Four armoured scale from Australia, South Africa, and Mexico : Quarantine interceptions in Japan Title (: Coccoidea: )

Author(s) Takagi, Sadao

Insecta matsumurana. New series : journal of the Faculty of Agriculture Hokkaido University, series entomology, 72, Citation 77-94

Issue Date 2016-09

Doc URL http://hdl.handle.net/2115/62829

Type bulletin (article)

File Information 4 Takagi2.pdf

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Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP INSECTA MATSUMURANA

NEW SERIES 72: 77–94 SEPTEMBER 2016

FOUR ARMOURED SCALE INSECTS FROM AUSTRALIA, SOUTH AFRICA, AND MEXICO: QUARANTINE INTERCEPTIONS IN JAPAN (STERNORRHYNCHA: COCCOIDEA: DIASPIDIDAE)

By SADAO TAKAGI

Abstract

TAKAGI, S. 2016. Four armoured scale insects from Australia, South Africa, and Mexico: Quarantine interceptions in Japan (Sternorrhyncha: Coccoidea: Diaspididae). Ins. matsum. n. s. ±¿JV Four species of armoured scale insects intercepted at quarantine inspection in Japan are recorded, and remarks are made about some taxonomic questions they arouse. One of the species, found on XanthorrhoeaVSLPSRUWHGIURP$XVWUDOLDLVLGHQWL¿HG with Chionaspis xanthorrhoeae Fuller and transferred to Poliaspis. Another species, on Chamelaucium from Australia, is very close to 3ROLDVSLVFHUDÀRUD Hardy and Henderson. $WKLUGVSHFLHVZDVFROOHFWHGRQFXWÀRZHUVRIParanomus, Serruria, and Berzelia from 6RXWK$IULFDDQGLVLGHQWL¿HGZLWKRolaspis lounsburyi (Cooley). The fourth species, Opuntiaspis sp., was found on cactus fruits from Mexico, and is apparently different IURPWKHWKUHHNQRZQVSHFLHVRIWKHJHQXV

Author’s address. Hukuzumi 3-3-4-16, Toyohira-ku, Sapporo, 062-0043 Japan (VGWNJ#MFRP]DTQHMS).

Contents,QWURGXFWLRQʊPoliaspis xanthorrhoeae, n.comb. = Chionaspis xanthorrhoeae)XOOHUʊPoliaspisVSʊRolaspis lounsburyi &RROH\ ʊOpuntiaspis VSʊ5HIHUHQFHVʊ)LJXUHV

77 INTRODUCTION

 5HFHQWO\,KDYHKDGWKHRSSRUWXQLW\WRH[DPLQHVRPHVFDOHLQVHFWVLQWHUFHSWHGDW quarantine inspection in Japan. These insects, all mounted on slide, were submitted to PHIRULGHQWL¿FDWLRQE\0U7,QRXH0RML3ODQW3URWHFWLRQ6WDWLRQ7KH\LQFOXGHIRXU species which are notable taxonomically, and the results of my study on these species are JLYHQLQWKHSUHVHQWSDSHU  2QHRIWKHIRXUVSHFLHVLVLGHQWL¿HGZLWKChionaspis xanthorrhoeae on the basis of WKHRULJLQDOGHVFULSWLRQDQRWKHUVSHFLHVLVYHU\VLPLODUWRWKHGHVFULSWLRQRIPoliaspis cerafloraEXWGRHVQRWH[DFWO\DJUHHZLWKWKHODWWHU LWLVFDOOHGµPoliaspis sp.Q’ in WKLVSDSHUµ4¶VWDQGLQJIRUTXDUDQWLQHLQWHUFHSWLRQ DWKLUGVSHFLHVLVLGHQWL¿HGZLWK Rolaspis lounsburyi,QH[DPLQLQJWKHVHWKUHHVSHFLHV,KDYHEHHQLPSUHVVHGZLWKWKH resemblance they show in some features of taxonomic importance, and I have considered WKHSRVVLELOLW\WKDWWKHVHVSHFLHVEHORQJWRRQHDQGWKHVDPHJHQXV,QWKHSUHVHQWSDSHU KRZHYHU,SUHIHUWRSODFHWKHIRUP,KDYHLGHQWL¿HGZLWKC. xanthorrhoeae in Poliaspis, and treat Poliaspis and RolaspisDVFORVHO\UHODWHGEXWGLIIHUHQWJHQHUDWHQWDWLYHO\7KH ODVWRIWKHIRXUH[DPLQHGVSHFLHVEHORQJVWROpuntiaspis, and is probably an undescribed IRUP µOpuntiaspis VS4¶LQWKLVSDSHU 7KHTXDUDQWLQHUHFRUGVVXJJHVWWKDWRolaspis lounsburyiLVQRWUDUHRQFXWÀRZHUVLPSRUWHGIURP6RXWK$IULFDOpuntiaspis sp.Q may also be worthy of attention from the viewpoint of its potential economic importance.  ,QWKLVSDSHUWKHWHUPµWUXOODH¶LVDGRSWHGLQSODFHRIµS\JLGLDOOREHV¶LQDXWKRUV,Q the description of Poliaspis xanthorrhoeae, WKHDEEUHYLDWLRQVµDEG,¶µDEG,,¶DQGVRRQ DUHXVHGIRUWKHFRUUHVSRQGLQJDEGRPLQDOVHJPHQWV  ,ZRXOGOLNHWRPDNHP\DFNQRZOHGJHPHQWWR0U7,QRXHIRUKLVJLYLQJPHWKH RSSRUWXQLW\WRVWXG\WKHVHLQWHUHVWLQJVFDOHLQVHFWV

RECORDS AND REMARKS

Poliaspis xanthorrhoeae, n.comb. =Chionaspis xanthorrhoeae Fuller, 1899 )LJV

Citations from Fuller (1899) p. 435 7KHIROORZLQJ1RWHVDQG'HVFULSWLRQVDUHWKHRXWFRPHRIVRPHPRQWKV¶UHVLGHQFHDW3HUWK RQWKH6ZDQ5LYHU:HVWHUQ$XVWUDOLDZKHUHLQWKHLPPHGLDWHYLFLQLW\RIWKHFLW\,FROOHFWHGWKH PDMRULW\RIWKHVSHFLPHQV p. 472 94. Chionaspis xanthorrhoeae, sp. n. 6FDOHRIƂVKLQLQJZKLWHS\ULIRUPH[XYLDHOLJKW\HOORZ $GXOWƂ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¿FHVDQWHULRUODWHUDOVWRSRVWHULRU

78 laterals 25 to 30. ƃSXSDULXPZKLWHQRWFDULQDWHGODUYDVNLQ\HOORZ On Xanthorrhoea, sp.

