'AKDELWXVGRUVDOYLHZBDQWHURODWHUDOPDUJLQRIFDUDSDFH

YHQWUDOYLHZC0;3DULJKW3FKHOLSHGEOHIW3FKHOLSHGFOHIW3GOHIW3HOHIW3I left 36FDOHEDUVPP%',VKDUHDVFDOHEDU

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ZLWKVSLQHVDQGIRUPLQJDFRQYH[OREHODWHUDOERUGHUVEHKLQGLWVWUDLJKWRUVOLJKWO\FRQYHUJLQJ

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Host corals. Fungicola utinomi LQKDELWVDWOHDVWQLQHFRUDOVSHFLHVEHORQJLQJWRHLJKWJHQHUD

EXWLVPRVWRIWHQUHFRUGHGIURPLithophyllon repanda )LJ7DEOH6,QL. repandaF. utinomi OLYHVLQDQRYDOFDYLW\ZLWKRYHUKDQJLQJFDQRS\ZKLFKFDQEHUDLVHGDERYHWKHFRUDOVXUIDFHVHH

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Distribution range. Fungicola utinomi KDVVRIDUEHHQGRFXPHQWHGIURP1KD7UDQJ9LHW- QDP)L]HDQG6HUqQH0DOXNX,QGRQHVLD5XPSKLXV,H[SHGLWLRQ6HUqQHet al

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Type locality.1KD7UDQJ9LHWQDP

Holotype.(¶7\SH·DFFRUGLQJWR)L]HDQG6HUqQHLQWKH,QVWLWXWHRI2FHDQRJUDSK\

LQ 1KD 7UDQJ 9LHWQDP )LJ $% DOORW\SH ( FROOHFWHG IURP Sandalolitha dentata 4XHOFKEXWVHHVHFWLRQRQKRVWFRUDOV

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QRODEHORQMDUJRRGFRQGLWLRQholotypeRYLJI(5WHLQMDU(UHVLGXH

( 5WH  LQ MDU ( 5WH  HPSW\ YLDO ( 5WH  LQ MDU

(5WHHPSW\YLDOallotype(5WHLQMDU(SRRUFRQ- GLWLRQ(LQMDU(5WHHPSW\YLDORQParahalomitra > San- dalolitha@robusta(5WHLQMDU(5WHHPSW\YLDORQPara- halomitra > Sandalolitha@robustaMDU(5WHFRQWDLQVRQHODEHOZLWKRXW

YLDO ( 5WH  Muséum national d’Histoire naturelle Paris France) 01+1

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Parahalomitra > Sandalolitha@robusta01+1,8ZDV(5WHJRRG

FRQGLWLRQRYLJIRQParahalomitra > Sandalolitha@robusta. Naturalis Biodiversity Center LeidenThe Netherlands)VHH7DEOH6

Diagnosis. &DUDSDFH VXEUHFWDQJXODU ORQJHU WKDQ EURDG GHSUHVVHG DQWHULRU KDOI EURDGHU WKDQ

SRVWHULRUKDOIVXUIDFHFRYHUHGZLWKÀQHJUDQXOHVPHGLDQJDVWULFUHJLRQIHHEO\FRQYH[LWVSRVWH- ULRUSDUWQRWVHSDUDWHGIURPFDUGLRLQWHVWLQDOUHJLRQEUDQFKLDOUHJLRQVVHSDUDWHGIURPPLGJDV- WULFDQGFDUGLRLQWHVWLQDOUHJLRQVE\VKDOORZIXUURZVIURQWPRGHUDWHO\FRQFDYHLQWHUQDORUELWDO

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Fig. 5. +RORW\SHFungicola syzygia VSQRY501+&UXV'AKDELWXVGRUVDOYLHZB anterolateral PDUJLQRIFDUDSDFHYHQWUDOYLHZC0;3DOHIW3FKHOLSHGEOHIW3FOHIW3GOHIW3H left 36FDOHEDUVPP%'+VKDUHVFDOHEDU

