AppL enlomo). Zoo1. 42 (3): 337-346 CW07) http:.,,'odokon.org Review

Applied evolutionary ecology of of the subfamily Bruchinae (Coleoptera: Chrysomelidae)

Midori TCDA * Institute of Biological Control, Faculty of Agriculture. Kyushu Lnivcrsity: Fukuoka 812-8581, Japan (Received 17 October 2006: Accepted 19 January 2007)

Abstract Bean ofthc subl:'lmily Bruchinae (formerly. the family I3ruchidae) incilide notorious pests of stored legumes, Callosohruchus, , Acunthoscelides and ZahlVles that arc able to feed and reproduce on dried beans and peas. Here, I revIew recent findings on the ecology, phylogeny, invasion and evolution in the bean heelles, based on field in­ vestigation of host plants and molecular studies. Possible future application or the new knowledge to weed and pest control is proposed, such as potential utility of the predators for modest control of beneficial yet invasive ('con­ flict') plants and new control methodology ofpest bean beetles.

Key words: Bruchidae: seed predators; stored product pests; biological weed control: molecular phylogeny

stored-bean pests. Strictly speaking, however, the !;\fTRODUCTlON genus Bmchus is different from other stored-bean Beetles of the subfamily Uruchinae (Coleoptera: pests in that it does not reproduce on dried hard­ Chrysomelidae) are specialized internal feeders of ened beans/peas and thus cannot cause further bean family (Fabaeeae). They are frequently damage in storage. and erroneously referred to as the bean 'wecvils' in spite of their aITmity to the leaf beetles PHYLOGENY (Chrysomclidae). Here, I usc the common name, the bean '' (or the seed 'beetle') to avoid tax­ Recent molecular studies support that the onomic confusion. I also follow the recent taxo­ Bruehinae are closely related to the frog-legged nomic consideration that the bean beetle is not beetle subfamily. Sagrinae, within the family unique enough as a famJiy but as a subfamily of Chrysomelidac (Farrell and Sequeira, 2(04). Chrysomelidae (Lingafelter and Pakaluk, 1997). Within the Bruehinae, the tribes Amblyeenlll and The Bruehinae consist of about 1.700 species Paehymerini are thought to be paraphyletic to a (Johnson et aL 2004). Although not very high in major tribe Bruchini (Farrell, 1998; Table 1). Al­ proportion of species, some of the bean beetles arc though recent evidence supports the previously notonous pests of stored beans \vith the highest in­ proposed phylogeny at the higher taxonomic levels, trinsic rate of increase among stored-products pests an intriguing nev·/ view of bean beetle phylogeny at Omura, 1990). They include several species of Cal­ the lower levels has been proposed, Kergoat et al. losobruchus, AC{/nihoscefides and Zabmtes that in­ (2005a, b) supported the previously suggested view fest beans (Tribe Phaseoleae: Vigna, Phmeolu\', that and 3ruchidills arc poly­ G(vcine, etc.) and Car:v'edon that feeds on peanuts phyletic groups (Johnson, 1981; Borowiec, 1987). or groundnuts (Arachis h}~pogaea) (Southgate, However, some monophyletic groups of Acan­ 1979; Johnson, 1981). Bruchlls species that infest thoscelides may be more closely related to those of broad beans and peas (Fabeae: Vicia and Pisum) of Hmchidills than the other congeneric groups (Al­ economic importance may also be categorized as varez et aI., 2006; Tuda, Kergoat, Kato, lto, Ru-

* F-mail: tuda(i;;gn.kyu.hll-ll.ac.jp DOl: IO.1303aez.2007.3.'\7 338 M. TL'DA

Table I. Taxonomic position and composition of Hruchinac (Balachowsky, 1962; Tuda, pers. obs.). In both cases, the host seeds arc too small to support bean Family beetle development into full adult. Finally. as an Suhfamily Trihe exception to the general pattern of pupating within seeds, the larvae of several species of Caryedon Chrysomelidae emerge out of seeds to form cocoons either on Bruehinae pods or on the ground for pupation (Center and Amblieerini 3 genera (5permopllilglls, Zahmtes) Johnson, 1974; Southgate, 1979; Tuda, pers. obs.). Rruehini 46 genera (Aeall1hoseelidl'.,', . Batch egg laying is knmvn only for a limited Bruehl/.I. Callfl,\(Jbrf/chl/l'. ,Hilllosesres) Eubaptini-l genus (tuha{!ms) number of species such as Pseudopachymerina Kylorhinini-l genus (KworhiI1I1S) spimj}es (Teran, 1962), Cwyedon jasciatus (Pre­ Paehymerini-12 genera (Caryed(Jnj vett, 1966). Cafyoborus serripes (Delobel et al., Rhaebini-l genus (Rhaehlls) 1995<:l) and Stator beali U