Material examined Four adult females, from Australia, on XanthorrhoeaVS ;DQWKRUUKRHDFHDH/LOLDOHV  LQWHUFHSWHGDWWKHSRUWRI

,GHQWL¿FDWLRQRIWKHVSHFLPHQV  7KHRULJLQDOGHVFULSWLRQLVDOPRVWOLPLWHGWRWKHS\JLGLXPDQGHVSHFLDOO\WRWKH PDUJLQDOIHDWXUHVRQWKHVL[WKDQGVXFFHHGLQJDEGRPLQDOVHJPHQWV,QWKHVHIHDWXUHV WKHIRXUVSHFLPHQVH[DPLQHGLQWKHSUHVHQWVWXG\DJUHHZHOOZLWKWKHGHVFULSWLRQZKLFK HYHQPHQWLRQVWKHRFFXUUHQFHRIµ7ZRYHU\VPDOOVSLQHVEHWZHHQWKHPHGLDQOREXOHV¶ WKHµPHVREDVDOSURFHVVHVRIWKHPHGLDQWUXOODH¶DVLQWHUSUHWHGLQWKHSUHVHQWVWXG\ see Remarks). However, the lateral lobule of the second trulla is somewhat variable in size and shape in the examined specimens: it is conical, with the apex blunt, in three VSHFLPHQV )LJ-$% ZKLOHLWLVUDWKHUURXQGHGDORQJWKHIUHHPDUJLQLQWKHRWKHU RQH )LJ& WKXVSUREDEO\DSSURDFKLQJWKHVWDWHPHQWLRQHGLQWKHRULJLQDOGHVFULSWLRQ :KDW)XOOHU¶VGHVFULSWLRQGHSLFWVLVDSDWWHUQRIFKDUDFWHUVZKLFKLVUDWKHUVSHFLDOLQWKH 'LDVSLGLQLVRWKDWWKHDJUHHPHQWRIWKHVSHFLPHQVZLWKWKHGHVFULSWLRQVXJJHVWVWKDW they are correctly referable to C. xanthorrhoeae unless more than one species which has such a pattern occurs on XanthorrhoeaDURXQG3HUWK7KHW\SHVHULHVLVSUHVXPHGWR have been lost (Miller and Gimpel, 2009). In the present circumstances, there may be QRDOWHUQDWLYHEXWWRUHJDUGWKHVSHFLPHQVDVEHORQJLQJWRC. xanthorrhoeae. However, WKLVLGHQWL¿FDWLRQLVQRPRUHWKDQDWHQWDWLYHWUHDWPHQWHLWKHU,WKHUHIRUHJLYHEHORZD GHVFULSWLRQEDVHGRQWKHIRXUVSHFLPHQVLQRUGHUWRDFFHSWFULWLFLVPRQP\LGHQWL¿FDWLRQ Needless to say, material from the type locality is essential to a critical study and for DGHILQLWHVHWWOLQJRIWKHPDWWHUC. xanthorrhoeae as understood in this study is not retainable in Chionaspis,RQFHFRQVLGHUHGWKHSRVVLELOLW\WKDWLWEHORQJVWRWKH$IULFDQ JHQXVRolaspis but in the present study it is transferred to Poliaspis, which is well represented in Australasia.

Description based on the four specimens  )XOO\JURZQDGXOWIHPDOHHORQJDWHZLGHVW across metathorax or abd I, about 2.2 to 2.4 WLPHVDVORQJDVZLGHSURVRPDPXFKORQJHUWKDQSRVWVRPDKHDGQDUURZLQJDQWHULRUO\ SURWKRUD[JUDGXDOO\DQGPHVRWKRUD[VOLJKWO\ZLGHQLQJSRVWHULRUO\PHWDWKRUD[DQGDEG ,±,,,OREHGODWHUDOO\S\JLGLXPPXFKZLGHUWKDQORQJURXQGLVKDORQJIUHHPDUJLQ  +HDGVWUHZQZLWKDERXW±PLFURGXFWVGRUVDOO\$QWHQQDHHDFKZLWKDVLQJOH VHWDRIPRGHUDWHOHQJWK$QWHULRUVSLUDFOHVHDFKZLWK±GLVFSRUHVZKLFKWHQGWRIRUP DVKRUWURZSRVWHULRUVSLUDFOHVZLWKQRGLVFSRUHVRUHDFKZLWKRU/HJYHVWLJHV SUHVHQWPHVRDQGPHWDWKRUDFLFOHJVHDFKUHSUHVHQWHGE\RUVPDOOVFOHURWL]HGFRQLFDO spines, at times much reduced.  &RQLFDOJODQGVSLQHVRQYHQWUDOVXUIDFHRIWKRUDFLFDQGSUHS\JLGLDODEGRPLQDO VHJPHQWVRUDQWHURODWHUDOO\DQG±SRVWHURODWHUDOO\WRDQWHULRUVSLUDFOH± RQPHVRWKRUD[IRUPLQJDORRVHJURXSLQLQWHUPHGLDWHDUHD±RQPHWDWKRUD[ SRVWHURODWHUDOO\WRSRVWHULRUVSLUDFOH±RQDEG,RQEDVHRIODWHUDOOREH±RQ,,RQ ODWHUDOOREHRURQ,,,DORQJPDUJLQRIODWHUDOOREH±RQ,9PDUJLQDOO\6OHQGHU