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FRQYH[ODWHUDOERUGHUVEHKLQGLWVOLJKWO\FRQFDYH&KHOLSHGQRWVWRXWPHUXVDVZHOODVXSSHUERU- GHUVRIFDUSXVSDOPFRYHUHGZLWKÀQHJUDQXOHV8SSHUIDFHRIFR[DHRIWKLUGWRÀIWKSHUHLRSRGV

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VHH)L]HDQG6HUqQH3O;,,,')$FFRUGLQJWR)L]HDQG6HUqQHFungicola fagei LVDVVRFLDWHGZLWKParahalomitra robusta > Sandalolitha robusta@%DVHGRQ3O;,,,ÀJ'(

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7DNHGDDQG7DPXUDFROOHFWHGIURPFungia > Pleuractis@paumotensisPRVWOLNHO\FRQ- stitutes F. syzygia VSQRYVHHUHPDUNVLQWKHVSHFLHVGHVFULSWLRQRIF. syzygia VSQRY

Remarks..URSSDVWDWHGWKDWWKHORFDWLRQRIWKHKRORW\SHZDVXQNQRZQKRZHYHUWKH

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Type material. Naturalis Biodiversity Center (Leiden, The Netherlands). Holotype:

501+&UXV'KRVWPleuractis granulosa .OXQ]LQJHU[RYLJIHPDOH

ðOHJ6(7YDQGHU0HLM%:+RHNVHPDWKHKRORW\SHZLOOEHGHSRVLWHGLQWKHMuseum Zoologicum Bogoriense, Bogor, Indonesia ZLWKFDWDORJXHQXPEHU0=%&UXallotype:

501+&UXV'PDOH7DQMXQJ3DVLU3XWLK7HUQDWH,QGRQHVLD1ƒ·µ(ƒ·

µKRVWPleuractis granulosa .OXQ]LQJHU[LPDOHðOHJ6(7YDQ

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1ƒ·µ(ƒ·µKRVWPleuractis granulosa .OXQ]LQJHU[LRYLJ

IHPDOHðOHJ6(7YDQGHU0HLMLee Kong Chian Natural History Museum (Singa- pore) =5&  H[ 501+&UXV' 6XODPDGDKD %D\ 7HUQDWH ,QGRQHVLD 1

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Fig. 6. $OORW\SHFungicola syzygia VSQRY501+&UXV'AKDELWXVGRUVDOYLHZB anterolateral PDUJLQRIFDUDSDFHYHQWUDOYLHZC0;3DDEGRPHQEJRQRSRGFWKRUDFLFVWHUQLWHVGOHIW3

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 $EGRPHQUKRPERLGORQJHVWDQGZLGHVWDWWKVHJPHQWWHOVRQURXQGHG)LJ'

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RFFXULQVSHFLPHQVRIF. fageiKHQFHFRORXUSDWWHUQLVQRWDGLDJQRVWLFFKDUDFWHU

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VHH9HKRIet alLQSUHVV)LJ$0DLQKRVWJHQHUDDUHCycloseris and PleuractisEXWWKHUHDUH

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7KLVLVWKHIRXUWKVSHFLHVUHFRUGHGIURPPXVKURRPFRUDOV

Table 3. &KDUDFWHUVGLVWLQJXLVKLQJFungicola fagei and F. syzygiaVSQRY

F. fagei F. syzygia VSQRY

KRVWJHQXV Podabacia Sandalolitha Cycloseris Pleuractis

FDUDSDFHVKDSH VXEUHFWDQJXODU VXEUHFWDQJXODUWRVTXDURLG

&/&:DOO “Q  “Q 

&/&:RYLJI “Q  “Q 

&/&:P Q  “Q 

ERUGHUEHWZHHQRUELWDODQJOHVI PLOGO\XQGXODWLQJ VWUDLJKWWRPLOGO\XQGXODWLQJ ODWHUDOPDUJLQFDUDSDFH VRPHZKDWGHÀQHGOHVVFRQFDYH QRWZHOOGHÀQHGPRUHFRQFDYH

than F. syzygia VSQRY than F. fagei

PHUXV0;3 PRGHVWGLVWRODWHUDO SURQRXQFHGGLVWRODWHUDO

SURMHFWLRQ SURMHFWLRQ



Diagnostic characters. Fungicola fagei and F. syzygia VSQRYFDQEHVHSDUDWHGEDVHGRQ

WKHLUKRVWVSHFLÀFLW\DQG'1$)RUWKHODWWHUEDVHGRQWKHUHVXOWVRIWKLVVWXG\DSRVLWLYHLGHQWL- ÀFDWLRQFDQEHDFKLHYHGE\DQDO\VLVRIDVLQJOHPLWRFKRQGULDOPDUNHUF. fagei and F. syzygia VS

