JapaneseJapaneseSociety Society of AppliedApplied Entomology and Zoology

AppL Entomol. ZooL 42 (3): 337-346 (2007) http/]todokon.orgl Review

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

Midori TuDA"

Institute ofBiological Centrol, Faculty of Agriculturc, Kyushu Univcrsity; Fukuoka 812-8581, Japan

(Reccis,ed 17 October 2006; Accepted 19 January 2007)

Abstract

Bean ofthe subfamily Bruchinae (formerly, the family Bruchidae) include notorious pests of stored legumes, Callosobrttchus, CaEyedon, Aeanthoscetides and Zdbrotes that are able to feed and reproduce on dricd bcans and peas. Here, I review recent findings on the ecology, phylogeny, invasion and evolution in the bean beetles, based on field in- vestigation of host plants and rnolecular studies. Possible future application ofthe new knowledge to weed and pest control is proposea such as potential utility of the predators for modest control of beneficial yet invasive (`con- flict') plants and new control methodology ofpcst bcan bcctlcs.

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

stored-bean Strictly speaking, however, the INTRODUCTION pests. genus is different from other stored-bean Beetles ofthe subfamily Bruchinae (Coleoptera:pests in that it does not reproduce on dried hard- Chrysomelidae) are specialized internal feeders of ened beanslpeas and thus cannot cause further bean family (Fabaceae). They are fi:equently damage in storage. `' and erroneously referred to as the bean

in spite of their aMnity to the leaf beetles PHYLOGENY (Chrysomelidae), Here, I use the common name, `' `beetle') the bean (or the seed to avoid tax- Recent molecular studies support that the onomic confusion, I also fo11ow the recent taxo- Bruchinae are closely related to the fi;og-legged

nomic consideration that the bean beetle is not beetle subfamily, Sagrinae, within the family unique enough as a family but as a subfamily of Chrysomelidae (Farre}1 and Sequeira, 2004). Chrysomelidae (Lingafelter and Pakaluk, 1997). Within the Bruchinae, the tribes Amblycerini and The Bruchinae consist of about 1,700 species Pachymerini 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 are though recent evidence supports the previously notorious pests of stored beans with the highest in- proposed phylogeny at the higher taxonomic levels, trinsic rate ofincrease among stored-products pests an intriguing new view of bean beetle phylogeny at (Imura, 1990). They include several species of Cat- the lower levels has been proposed. Kergoat et al. losohruchus, Acanthoscelicles and Zkebrotes that in- (2005a, b) supported the previously suggested view fest beans (Tribe Phaseoleae: Vigna, Phaseolus, that and are poly- Clycine, etc,) and Ckeilyedon that feeds on peanuts phyietic groups (Jehnson, 1981; Borowiec, 1987). or groundnuts (AJuchis Itypagaea) (Southgate,However, some monophyletic groups of Acan- 1979; Johnson, 1981). Bruchus species that infest thoscelides may be more closely related to those of broad beans and peas (Fabeae: Vicia and Pisum) of Bruchidius than the othcr congencric groups (Al- economic importance may also be categorized as varez et al., 2006; TIJda, Kergoat, Kato, Ito, Bu-

"E-mail]tuda@grt,kyLishu-u,ac.jp DOI/IO.13031acz.2007.337

337

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Table 1. Tlaxonomicposition and eompositionol'Bruchinae (BalachowskM 1962; [[Uda, pers. obs.). In both cases, the host seeds are too small to support bean Family beetle development into fu11 adult. FinallM as an Subfamily exception to the ofpupating within Tribe generalpattern seeds, the larvae of several species of CaFyedon