79 PDUJLQDOJODQGVSLQHVRQDEG9±9,,,HDFK WKHODVWEHLQJVLWXDWHGMXVWODWHUDOO\WR median trulla). Small ducts on meso- and metathorax, strewn on body side dorsally to ventrally, ±RQPHVRWKRUD[DQG±RQPHWDWKRUD[/DWHUDOPDFURGXFWVRQDEG,±,,, FRQÀXHQWZLWKVXEPDUJLQDOGRUVDOPDFURGXFWVZKLFKRFFXUQRWRQO\DORQJWKHSRVWHULRU PDUJLQ VHJPHQWDOO\ EXWDOVRRQWKHLQQHUDUHD LQIUDVHJPHQWDOO\ RQHDFKVHJPHQW ODWHUDODQGVXEPDUJLQDOPDFURGXFWVFRPELQHG±RQ,±RQ,,DQG± RQ,,,6XEPDUJLQDOPDFURGXFWVVHJPHQWDOO\DQGRULQIUDVHJPHQWDOO\RULQ WRWDORQDEG,9±VHJPHQWDOO\DQGRULQIUDVHJPHQWDOO\±LQWRWDORQ9 6PDOOVXEPHGLDQGXFWVRQDEG,DQG,,DORQJSRVWHULRUPDUJLQRIHDFKVHJPHQW± RQ,DQG±RQ,,6XEPHGLDQPDFURGXFWVRQDEG,,,±9,±VHJPHQWDOO\DQG± LQIUDVHJPHQWDOO\WRWDO±RQ,,,±DQG±WRWDORURQ,9±DQG± WRWDO±RQ99,ZLWK±LQDURZ LQFDVHDOVRZLWKVXEPDUJLQDOO\ 0DUJLQDO PDFURGXFWVRQDEG,9HDFKZLWKDUREXVWSRUHSURPLQHQFHRQ9DQGDOVRRQ9, PHVDORQHZLWKDUREXVWSRUHSURPLQHQFHRQHDFKVHJPHQWRQ9,, MXVWPHVDOO\WR VHFRQGWUXOOD ZLWKDVPDOODQGVOHQGHUVSLQH6XEPDUJLQDOGRUVDOERVVHVRQHRQHDFKRI abd I, III, and V, removed much mesally on I.  3HULYXOYDUGLVFSRUHVLQJURXSV±LQPHGLDQ±LQHDFKDQWHURODWHUDODQG ±LQHDFKSRVWHURODWHUDOJURXSWRWDO±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conical processes laterally to pore prominence.

Remarks Hardy and Henderson (2011) made a nearly complete revision of Poliaspis and UHFRJQL]HGVSHFLHVRIWKHJHQXVRQO\WZRRWKHUVNQRZQDWWKDWWLPHKDYLQJEHHQOHIW unrevised. P. xanthorrhoeaeDVXQGHUVWRRGLQP\VWXG\GRHVQRWDJUHHDWOHDVWLQWKUHH points with the concept of Poliaspis presented by them. First, this species has no extra abdominal disc pores, whereas all the species examined by them are provided with H[WUDµSHULYXOYDU¶GLVFSRUHV6HFRQGO\WKLVVSHFLHVKDVWKHPHGLDQWUXOODHDSSDUHQWO\ QRWFRQÀXHQWEDVDOO\ZKHUHDVDFFRUGLQJWRWKHPWKHPHPEHUVRIWKHJHQXVH[FHSWP. FHUDÀRUDKDYHµ]\JRWLF¶PHGLDQWUXOODH7KLUGO\WKLVVSHFLHVSRVVHVVHVDSDLURIVPDOO slender spinous processes on the mesal bases of the median trullae, whereas the presence of such processes is not clearly mentioned in all the species examined by them. The question, therefore, may arise why Chionaspis xanthorrhoeae should be transferred into PoliaspisLQVSLWHRIDOOWKHVHGLVDJUHHPHQWV$QDWWHPSWLVPDGHEHORZWRFKHFN WKHGLVDJUHHPHQWVDQGWRFOHDUXSWKHTXHVWLRQ,QWKHSUHVHQWVWXG\WZRRWKHUVSHFLHV Poliaspis incisa and the form treated as Poliaspis sp.Q, have been available for my examination. P. incisaZDVGHVFULEHGLQ7DNDJLDQG'H)DYHUL  DQGDFFHSWHGE\ Hardy and Henderson (2011) as a member of PoliaspisP. sp.Q is probably very close to 3FHUDÀRUD, which was described by Hardy and Henderson as a species of Poliaspis in 80 VSLWHRIWKHµQRQ]\JRWLF¶PHGLDQWUXOODH First, extra disc pores occur in P. incisa and P. sp.Q as usual in Poliaspis. In SULQFLSOHWD[DDUHUHFRJQL]HGRQWKHEDVLVRIVWDWLVWLFDOO\FRUUHODWHGFKDUDFWHUV,QWKH Diaspididae, the occurrence of extra abdominal disc pores is scattered in restricted parts RIWKHIDPLO\DQGRIWHQDUELWUDU\ZLWKWKHFRQWH[WRIWD[D7KLVIDFWVXJJHVWVWKDWWKHVH GLVFSRUHVUHSUHVHQWDSULPLWLYHIHDWXUHZKLFKPD\KDYHEHHQSHUVLVWLQJ DVDUHOLF RU PD\KDYHHPHUJHGDEUXSWO\DQGDWDYLVWLFDOO\ DVDUHFDOO LQWKHFRXUVHRIHYROXWLRQ The taxonomic value of this feature, therefore, should depend on the manner of its appearance in taxa under study. Lepidosaphes piniRULJLQDOO\GHVFULEHGDVPoliaspis piniE\0DVNHOOPD\DIIRUGDJRRGH[DPSOHIRUFRQVLGHULQJWKHPDWWHU7KLVVSHFLHVLV H[FHSWLRQDOLQKDYLQJH[WUDµSHULYXOYDU¶GLVFSRUHVEXWRWKHUZLVHQRWH[WUDRUGLQDU\LQ WKHJHQXVLepidosaphesDQGLQWKHWULEH/HSLGRVDSKLGLQL,QWKLVFDVHWKHRFFXUUHQFHRI WKHVHGLVFSRUHVSURYLGHVDGLDJQRVWLFFKDUDFWHUXVHIXOIRUUHFRJQL]LQJWKHVSHFLHVDPRQJ WKHPHPEHUVRIWKHJHQXVDQGWKHWULEHEXWZKHWKHULWUHSUHVHQWVDUHOLFRUDUHFDOOLWKDV no further taxonomic value. Occasionally, extra disc pores occur only to the extent of DEQRUPDOLQGLYLGXDOYDULDWLRQ DSSDUHQWO\DWDYLVWLFDOO\ZKHQRFFXUULQJLQGLDVSLGLGVRI derivative taxa) and in such a case their occurrence has no taxonomic value. Secondly, in PVS4WKHPHGLDQWUXOODHDUHREYLRXVO\QRQ]\JRWLFDVLQ3FHUDÀRUD. In P. incisa, the bases of the median trullae are separated from each other by a wide VSDFHDQGWKHYHQWUDOS\JLGLDOPDUJLQLQWKLVVSDFHLVGHHSO\LQFLVHGDQGRQO\QDUURZO\ sclerotic. In P. xanthorrhoeae, the bases of the median trullae extend anteromesally, but WKHLUH[WHQVLRQVGRQRWDGMRLQHDFKRWKHUWRIRUPD\RNH,QWKHVWDWHDURXQGWKHEDVHV RIWKHPHGLDQWUXOODHWKHVHWKUHHVSHFLHVDUHQRWWKHVDPHDQGDFFRUGLQJO\Poliaspis VKRXOGQRWEHXQLIRUPWKURXJKRXW7KHPHGLDQWUXOODHLQWKHW\SHVSHFLHVRIPoliaspis, P. media (or P. cycadis or P. gaultheriaeERWKKDYLQJEHHQV\QRQ\PL]HGZLWKP. media), ZHUHGHVFULEHGDVEDVDOO\µFRQÀXHQW¶µ\RNHGWRJHWKHU¶RUµ]\JRWLF¶E\DXWKRUVWKRXJK DOVRDVµQRWFRQWLJXRXV¶E\0RUULVRQDQG0RUULVRQ  7KHGHVFULSWLRQRIP. cycadis in Balachowsky (1954, p. 428) may be pertinent to the matter under consideration. He