QRYDUHPRUHGLIÀFXOWWRVHSDUDWHEDVHGRQPRUSKRORJ\%RWKVSHFLHVVKRZFRQVLGHUDEOHLQtra- VSHFLÀFYDULDWLRQLQWKHVKDSHDQJOHRIWKHH[WHUQDORUELWDODQJOHDQGWKHERUGHUEHWZHHQWKH

RUELWDODQJOHV,QWUDVSHFLÀFYDULDWLRQLVDOVRREVHUYHGRQWKHODWHUDOPDUJLQV,QJHQHUDOLQF.

syzygia VSQRYWKHODWHUDOPDUJLQVDUHPRUHFRQFDYHWKHQLQF. fagei%DVHGRQWKHFDUDSDFH

OHQJWKFDUDSDFHZLGWKUDWLRQLWLVFOHDUWKDWF. fagei LVPRUHUHFWDQJXODUWKDQF. syzygia VSQRY

ZKLFKLVDOPRVWVTXDURLG7KHRQHDYDLODEOHPDOHRIF. fagei LVVRPHZKDWRIDQRXWOLHUSHUKDSV

WKHUHVXOWVRIVH[XDOGLPRUSKLVP$IWHUUHPRYDORIWKLVVLQJOHPDOHIURPWKH&/&:DOOFDWH- JRU\WKHPHDQUDWLREHFRPHV“Q 7DEOH

Distribution range. Fungicola syzygia LVVRIDUNQRZQIURPYDULRXVORFDWLRQVLQ,QGRQHVLD

0DOD\VLDDQGWKH0DOGLYHV7KHPDOHVSHFLPHQVRI7DNHGDDQG7DPXUDPD\EHORQJWRWKH

QHZVSHFLHVF. syzygia VSQRYVHHUHPDUNVZKLFKZRXOGDGG-DSDQWRWKHNQRZQGLVWULEXWLRQ

UDQJHLIWKHLGHQWLÀFDWLRQZHUHFRQÀUPHG7KHÀJXUHGVSHFLPHQLQ1Jet alZDVFROOHFWHG

LQ9DQXDWXVHHUHPDUNV7KHGLVWULEXWLRQRIF. syzygia VSQRYSRVVLEO\RYHUODSVWKHGLVWULEXWLRQ

RIWKHLUKRVWFRUDOVIRUIXQJLLGGLVWULEXWLRQVVHH+RHNVHPDEXWODUJHDUHDVRIWKHLUKRVW·V

GLVWULEXWLRQKDYHQRW\HWEHHQH[DPLQHGIRUJDOOFUDEV

Remarks.$VLEOLQJVSHFLHVLVDFU\SWLFVLVWHUVSHFLHVWZRVSHFLHVWKDWDUHWKHFORVHVWUHODWLYH

RIHDFKRWKHUDQGKDYHQRWEHHQGLVWLQJXLVKHGIURPRQHDQRWKHUWD[RQRPLFDOO\%LFNIRUGet al

Fungicola syzygia VSQRYLVWKHUHIRUHDVLEOLQJVSHFLHVRIF. fagei,WLVWKHWKLUGVSHFLHV

DVVLJQHGWRWKHJHQXVFungicola3UHYLRXVZRUNHUVPRVWOLNHO\GLGQRWHQFRXQWHUWKLVQHZVSHFLHV

LQWKHLUPDWHULDO)L]HDQG6HUqQHFROOHFWHGF. fagei VSHFLPHQVIURPS. dentata DQGSRVVL- EO\S. robustaVHHKRVWFRUDOVVHFWLRQRIF. fageiDQGGLGQRWFROOHFWJDOOFUDEVIURPCycloseris or Pleuractis7KHLUPDWHULDOWKHUHIRUHPRVWOLNHO\FRQVLVWVRIVROHO\F. fagei6SHFLPHQVRIF.