(sensu Jenny, 1984) or sequential radiation (sensu ture studies. hardness of dried seeds will be one of Abrahamson et al., 2(03) that follows plant diversi­ the foci ofresearch because the hardness per se can fication, \'.'hieh contrasts to the reciprocal coevolu­ serve as a deterrent against seed predators, includ­ tion model that assumes plant evolution in re­ ing bean beetles (Janzen, 1977; Southgate, 1979; sponse to lOsect evolution against plant defensive Kitch et aJ.. 1991; Dongre et al., 1993). The loss of traits (sensu Ehrlich and Raven, 1964: Futuyma, toxic chemicals during post-maturity drying 1983; Becerra, 2003). processes may also increase survival of such grani­ vores, \vhich promotes host range expansion. EVOLUTION OF STORED PRODUCT PESTS INVASION Evolution of stored bean pests IS likely a t\\'o­ step process. The first is preadaptation to utilize With human aid, dispersal and invasion of living dried hard seeds (Watanabe, 1985) and the second organisms in recent centuries are occurring at a is the evolution ofdispersal, reproductive and com­ speed that has been unattainable v,'ith plate tecton­ petitive polymorphism to adapt to cultivated seeds ics and glacial dynamics. In Japan, examples of and storage environments (Utida, 1954, 1981; bean beetle invaders arc Bruchus I"/!jiman/ls and Cas\vell, 1960; "Nakamurd, 1966; Sano, 1967: Oue­ Bruchus pisanl/n, the pests of broad beans and draogo and Huignard, 19H 1; Messina, 1984: Toque­ peas. respectively (Table 2). Bruchus !oti has also naga, 1990: Tuda, 1997, 1998: Tuda and Iwasa, been considered to be an aiJen species but our re­ 1998; Takano et aI., 2(01). lIenee, an ability to cent fmding indicates the current status as an intro­ feed on dried mature host seeds IS an inevitable duced species may require reconsideration, as de­ prerequisite for bruchids to become pests. This re­ scribed later in this section. We found two addi­ quires first, female adults to deposit eggs on/near tional bean beetle species havc invaded Japan dried mature seeds and second, hatched larvae to (Tuda et a1.. 2001; Tuda, Tateishi, Niyomdham. burrow into the hard seeds to feed on. Monmoto. Chen, Zhu. Zhang. t\lurugan, Chou and What is it then that promotes evolution of uti­ Johnson, unpublished). lization of dry, hard seeds? Using a comparative One recent invader is a bean beetlc Acan­ approach to study Ca!losohruchus pest and non­ Ihoscelides pallidipennis (=.4. collusus) that feeds pest species, the potential effects of climate, host on a fast-growing. nitrogen-fixing legume, In­ plants, phylogeny and endosymbiotie bacterium digobush (or False indigo; ltaehi-hagi). Amol"pha were examined (Tuda et aL 2(06). Long dry sea­ fruticosa. The legume had invaded from North son was shown to be the most important factor for America and established in England, Europe and evolution towards preadaptive stage to become East Asia except Japan by the mid 20th century stored bean pests (i.e., ability to use dry hard (Szentesi. 1999: Tuda et al.. 20(1). Tn Japan. the beans) when evolutionary history \vas accounted bean beetle \vas first found at two locations in the for (Tuda et aI., 2006). Phylogenetic history of Caf­ central and southwestern parts of the country ltl losobruchus and idiosyncrasy of their host plants 1997 (Tuda et al., 2001) and later in several areas also had significant effects on the evolution to­ in the southwestern Japan as well (Ishihara, unpub­ wards stored bean pests (Tuda et aI., 2006). Tn fu- lished). A molecular analysis of A. pallidipennis