VWDWHVµ/1«DFFROpHVO¶XQHjO¶DXWUHVHXOHPHQWSDUXQHPHPEUDQHLQWHUQHVFOpURVpH dans leur partie saillante, les palettes sont séparées à leur base par un espace médian FRPSOHW¶:KDWLVVWDWHGKHUHPD\EHXQGHUVWRRGDVIROORZVWKHPHGLDQWUXOODHDUH SURGXFHGZLWKWKHLUEDVHVVHSDUDWHGIURPHDFKRWKHUE\DJRRGVSDFHZKLOHWKHGRUVDO GHUPFRYHULQJWKHEDVHVDQGWKHVSDFHLVVRVFOHURWL]HGDVWRPDNHWKHWUXOODHDSSHDU WREHFRQÀXHQWEDVDOO\$FFRUGLQJWRWKLVLQWHUSUHWDWLRQWKHPHGLDQWUXOODHLQWKHW\SH VSHFLHVDUHQRWXQLWHGE\DVFOHURWL]HG\RNHQRUDUHWKH\FRPELQHGE\D]\JRWLFVFOHULWH (In this connection, see Remarks under Rolaspis lounsburyi and also Remarks under Poliaspis sp.Q.) Thirdly, P. sp.Q is provided with a pair of small spinous processes between the median trullae, and P. incisa with a pair of much reduced processes on the mesal bases of the median trullae. P. FHUDÀRUD, which is probably very close to P. sp.Q, is also provided ZLWKDSDLURIVKRUWVSLQHVMXVWPHVDOO\WRWKHPHGLDQWUXOODHVRIDUDVSUHVXPHGIURP WKH¿JXUHLQ+DUG\DQG+HQGHUVRQ ¿J +HQGHUVRQ  VWXGLHGWKHVHFRQG instar females of six species and the second instar males of four species of Poliaspis RFFXUULQJLQ1HZ=HDODQG+HU¿JXUH ¿J RIWKHVHFRQGLQVWDUIHPDOHRIP. media, the type species, shows a pair of processes on the mesal bases of the median trullae. 6XFKSURFHVVHVZKLFKPD\EHWHUPHGµPHVREDVDOSURFHVVHVRIWKHPHGLDQWUXOODH¶ DUHQRWVKRZQLQKHU¿JXUHVRIWKHVHFRQGLQVWDUIHPDOHVRIWKHRWKHU¿YHVSHFLHV+HU