fagei ZHUHIXUWKHUPRUHFROOHFWHGGXULQJWKH5XPSKLXV,DQG,,H[SHGLWLRQVWR,QGRQHVLD.URSS

UHH[DPLQHGWKH5XPSKLXVPDWHULDOEXWPHQWLRQHGWKDWWKHVSHFLPHQVRIF. fagei ZHUHQR

ORQJHUDYDLODEOHDQGWKDWWKHKRVWFRUDOVZHUHQRWLQFOXGHG7DNHGDDQG7DPXUDGLGFROOHFW

WZRPDOHVSHFLPHQVZKLFKEDVHGRQWKHKRVWVSHFLÀFLW\GDWDFungia paumotensis > Pleuractis paumotensis@DQGWKHLOOXVWUDWLRQVSRVVLEO\FRXOGEHLGHQWLÀHGDVFungicola syzygia 1DWLRQDO

0XVHXPRI1DWXUHDQG6FLHQFH7RN\R1607&U1Jet al)LJLOOXVWUDWHGD

VSHFLPHQ RI Fungicola VS FROOHFWHG IURP P. paumotensis %: +RHNVHPD SHUV FRPP LQ

6DQWR9DQXDWXPleuractis paumotensis LVRQO\NQRZQWRKRVWFungicola syzygiaWKHUHIRUHWKLV

PDWHULDOLVWHQWDWLYHO\LQFOXGHGLQWKHV\QRQ\P\RIWKLVVSHFLHV

Etymology.6\]\JLDLQUHIHUHQFHWRWKHW\SHORFDOLW\7HUQDWHRQFHWKHZRUOG·VPDMRUSURGXF- HURIFORYHVSyzygium aromaticum /0HUUDQG3HUU\7KH/DWLQV\]\JLDLVGHULYHGIURPWKH

DQFLHQW*UHHNƱƳƥƳƢрƠ>VX]XJLD@IRUDSDLURIUHODWHGWKLQJVRUXQLRQUHIHUULQJWRWKHREOLJDWH

V\PELRVLVEHWZHHQWKHJDOOFUDEDQGLWVKRVWFRUDO

Discussion

6SHFLHVKDYLQJDVPDOOVL]HRUKLGGHQKDELWDWDUHRIWHQUHIHUUHGWRDV¶FU\SWLF·$OWKRXJKJDOOFUDEV

DUHVPDOODQGOLYHDKLGGHQOLIHLQVLGHWKHLUFRUDOVKRVWVDQGKHQFH·FU\SWLF·FU\SWLFVSHFLHVLV

KHUHLQUHIHUUHGWRDVWZRRUPRUHVSHFLHVWKDWZHUHSUHYLRXVO\FODVVLÀHGDVDVLQJOHRQHEHFDXVHRI

DWOHDVWVXSHUÀFLDOPRUSKRORJLFDOVLPLODULWLHV%LFNIRUGet al7KLVGHÀQLWLRQLVDSSOLFDEOH

to F. syzygia VSQRYZKLFKLVGLIÀFXOWWRVHSDUDWHIURPF. fagei EDVHGRQPRUSKRORJ\DORQH'1$

WD[RQRP\LVEHFRPLQJDQLQGLVSHQVDEOHWRROIRUXQUDYHOOLQJFU\SWLFVSHFLDWLRQ7KLVHVSHFLDOO\



KROGVWUXHIRUHQGRV\PELRQWVZKLFKKDYHSURORQJHGUHODWLRQVKLSVZLWKWKHLUKRVWVWKDWFDQOHDGWR

PRUSKRORJLFDOVWDVLVHJ*LWWHQEHUJHUDQG*LWWHQEHUJHU

Fungicola syzygia VSQRYDQGF. fagei DUHFOHDUO\VHSDUDWHGEDVHGRQ'1$DQGKRVWVSHFLÀF- LW\)LJFungicola syzygia VSQRYDQGF. fagei DUHPRUHFORVHO\UHODWHGWRHDFKRWKHUWKDQWR