Table 1. Bean beetle invasion to Japan

Bean beetle Time lIfinvasion Source population

Srue/HIS pi.l'orllm' 1888 l;SA Bruchus TI(fimunlls" 1921 England/Europe Cal/osohnlchlls macil/alush 1950 o .1uillrho,lcc!ides pal/idipei/llis' 1<)70s 199i Korea and China .'Icanlho.\'ci'lides iIIlicrophthafmu.\d recent ( 2000)

'Yoshida, 1990; "'v!orimoto and Kiritani. t995. 'Tuda el al.. 2001. dTuda. Tateishi, "\iiyomdham. Morimoto. Chen. Zhu. Zhang. Murugan, Chou and Johnson, unpllbli~hcd. 340 M_ TLn..\ from four States of the USA and from East Asia re­ lotypes found in the Japanese populations are vealed that Japanese populations consist of several dearly different from those in European popula­ unrelated haplotypes that are shared by Chinese tions. which mdlcates that Japanese B. loti is not and Korean populations and individuals intercepted introduced from Europe and llkely originated from by import inspection at Japanese seaport quaran­ the Far East including Japan (Tuda. lchita and Ker­ tines (Tuda, Ishihara, Wasano, Morimoto, Paik, goat, unpublished). The genetic uniqueness of the Houck and Johnson, unpublished). This molecular Japanese population mdicates that B. loti should cv'idcnce supports our hypothesis (Tuda et aI., perhaps be consen-ed m tillS country. The status of 2001) thal A pallidipennis established in Japan Japanese Callosobmehus chillensis is also contro­ was transported from the introduced populations in versial. Our molecular study shO\vs that popula­ nearby East Asian countries. Indeed, from these re­ tions of C chinel1sis m agricultural habitats are ge­ gions, the host seeds (A. jhaicosa) have been im­ netically homogeneous. \\-hereas those from natural ported for erosion control since the 1970s (Tuda ct habitats arc heterogeneous. indicating a bottleneck aI., 2001.), e'ient caused by human agricultural practice and The other invader is Acanthoseelides maeroph­ transportation (Tuda el a1.. 2004). thalmus (Tuda, Tateishi, Niyomdham, Morimoto, Chen, Zhu, Zhang, Murugan, Chou and Johnson, APPLICATlO" TO BIOLOGICAL CONTROL unpublished). It is a Neotropical species that feeds on Wild tamarind, or Gin-nemu, Lel/eaClla leucu­ As natural cncm~- of conftict plants. A control cephala. L('ueaena leucocephala is a fast-growing agent should ideally be a specialist feeding only on nitrogen-fixing leguminous tree that is cultivated targets of control and not harm beneficial organ­ for fodder (Elharith ct a1., 1980), grccn manure isms (Sweetman, 1936: Huffaker, 1964). Internal (Chagas, 1981), rcforestation, \vindbreak, fucl, feedcrs ('endophages') tend to have narrower diet pulp and erosion control (Kondo et aI., 1987: Sa­ breadth than external feeders Cexophages') take ct al., 1989). Because A. macrophthalllJlIs is (Lewinsohn, 1991: Frenzel and Brandl, 1998) and specialized to Lmcaenu species (e.g., Johnson, are probably more suitable as biological control 1979: Hughes and Johnson, 1996; Delobel and agents. Indeed, bean beetles have been used as bio­ Johnson, 1998), L. leucocephalu is the only Leu­ logical control agents of weedy plants and proven caCl1a species that has been introduced to Japan, to be effective, to some extent, once established in and airborne long-distance dispersal is highly un­ the release areas (Table 3: c.L Julien, 1992). likely for bean beetles (Tuda et al., 2m)]), there is 1propose tv,'O bean beetle species that have been little doubt that A. mucrophthalmus has been intro­ accidentally introduced and can be used for biolog­ duced with the host seeds. ical control of new leguminous weeds in Japan. Accelerated long-range human transportation The first is A. pallidipel1nis. The narrO\v diet range promotes unexpected dispcrsal of and of A. pallidipennis and the absence of congeneric plants that eventually become pests in the areas relatives of lndigobush and naturally associated where they are introduced. As a consequence, it is parasitoids ofthe beetle arc preferable features as a sometimes difficult to dctermine geographic ori­ control agent, in contrast \vith the case of Bruchid­ gins of widcly distributed organisms. Whcther ius vi!losus that attacked not only the target but local populations arc native or of relatively rcccnt also non-target (but non-indigenous) congeners introduction from foreign populations can be esti­ after its introduction to '\Jew Zealand (Paynter et mated by statistics bascd on population genetics aI., 2004). In this respect, A pallidipennis is suit­ theory if Dr\A sequcnce or restriction site data arc able as an agent that controls escape of AmOlpha available (c.g.. Templeton et a1., 1992; Tcmpleton, fruticosa from cultivation by seed dispersal. The 1998). Our recent fmdings regarding Japanese pop­ suitability of A. pallidipennis as a control agent has ulations of Bruchus lori may be a good example. also been pointed out in the native distribution area Palearctic fauna and flora share many species in­ of Indigobush (Rogers and Garrison, 1975). The cluding B. loti. The Japanese populations of B. loti bean beetle would be even more suitable if it would are currently listed as an alien species that would be applied for control in introduced areas \vhere become a target of eradication. We found that hap- its naturally associated enemies (i.e., parasitoids, Applied evolutionary Ecology of Bruchinae 341