81 ILJXUHV ILJV RIWKHVHFRQGLQVWDUPDOHVRIWKHIRXUVSHFLHV are provided on the mesal bases of the median trullae with a pair of small processes, RIZKLFKWKHSUHVHQFHLVPHQWLRQHGE\KHUDVIROORZVµPHGLDQOREHVELOREDWHZLWKD VPDOOSRLQWHGOREXOHPHGLDOWRHDFKPDLQOREH¶µDVPDOOSRLQWHGOREXOH¶KHUHDSSDUHQWO\ FRUUHVSRQGLQJWRHDFKPHVREDVDOSURFHVV,QRQHRIWKHIRXUVSHFLHV3ÀRFFRVD ¿J 183), a macroduct is associated with each of the mesobasal processes. The view may be adopted, therefore, that these processes are not lobules of the median trullae but UHSUHVHQWWKHSRUHSURPLQHQFHVRIWKHHLJKWKDEGRPLQDOVHJPHQWDQGWKDWWKHDEVHQFHRI PDUJLQDOPDFURGXFWVRQWKHSURFHVVHVLQWKHVHFRQGLQVWDUQ\PSKVRIWKHRWKHUVSHFLHV she examined is due to their evolutionary disappearance. If this view is correct, the VOHQGHUVSLQRXVSURFHVVHVRFFXUULQJRQWKHPHVDOEDVHVRIWKHPHGLDQWUXOODHLQWKHDGXOW female of P. xanthorrhoeaeVKRXOGDOVREHWKHSRUHSURPLQHQFHVRIWKHHLJKWKDEGRPLQDO VHJPHQWRULJLQDOO\ HDFKRIWKHPEHLQJQRORQJHUDFFRPSDQLHGZLWKDPDUJLQDO PDFURGXFW EHFDXVHWKH\DSSDUHQWO\FRUUHVSRQGWRWKHPHVREDVDOSURFHVVHVRFFXUULQJ in the second instar nymphs. In fact, the adult female of this species has the pore SURPLQHQFHVRIWKHVHYHQWKDEGRPLQDOVHJPHQWWRRPRGLILHGLQWRVOHQGHUSURFHVVHV HDFKRIWKHPEHLQJWRRVOHQGHUWRDFFRPPRGDWHWKHRUL¿FHRIWKHPDUJLQDOPDFURGXFW combined with it). It should be added that in P. incisa not only the adult female but also the second instar female and male are provided between the median trullae with a pair RIVPDOOSURFHVVHV HDFKEHLQJDFFRPSDQLHGZLWKQRPDFURGXFWLQHLWKHUVH[ WKRXJK these processes in the female nymph are sometimes so rudimentary that they may easily be overlooked. Furthermore, in the character pattern of the second instar male, P. incisa is very similar to two of the four species studied by Henderson, P. lactea ¿J DQGP. media ¿JV  $VWRWKHVSLQRXVSURFHVVHVRFFXUULQJEHWZHHQWKHPHGLDQWUXOODH in the adult females of P. sp.Q and3FHUDÀRUD, see under Poliaspis sp.Q.)  7DNLQJDOOWKHVHFRQVLGHUDWLRQVLQWRDFFRXQW,KDYHFRQFOXGHGWKDWWKHUHLVQR GH¿QLWHUHDVRQIRUH[FOXGLQJP. xanthorrhoeae from Poliaspis)XUWKHUPRUHLQFKHFNLQJ published descriptions, I have had the impression that many species of Poliaspis are to EHFDUHIXOO\UHH[DPLQHGIRUWKHVWDWHDURXQGWKHEDVHVRIWKHPHGLDQWUXOODH&RPSDULQJ ZHOOVWDLQHGVSHFLPHQVZLWKOLWWOHRUQRQVWDLQHGRQHVPD\EHXVHIXOIRUHOXFLGDWLQJWKH state of this part (see Remarks under Rolaspis lounsburyi).

Poliaspis sp. (PµVS4¶ )LJV

Material examined (LJKWDGXOWIHPDOHVIURP$XVWUDOLDRQChamelauciumVS 0\UWDFHDH OHDILQWHUFHSWHGDW Narita Airport, 26.VI.1996, M. Takada. 7KHPRXQWHGVSHFLPHQVDUHJHQHUDOO\QRWLQJRRGFRQGLWLRQREVHUYDWLRQVRQWKHGHWDLOVRI features have been restricted to a few better specimens.

&RPSDULVRQZLWK3ROLDSLVFHUDÀRUD Poliaspis sp.Q is probably very close to Poliaspis ceraflora. The latter was GHVFULEHGE\+DUG\DQG+HQGHUVRQ  RQWKHEDVLVRIPDWHULDOFROOHFWHGDW3HUWK :HVWHUQ$XVWUDOLDRQChamelaucium uncinatum VL[DGXOWIHPDOHVLQFOXGLQJWKH holotype) and MelaleucaVS VL[DGXOWIHPDOHV ERWKWKHVHSODQWVEHORQJLQJWRWKH family Myrtaceae. P. sp.Q and 3FHUDÀRUDWKHUHIRUHPD\QRWGLIIHULQWKHUDQJHRIKRVW 82 plants. P. sp.Q and P. cerafloraDJUHHLQKDYLQJWKHPHGLDQWUXOODHVPDOODQGZLGHO\ separated from each other. P. sp.Q is provided between the median trullae with a pair of spinous processes, which are smaller than the median trullae but may not easily be RYHUORRNHGWKHGHVFULSWLRQRIP. ceraflora makes no mention of such processes, but WKH¿JXUH ¿J VXJJHVWVWKHRFFXUUHQFHRIDVPDOOVSLQHPHVDOO\WRWKHPHGLDQWUXOOD These processes in both forms occur closely to the median trullae rather than on the PHVDOEDVHVRIWKHODWWHUEXWWKH\XQGRXEWHGO\FRUUHVSRQGWRWKHµPHVREDVDOSURFHVVHV¶ discussed in Remarks under Poliaspis xanthorrhoeae. The two forms remarkably differ in the shape of the median trullae: in P. sp.Q these WUXOODHDUHHORQJDWHQDUURZO\FRQLFDODQGSRLQWHGDWWKHDSH[ZKHUHDVLQ3FHUDÀRUD WKHVHWUXOODHDUHVKRUWDVORQJDVZLGHDQGEURDGO\URXQGHGDORQJWKHDSLFDOPDUJLQ VRIDUDVEDVHGRQWKHGHVFULSWLRQ µHDFKOREHZLWKURXQGHGDSH[¶ DQGWKH¿JXUH7KH occurrence of the conical median trullae in P.VS4KDVEHHQFRQ¿UPHGLQVL[VSHFLPHQV (but not in the other two specimens, which are badly distorted), and the rounded ones in P. FHUDÀRUD should have been based on the 12 specimens. The shape of the median trullae, therefore, should be stable in either form. P. sp.Q and 3FHUDÀRUDDJUHHLQKDYLQJRQO\WZRVPDOOPDUJLQDOJODQGVSLQHVRQ HDFKVLGHRIWKHS\JLGLXPRQHMXVWODWHUDOO\WRWKHPHGLDQWUXOODDQGWKHRWKHUODWHUDOO\WR WKHVHFRQGWUXOOD7KLVFKDUDFWHULVUDWKHUUDUHLQWKHJHQXVEHLQJUHVWULFWHGWRWKHVHWZR forms and another species (P. callitris, see Remarks below). P. sp.Q may be similar to 3FHUDÀRUD in the occurrence of ducts, but it is not easy WRPDNHDFRPSDULVRQZLWKWKH¿JXUH ¿J RIWKHODWWHUDVWRWKHGHWDLOVRIPDFURDQG PLFURGXFWVRFFXUULQJLQWKHSUHS\JLGLDOUHJLRQ7KHVHIRUPVDJUHHLQKDYLQJDJRRG QXPEHURIPLFURGXFWVVFDWWHUHGRQWKHGRUVDOVXUIDFHRIWKHKHDGP. sp.Q, in addition, has a variable number of microducts on the ventral surface of the meso- and metathorax LQDEURDGPHGLDQDUHD WKHVHPLFURGXFWVEHLQJQXPHURXVLQWKHVSHFLPHQVKRZQLQ)LJ DQG ZKHUHDVWKH¿JXUHRI3FHUDÀRUD shows no such microducts. In P. sp.Q each DQWHQQDLVSURYLGHGZLWKWZRVHWDHDWOHDVWXVXDOO\RQHRIWKHPEHLQJPXFKVPDOOHUWKDQ WKHRWKHUZKHUHDVWKH¿JXUHRI3FHUDÀRUDVKRZVDVLQJOHVHWDRQWKHDQWHQQD The difference in the shape of median trullae is so remarkable between P. sp.Q DQGWKHGHVFULSWLRQDQG¿JXUHRI3FHUDÀRUD that these forms seem to represent distinct species (provided that this difference is real and stable). If this view is correct, there should be noticeable differences between the two forms also in other features, especially in the occurrence of dorsal macro- and microducts and also of ventral microducts. Detailed examinations on these forms, preferably based on further samples, and HVSHFLDOO\RQYDULDWLRQLQHDFKIRUPPD\EHQHFHVVDU\IRUFOHDULQJXSWKHUHODWLRQ between them.