F. utinomi7KHLQWUDVSHFLÀFYDULDWLRQLVORZ7DEOHV7KHPLWRFKRQGULDOPDUNHUV6DQG

&2,VKRZHGKLJKUHVROXWLRQDWVSHFLHVOHYHO)LJ6FRQWUDU\WRWKHQXFOHDUPDUNHU+)LJ6

7KHODWWHUPDUNHUUHFRYHUHGF. utinomi DVDPRQRSK\OHWLFFODGHH[FHSWIRU501+&UXV'

ZKLFKFOXVWHUHGEDVDOO\WRDOORWKHUVSHFLPHQV+GLGQRWGLVWLQJXLVKEHWZHHQF. fagei and F.

syzygia VSQRY$VWXG\E\'LQDSROLet alVKRZHGWKDW+LVPRVWLQIRUPDWLYHRQJHQXV

OHYHO LQ KHWHUREUDQFK *DVWURSRGD PDNLQJ WKLV PDUNHU D SRVVLEOH FDQGLGDWH IRU IXUWKHU UHFRQ- VWUXFWLRQVRIWKHGHHSHUSK\ORJHQHWLFUHODWLRQVKLSVZLWKLQWKH&U\SWRFKLULGDH

Fungicola syzygia VSQRYDQGF. fagei FDQDOVREHVHSDUDWHGEDVHGRQWKHLUKRVWRFFXUUHQFH

7KHXVXDOSDWWHUQDPRQJKRVWVSHFLÀFRUJDQLVPVLVDFRPPRQRU¶SUHIHUUHG·KRVWZLWKDQXPEHU

RIRWKHUKRVWVOHVVIUHTXHQWO\LQKDELWHG1RUWRQDQG&DUSHQWHU7KLVVHHPVWREHWKHFDVH

LQ IXQJLLGDVVRFLDWHG JDOO FUDEV DV ZHOO 7KH SUHVHQW UHVXOWV VKRZHG WKDW Fungicola fagei ZDV

VWULFWO\DVVRFLDWHGZLWKWKHVLVWHUJHQHUDPodabacia and SandalolithaZLWKP. crustacea as the most common host. Fungicola syzygia VSQRYLVPRVWO\DVVRFLDWHGZLWKWKHJHQHUDCycloseris and PleuractisZLWKP. granulosa and P. paumotensis as most common hosts. Fungicola utinomi LVDVVRFLDWHGZLWKDZLGHUUDQJHRIUHODWHGJHQHUDEXWLVPRVWRIWHQHQFRXQWHUHGLQLithophyllon repanda )L]HDQG6HUqQHYDQGHU0HLMDQG+RHNVHPD)LJ7DEOH6+RHNVHPD

et alPHQWLRQHGWKDWJDOOFUDEVDUHQRWYHU\KRVWVSHFLÀFEXWDWWKHWLPHRIZULWLQJWKHQHZ

VLEOLQJVSHFLHVRIFungicola fagei ZDVQRW\HWGLVFRYHUHG,WWXUQVRXWWKDWFungicola VSHFLHVDUH

VSHFLÀFWRDWOHDVWJHQXVOHYHO7KHVSHFLPHQVUHIHUUHGWRDV¶FU\SWRFKLULGVS·LQ+RHNVHPDet al.

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WKHLUFRUDOPRUSKRORJ\PDNHVIRUDQXQVXLWDEOHKDELWDWIRUJDOOFUDEVHJORQJWHQWDFOHVLQHelio- fungia actiniformis RUYHU\WKLQSODWHOLNHPRUSKRORJ\LQHalomitra clavator and Zoopilus echi- natus2WKHUVSHFLHVKDYHGLVWULEXWLRQVRXWVLGHWKHUDQJHLQZKLFKWKHÀHOGZRUNZDVFDUULHGRXW

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Acknowledgements

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Appendices S1-3 are available in the digital academic repository of Naturalis Biodiversity Center.