Tahle 3. Bean heetles used as biological control agents of weeds (modified from Julien. 1992: ARC-I'I'RI. 20(3)

Control agent Target weed Location of release

Aeanthosce!ide.l· macrophlhalmus Leueaena leucocepha/a South Africa f1canthosce!idcs puniceus Mimosa pigru Aus1ralia. Thailand, Vielnam (Malaysia. lvlyanmilr) Aeanthoscefides quadridentmu,I' ....limosa pigru Australia. Thailand, Vietnam (rvlitlaysia, Myantllilf.lJSA) /1lgarohius ho/limeri Prnsopis glanda/osa South Africa AfgarohiuY prosopis F'rosopis vetil/ina South Ali-iea Bnlchidius vilfosus CHis'us scopariu.\' 1\ew Zealand BrllchidiliS sahlhergi Acacia nifotiea Australia Slllcohl"1lchu.l.lubsll/llmlis Caesa/pillia drxaperala South Africa

Locations in parelltheses are due 10 naturdl spread ofreleased agents.

predators, parasites, and pathogens) and competi­ e.g.. Southgate, 1979; Steffan, 1981; Fujii and Wai, tors (e.g., Acanthoscelides slIbmulicus) are rare or 1990; Mitsunaga and Fujii, ]999; Schmale et aI., absent (Tuda et aI., 200 I). Acanthoscefides pal­ 2001; Tuda et a1., 2001; Kobayashi et al., 2003; IIdipenni.\" feeds only on Amorpha and related gen­ Jaloux et aI., 2004; Tuda and Shimada, 2005; Wu era (Amorpheae: Erra=llrizia and Parryella) that et al., 2005; Vamosi, Hollander and Tuda, unpub­ distribute naturally only in North America (Center lished) and plant extracts (e.g., Lambert, 1985; and Johnson, 1974). Rahman, ]990) have been actively explored for bi­ The second is A. macruphthalmus. Lellcaena ological control (sec Fujii et £11.,1990; Huis, 1991 leucocephala, initially introduced as a beneficial for review). In addition to biological and botanical tree in the 19th century, escaped from cultivation control, controlled temperature (e.g., Rahman, by seed dispersal and has become weedy in tropical 1990). mechanical control (e.g., Quentin et a1.. regions of Japan and other introduced areas (e.g., 1991), controlled atmosphere (e.g., Oosthuzien and Smith, 1985; Henderson, 2001; Wu et a1., 2003). Schmidth, 1942) and radiation (e.g., Kiyoku and Its seed predator, A. macruphlhalmu.\", satisfies the Tsukuda, 196R; Hossain et aI., ]972; Reddy et aI., above criteria as a control agent in Asia, i.e., nar­ 20(6) have been proposed as effective control row host range and scarcity of parasitoids. In fact, methodology (see also Highley et a1., 1994). 1 sug­ this bean beetle has already been deliberately intro­ gest an endosymbiotic bacterium, H/olhachia, front duced to South Africa for the control of L !ellco­ bean beetle pests and its functional relation with cephala (ARC-PPR1, 2003; Olckers, 2004; Tabl~ the host (Kondo et a1., 20(2) may be applied to bi­ 3). Nevertheless, there remains a flsk that the con­ ological control of stored product pests. When the trol agent could easily accumulate local generalist above-mentioned multiple methods arc used III parasitoids and in the long term the control ~frect combination, the possibility of one agent negating could decrease as observed in West Africa (Delo­ the other must be taken into account (e.g.. Boeke et bel and Johnson, 1998). aI., 200]). Control of bean beetles. Fumigation has been Identification of pest species. External and in­ widely applied to stored products as an effective ternal morphology of the adult stage is used for method of control of stored products pests. How­ identification to species. Most recently male g~nital ever, methyl bromide, used for fumigation of gram traits for were compared and sum­ legumes (Yoneda et al., 1990) has been recognized marized by Tuda et al. (2006). Compared to adult as one of the chemicals that deplete the stratos­ morphology, egg and larval morphology is less pheric ozone layer and hence, was called for its well studied because they arc not readily available controlled use by the Montreal Protocol (U~[P. (but see Pfaffenberger and Johnson, 1976; Arora, 2000). 197R; Delobel et a1., 1995b), \vhich makes identifi­ Alternatively, parasitoids (e.g., a trichogram­ cation based on pre-adult stages difficult. Tn con­ matid egg parasitoid U~cana, larval-pupal para­ trast to the differential accessibility among devel­ sitoids, pteromalid Dinarmus, Anisopteromafus, an opmental stages of morphological characters, mo­ eupe1mid Eupelmus, and a braconid Heterospillls; lecular characters can be more stable and useful a.s 342 \1. ltD,,,