Remarks Poliaspis callitris was described from CallitrisLQ/DLQJ  DQGUHFRUGHG from Callitris and CupressusLQ+DUG\DQG+HQGHUVRQ  ERWKWKHVHSODQWJHQHUD EHORQJLQJWRWKHIDPLO\&XSUHVVDFHDH7KHPHGLDQWUXOODHLQWKLVVSHFLHVDUHµ]\JRWLF¶ in Hardy-Henderson’s description, whereas separated by a broad space from each other, ZLWKQRFRQQHFWLQJVFOHURWL]HG\RNHEHWZHHQWKHPLQ/DLQJ¶VILJXUH ILJ 7KLV difference should not be real, because the material examined by Hardy and Henderson

83 included specimens from the type series (in this connection, see Remarks under Poliaspis xanthorrhoeae and also Remarks under Rolaspis lounsburyi). P. sp.Q, 3FHUDÀRUD, and P. callitrisDUHFRPPRQO\FKDUDFWHUL]HGLQKDYLQJVPDOO median trullae, which are distinctly separated from each other in the former two and SUREDEO\DOVRLQWKHWKLUGVSHFLHVDQGLQKDYLQJRQO\WZRSDLUVRIVPDOOPDUJLQDO JODQGVSLQHVRQWKHS\JLGLXP7KH\PD\IRUPDFORVHJURXSDQGDUHWREHUHYLVHGDQG compared from this viewpoint.

Rolaspis lounsburyi =Chionaspis lounsburyi Cooley, 1898  )LJV

Material examined Two adult females, from South Africa, on cut flowers of ParanomusVS 3URWHDFHDH  intercepted at Narita Airport, 3.VII.2005, H. Hayashi. 2QHDGXOWIHPDOHIURP6RXWK$IULFDRQFXWÀRZHURISerruriaVS 3URWHDFHDH LQWHUFHSWHG at Narita Airport, 15.VII.2011, K. Kaneshiro. Three adult females, from South Africa, on cut flowers of BerzeliaVS %UXQLDFHDH  intercepted at Fukuoka Airport, 19.VIII.2012, T. Inoue.

,GHQWL¿FDWLRQRIWKHVSHFLPHQV  7KHVSHFLPHQVH[DPLQHGLQWKHSUHVHQWVWXG\QHDUO\DJUHHZLWKWKHGHVFULSWLRQ DQGILJXUHRIRolaspis lounsburyiLQ0XQWLQJ ILJ 7KH\GLVDJUHHZLWK WKHODWWHULQKDYLQJWKHERG\QRWµKHDYLO\VFOHURWL]HGDFURVVWKHPHWDWKRUD[DQGILUVW DEGRPLQDOVHJPHQW¶DWIXOOJURZWKLQODFNLQJVPDOOGXFWVDORQJWKHSRVWHULRUPDUJLQ RIWKHPHWDWKRUD[ ZKHUHDVILJVKRZVVRPHGXFWVVFDWWHUHGWKHUH LQKDYLQJRQH VXEPDUJLQDOGRUVDOERVVRQHDFKRIWKHILUVWDQGILIWKDEGRPLQDOVHJPHQWVRURQO\ RQWKHILUVW ZKHUHDVµ'RUVDOVXEPDUJLQDOERVVHVSUHVHQWRQVHJPHQWV,DQG,,,¶LQ 0XQWLQJ¶VGHVFULSWLRQ ,GRQRWWKLQNWKDWWKHVHGLVDJUHHPHQWVSXWVHULRXVGLI¿FXOWLHV LQP\LGHQWLILFDWLRQWKHVFOHURWL]DWLRQRIWKHGHUPLVDJURZWKSKHQRPHQRQDQGLQ JHQHUDOOLDEOHWRYDU\ZLWKVSHFLPHQVLQGHJUHHDQGH[WHQWWKHGRUVDOGXFWVDUHµYHU\ YDULDEOHLQQXPEHU¶ 0XQWLQJ LQR. lounsburyi (somewhat variable on anterior DEGRPLQDOVHJPHQWVHYHQLQWKHVL[VSHFLPHQVDYDLODEOHIRUWKHSUHVHQWVWXG\ DQGD IRUPKDYLQJVRPHFKDUDFWHUVRIGRUVDOGXFWVDQGRQFHGLVWLQJXLVKHGWKHUHE\IURP R. lounsburyiLVFRQQHFWHGZLWKWKHODWWHUWKURXJKµDOOLQWHUPHGLDWHFRQGLWLRQV¶ 0XQWLQJ  WKHRFFXUUHQFHRIGRUVDOERVVHVLVDSSDUHQWO\XQVWDEOHLQWKHVSHFLPHQVH[DPLQHG in the present study. R. lounsburyiKDVEHHQUHFRUGHGIURPYDULRXVSODQWVEHORQJLQJWR QLQHIDPLOLHV 0LOOHUDQG*LPSHO LQWKHSUHVHQWVWXG\DQRWKHUIDPLO\DQGWKUHH RWKHUJHQHUDDUHDGGHGWRWKHOLVWRIKRVWSODQWV%HLQJEURDGO\GLVWULEXWHGLQVRXWKHUQ $IULFDDQGRFFXUULQJRQYDULRXVNLQGVRISODQWVWKLVVSHFLHVVKRXOGEHYDULDEOHLQVRPH PRUSKRORJLFDOIHDWXUHVDFFRUGLQJO\  0XQWLQJ  PDGHQRPHQWLRQRIWKHVWDWHDURXQGWKHEDVHVRIWKHPHGLDQWUXOODH in R. lounsburyi. In the course of the present study, I have been under the impression that the state of this part is to be re-examined carefully and critically not only in Poliaspis (see Remarks under Poliaspis xanthorrhoeae) but also in Rolaspis and some other African JHQHUD VHHRemarks below).