keys for identification. Using the PCR-RFLP tech­ 44: 63 74. nique. which is relatively easy and inexpensive, Amon, K.-W (1999) Rcvision of the genus Sulcohl"lIchul' species of all developmental stages of important Chujo 1937 and description of ParaslIlcobrllchllS nov, gcn, (Coleoplera. Bru~hidae. Bruchinae). I.in:, Rio/. stored bean pests in the genus Callosohruchus have Beirr. 31: 629-650. been successfully distinguished (Tuda et a1., 1995). Anton. K.-W. (2000) riv,; ncw spccics of the Cu/losohruchlls chinensi,\' group from thc Oricntal Region and Australia (Coleoptera: Aruchida,;: Bruchinae). Genus 11. 13-28. AnIon. K.-\V, J. lIalp';rJn and 1\.1 Caldcron (1997) An anno­ Bean beetles. as other phytophagous insects, ex­ taled list ofthe Aruchidac (Co!,;optera) of Israel and adja­ ccnt arcas. Israel 1 Enllil/wl. 31: 59-96, hibit significant conservatism in host utilization. ARC-PPRI (2003) Releases (!/ Biological CO/ltrol Agents Feeding on dried seeds not only enables repeated againsr If-ceds in Soulh A/i'im, http:/'155.240.199.39, generations but also broadens the diet of some inslitules/ppriimain/divisiolls/w,;cdsdiy/releases.htm. bean beetle pests. Our study based on molecular Arora. G. L. 11'177) of Aruchida,; (Coleoptera) of phylogenetics indicates this preadaptation to use Korthwest India. ParI 1. Adults. Orienl. InseC! Slippl, 7: I 132. dry. hard beans that precedes human store of beans Arora. G. L. (1978) Taxonomy or Bruchidac (Co!coptiOra) of has been selected for by arid habitat climate (Tuda "Iorthwest India, Pan 11. Larvac. Orielll, InseC! Supp!. et al., 2006). Recent investig':ltions of Asian bean S: 1·48. beetle fauna have revealed diversity of the species Arora. G, L. (1(80) A Stud\' o/Ihe Biolog.l anJ Tiuonomy of and ecology of the seed predators. This also led to Ihe Genlls Bruchidius (Co!eoplera. Bl"uchidaej from the finding of invasions of alien bean beetles (Tuda India, filial Tech. Rep' (/974 1(79) [.is. PL-4(10 Res. Proj. AJ·L,Vj:J(!3. Department ofZoology. Punjab Uni­ et a\., 2001; Tuda et al., unpublished). Invasion vcrsity. Chandigarh. '16 pp, routes estimated by our molecular approach indi­ Balachowsky. A. S. (1962) t:ntomo!ogie Appliquee a !"Agri­ cated the invasion of the bean beetle was associ­ ell/wre Tome I. Col(optel"es. Vol. I. Masson ct Cic ated with imported alien host seeds. 1 propose the fdileurs. Paris. 564 pp. reeent bean beetle invaders may be used for modest Bc,;crra. J. X, (2003) Synchronous coadaptation ltl an an­ cienl case of herbivory. Proc. ,Io-,'al!, Acud. Sci. ['SA 100 control of the sced dispersal of beneficial but inva­ 12804-12X07, sive alien plants ('conflict plants' (Neser, 1994)). Boeke, S. 1.. A, ,1\. C Sinzogan, R. P. de Almeida. P. \V. M. dc Boer. G. Jeong. D. K. Kossou and J. J. A. van Loon ACK]\"OWLEOCE.