84 Remarks  7KLVVSHFLHVZDVRULJLQDOO\GHVFULEHGLQChionaspisDJHQXVQDPHGHDUO\LQ armoured scale . It was transferred by authors to Phenacaspis, Dinaspis, and Trichomytilus but, needless to say, all in misconception. Hall (1946) in his study on the 'LDVSLGLQLRIWKH(WKLRSLDQ5HJLRQHUHFWHGRolaspisWRJHWKHUZLWKRWKHUJHQHUDFORVHO\ related to it, and since that time the species has been treated as a member of Rolaspis. In WKHFRXUVHRIWKHSUHVHQWVWXG\KRZHYHU,KDYHUHDOL]HGWKHQHFHVVLW\RIUHH[DPLQLQJ WKHWD[RQRPLFUHODWLRQVQRWRQO\DPRQJWKHVH$IULFDQJHQHUDEXWDOVREHWZHHQWKHP and Poliaspis (see Introduction and Identification of the specimens under Poliaspis xanthorrhoeae).  :KHQ+DOOHUHFWHG RolaspisKHGHVFULEHGWKHPHGLDQWUXOODHLQWKHJHQXVDV µFOHDUO\\RNHGWRJHWKHU¶7KUHHRIWKHVSHFLPHQVH[DPLQHGLQWKHSUHVHQWVWXG\DUH well stained, while the other three have no trace of stain. In the latter three specimens observed in phase-contrast microscopy the median trullae have their mesal bases H[WHQGLQJDQWHURPHVDOO\EXWWKHVHH[WHQVLRQVGRQRWDGMRLQHDFKRWKHUWRIRUPD\RNH ,QWKHZHOOVWDLQHGVSHFLPHQVWKHGRUVDOGHUPFRYHULQJWKHEDVHVRIWKHPHGLDQWUXOODH LVVRVFOHURWL]HGDVWRPDVNWKHGLVMRLQHGEDVDOH[WHQVLRQVDQGWRPDNHWKHWUXOODH DSSHDUFRQÀXHQWEDVDOO\7KLVVWDWHDURXQGWKHEDVHVRIWKHPHGLDQWUXOODHLVSUREDEO\ the same as that described by Balachowsky (1954) for Poliaspis cycadis (which was synonymized with P. media, the type species of Poliaspis) (see Remarks under Poliaspis xanthorrhoeae).  +DUG\DQG+HQGHUVRQ  PHQWLRQHGWKHRFFXUUHQFHRIH[WUDµSHULYXOYDU¶GLVF pores in all the species of Poliaspis they examined and in some species of the African JHQHUD Rolaspis, Tecaspis, and DentachionaspisVXJJHVWLQJFORVHUHODWLRQVDPRQJWKHVH JHQHUD 7KH\KRZHYHUUHIUDLQHGIURPµWDNLQJDQ\QRPHQFODWXUDODFWLRQ¶RQWKHVHJHQHUD for a certain reason.) So far as based on published descriptions in addition to my limited REVHUYDWLRQVDOOWKHVHJHQHUD DQGVoraspisDQRWKHU$IULFDQJHQXV DUHQRWLFHDEO\ VLPLODUDSSDUHQWO\VKRZLQJWKHLUFORVHUHODWLRQVLQWKHVWUXFWXUHRIWKHS\JLGLDOPDUJLQ DQGWKHDUUDQJHPHQWRIWKHGRUVDOPDFURGXFWVIHDWXUHVXQGRXEWHGO\RIWD[RQRPLF LPSRUWDQFH7KHIUHTXHQWRFFXUUHQFHRIH[WUDµSHULYXOYDU¶GLVFSRUHVFHUWDLQO\UHÀHFWV WKHFORVHUHODWLRQVDPRQJPoliaspisDQGWKH$IULFDQJHQHUD ,WVKRXOGKDYHKRZHYHU QRIXUWKHUWD[RQRPLFVLJQLILFDQFHVHHRemarks under Poliaspis xanthorrhoeae.) 7KHµPoliaspis complex’, comprised of all these forms, may afford another clue for HOXFLGDWLQJWKHIDXQDOFRQQHFWLRQEHWZHHQ$IULFDDQG$XVWUDODVLD

Opuntiaspis sp. (OµVS4¶ )LJV

Material examined 2QHDGXOWIHPDOHIURP0H[LFRRQ3LWD\DIUXLW &DFWDFHDH LQWHUFHSWHGDW1DULWD$LUSRUW 23.V.1996, G. Tokihiro. 2QHDGXOWIHPDOHIURP0H[LFRRQGUDJRQIUXLWHylocereus undatus &DFWDFHDH LQWHUFHSWHG at Narita Airport, 9.VIII.2006, S. Sekimoto.

Remarks  1DNDKDUD  UHFRJQL]HGWKUHHVSHFLHVLQOpuntiaspis. The two specimens H[DPLQHGLQWKHSUHVHQWVWXG\GLIIHUIURPSXEOLVKHGGHVFULSWLRQV DPRQJZKLFK*UHHQ 85 1905, Ferris, 1937, and Balachowsky, 1954, may be mentioned here) of the three species PDLQO\LQKDYLQJWKHVHFRQGWUXOODHXQLOREHGDQGLQKDYLQJWKHOHJYHVWLJHVUHVWULFWHG WRWKHPHVRDQGPHWDWKRUD[ WKHUHEHLQJQRWUDFHRQWKHSURWKRUD[ DQGPXFKPRUH rudimentary. They are similar especially to Opuntiaspis philococcus, which also occurs on Cactaceae. The available specimens are too few to show whether they represent some variation or abnormality within O. philococcusRUEHORQJWRDQRWKHUVSHFLHVZKLFKLV VWDEOHLQKDYLQJWKHFKDUDFWHUVPHQWLRQHGDERYH)XUWKHUPRUHERWKVSHFLPHQVDSSDUHQWO\ UHSUHVHQWJURZLQJDGXOWIHPDOHVIDLOLQJWRVKRZDQ\VSHFL¿FFKDUDFWHUVZKLFKPD\EH UHYHDOHGRUFRPSOHWHGDWIXOOJURZWK  ,KDYHIDLOHGWR¿QGLQOLWHUDWXUH any article or comment about this form, which may EHWUHDWHGDVDQXQGHVFULEHGVSHFLHVWHQWDWLYHO\7KHUHSHDWHGTXDUDQWLQHUHFRUGVVXJJHVW that Opuntiaspis sp.Q is an established , if not a serious pest, in some districts of cactus fruit production.

REFERENCES

%DODFKRZVN\$6/HV&RFKHQLOOHV3DOpDUFWLTXHVGHOD7ULEXGHV'LDVSLGLQL ,QVWLWXW3DVWHXU &RROH\5$1HZVSHFLHVRIChionaspis and notes on previously known species. &DQDGLDQ(QWRPRORJLVW± Ferris, G. F., 1937. Atlas of the Scale Insects of North America, I. Stanford University 3UHVV )XOOHU&1RWHVDQGGHVFULSWLRQVRIVRPHVSHFLHVRI:HVWHUQ$XVWUDOLDQ&RFFLGDH 7UDQVDFWLRQVRIWKH(QWRPRORJLFDO6RFLHW\RI/RQGRQIRUWKH

86 )LJPoliaspis xanthorrhoeaeDGXOWIHPDOHIXOO\JURZQ%GRUVDOERVVDQGGXFWVRQDEG, &ODWHUDOOREHRIDEG,,,GRUVDOVXUIDFH'S\JLGLDOPDUJLQDEG9DQG9,(DQWHQQD) DQWHULRUVSLUDFOH*OHJYHVWLJHRQPHVRWKRUD[+FRQLFDOJODQGVSLQHVRQPHWDWKRUD[,OHJ YHVWLJHRQPHWDWKRUD[-PHGLDQDQGVHFRQGWUXOODH6FDOHEDUȝP$ȝP%±-

87 )LJPoliaspis xanthorrhoeaeDGXOWIHPDOHDVHFRQGVSHFLPHQ $% $DEGRPHQ%S\JLGLDO PDUJLQ&RXWOLQHRIWUXOODHLQDWKLUGVSHFLPHQ6FDOHEDUȝP$ȝP%& 88 )LJPoliaspisVS4DGXOWIHPDOH%DQWHQQD&DQWHULRUVSLUDFOH6FDOHEDUȝP$ȝP% C.

89 )LJPoliaspisVS4DGXOWIHPDOH$DEGRPHQ%S\JLGLDOPDUJLQ6FDOHEDUȝP$ȝP% 90 )LJRolaspis lounsburyiDGXOWIHPDOHIXOO\JURZQ2QSerruria%GRUVDOGXFWVRQDEG,& GRUVDOPDFURGXFWVRQDEG,,'GRUVDOPDFURGXFWVRQDEG,,,(S\JLGLDOPDUJLQDEG9DQG 9,GRUVDOVXUIDFH)DQWHQQD*DQWHULRUVSLUDFOH+FRQLFDOJODQGVSLQHVRQPHVRWKRUD[, FRQLFDOJODQGVSLQHVDQGDGXFWRQDEG,-ODWHUDOOREHRIDEG,,,YHQWUDOVXUIDFH.PHGLDQ DQGVHFRQGWUXOODH6FDOHEDUȝP$ȝP%±.

91 )LJRolaspis lounsburyi, adult female. On Paranomus$DEGRPHQ%S\JLGLDOPDUJLQ6FDOH EDUȝP$ȝP%

92 )LJOpuntiaspisVS4$GXOWIHPDOHLQDJURZLQJVWDJH2QGUDJRQIUXLW H[FHSWIRU' % DQWHULRUVSLUDFOH&GXFWVRQPHVRWKRUDFLFPDUJLQ'DQWHQQD GUDZQIURPWKHVSHFLPHQ RQ3LWD\DIUXLW (SRVWHULRUVSLUDFOHZLWKGXFWVDQGFRQLFDOJODQGVSLQHVODWHUDOO\)GXFWV DQGFRQLFDOJODQGVSLQHVRQDEG,*PDUJLQRIDEG,,,YHQWUDOVXUIDFH+OHJYHVWLJHRQ PHVRWKRUD[,OHJYHVWLJHRQPHWDWKRUD[6FDOHEDUȝP$ȝP%±,

93 )LJOpuntiaspisVS4$GXOWIHPDOHLQDJURZLQJVWDJH2QGUDJRQIUXLW$DEGRPHQ%GRUVDO PDFURGXFWVRQDEG9,,&S\JLGLDOPDUJLQDEG9DQG9,GRUVDOVXUIDFH'PHGLDQDQG VHFRQGWUXOODH(PDUJLQRIDEG,,,GRUVDOVXUIDFH6FDOHEDUȝP$ȝP%±( 94