\IENTS (200]) Sidc-cffe,;ts of treatmem wilh hotanical ! am gratdul to the Japanese Society of Applied fntonlOl­ insecticides on two parasilOids of Cal/o\'ohrlli.'hlls IlJaClI­ ogy and Zoology for the honorable award and the opportunity IU/lis. t:IJ£O/I1O/. F.xp. App/. lOS: 43-51. (I~n) to writiO this r,;vi,;w, Thanks are also due to Alex Delobel and Borowiec, L. II The gcnera of sccd-bc,;t1cs (Coleoptera, Sleven Vamosi for their valuable ,;ommcnts on lhe manuscript Bruchidae), Po/sk. Pismo t:I1[O/no/' 57: 3 207, Ihis sludy was supported partly by thc FUJiwara :'-Jatural Caswell. G. H, (1960) Observations on an ahnOnllal form of History Foundation. lhe Sumitomo Foundation. and Grant-in­ Caffosobruchus mamlalus (E). Bufl. t:nromo1. Res. 50: Aids for 1nlernalional Scientific Rescarch (ficld R,;scarch 671-6S0, 09041145) and for Scientific Rcscarch (A) (08304049, Center. T D. and C D. Johnson (1972) Comparative life his- 15208007). (13) (14405003,17405005) and for 'roung Scien­ torics ofSenniu.\ Em·lm/!. t:II/0/110/. 2: 669 672. Ccnter, T. D, and C. D. Johnson (1974) COiOvolution of some lists (I:l) (15770011) from MEXT. This paper is dedicated [0 my collahorators, the late Prof Shwu-Bin Horng for his work sccd bcetles (Coleoptera: 13ruchidal;) and thcir hosts, on behavioral ecology of Ca/losohmcflUs and the late Dt, Fc%gy 55: 1096-1103. l.iang-Yih Chou for his conlribution to applil.:d cntomology. Chagas, J M. ( 1981) LClfcuena Icucocepha/a as a green ma­ nure for bean growing in c,;rrado soil. Pesq. Agl: Bras. REFEREI\CES 16: 809-S14. Delgado. c.. G. Couturier and A. Dclobcl (1997) Oviposi­ Abrahamson, W. G., C. l' Blair, M. !l Eubanks and S. A. tion of seed-beelle Caryohor!ls .)erripes (Sturm) (Co­ Morehead (20lH) Sequential radiation of unrdatcd or­ leoptera: Bruchidae) on palm (As[rocaryum cham· ganisms: lhe gall lly L'urosra solidaginis and th,; tumbling bira) fruils under natural conditions in P,;ru, AliI/. Soc. flow,;r bcctle Jfol"de//islcnu cm/victu. J. Frol. Bioi, 16: Elllol/Jo!. Frail. 33: 405--409. 781-789. DelobeL A. and I:l. Delobl.:1 (2003) L,;s plantes holes des Alvarez.1\., 1. R, Napoles. K.-\V. ,I\nton. B. Bemey and \1 bruches (Colcoptcra: Bruchidac) de la [aune de hance. Hossaert-!'v1cKey (2006) Phylogcnctic rdationships in une analyse critiquc. Rull. ,Hens, Soc Linn, Lm!! 72: Ih,; Kcotropical bruchid genus Acanthosce/ides (Aruchi­ 199221. nae. Aruchidac. Coleoptera). J Lool. .\1'\'1. Emf Res. Delohl;l. A. and C. D, Johnson (1998) first record of a sccd- /\pplied Evolutionary Ecology ofBruchinae 343

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