Chrysomelidae emerge out of seeds to form cocoons either on Bruchinae pods or on the ground fbr pupation (Center and Amblicerini-3 genera (fpermophagtts, Zbbrotes) Johnson, 1974; Southgate, 1979; IUda, pers, obs,). Bruchini--46 genera (Aeanthoseelides, Bruchiditts, Batch egg laying is known only fbr a limited Bruchus, (;Zillesobruchus, il4imosestes) number of species such as Rseudopacbymerina Eubaptini-1 genus

et al,, 2007a,b), although there are a few ECOLOGY general- ists that utilize different subfamilies of legumes. ln most bean beetles, female adults deposit eggs Evolutionary relation between bean beetles and en pods andlor seeds, hatched larvae burrow into their host plants has been actively discussed as a seeds and emerge out as adults (Southgate, 1979). model ofplant- evoLution, Proximate mecha- Adults lay eggs singly and larvae feed only single nisms for host-plant shifts are probably associated seeds (Center and Johnson, 1974). Exceptions to with chemical (Janzen, 1969), morphological the typical life history are Conicobrttchus (Prevett,(Janzen, 1969; Szentesi and Jermy, 1995) and geo- 1967), A,ferobruchus (Johnson, 1967) and Sbnnius graphical (Johnson and Siemens, 1991) proximity (Center and Johnson, 1972), which feed on several between the current and potential hosts. Delobel seeds in a single pod and Bruchidius, which feeds and De]obel (2006) suggest that the speciation in on multiple pods in single 7}'ijblium inflorescenccs European bean beetles is sequential evolution

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Applied Evolutionary Ecology ofBruchinae 339

(sensu Jermy, 1984) Qr sequential radiation (sensu ture studies, hardness ofdried seeds wM be one of Abrahamson et al., 2003) that fo11ows plant diversi- the fbci ofresearch because the hardness per se can fication, which contrasts to thc 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 insect evolution against plant defensive Kitch et al., 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, which promotes host range expansien.

EVOLUTION OF STORED PRODUCT PESTS INYASION Evolution of stored bean pests is likely a two- step process. The first is preadaptation to utilize With human aid, dispersal and invasion ofliving dried hard seeds (Watanabe, 1985) and the second organisms in recent centuries are occurring at a is the evolution of dispersal, reproductive and com- speed that has bcen unattainable with plate tecton- petitive polymorphism to adapt to cultivated seeds ics and glacial dynamics. In Japan, examples of and storage environments (Utida, 1954, 1981; bcan beetle invaders are Bruchus ntfimanus and Caswell, 1960; Nakamura, l966; Sano, 1967; Oue- , the pests of broad beans and draogo and Huignard 1981; Messina, 1984; Toque- peas, respectively (Tlable 2). Bruchtis loti has also naga, 1990; [IUda, 1997, 1998; [fuda and Iwasa, been considered to be an alien species but our rc-

1998; fakano et al., 2001). Hence, an ability to cent finding indicates the current status as an intro-

feed on dried mature host seeds is an inevitable duced species may require reconsideration, as de- prerequisite fbr bruchids to become pests. This re- scribed later in this section, We fbund two addi- quires first, female adults to deposit eggs onfnear tional bean beetle species have invaded Japan dried mature seeds and seconct hatched larvae to (Tuda et al., 2001; [IUda, [Ibteishi, Niyemdham, burrow into the hard seeds to feed on, Morimoto, Chen, Zhu, Zhang, Murugan, Chou and What is it then that promotes evolution of uti- Johnson, unpublished). Iization of dry, hard seeds? Using a comparative One recent invader is a bean beetle Ac'an- approach to study pest and non- thoscelides pallidipennis (=A. coUusus) that feeds pest species, the potential effects of climate, host on a fast-growing, nitrogen-fixing legume, In- plants, phylogeny and endosymbiotic bacterium digobush (or False indigo; Itachi-hagi), Amorpha were examined ([[Uda et al,, 2006), Long dry sea- .fruticosa, The legume had invaded from North son was shown to be the most important factor fbr 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., 2001). In Japan, the beans) when evolutionary history was accountcd bean beetle was first found at two locations in the for ([[lida et al., 2006). Phylogenetic history of Cal- central and southwestern parts of the country in tosobruchus and idiosyncrasy of their hQst plants 1997 (TUda et al., 2001) and later in several areas also had significant effects on the cvolution to- in the southwestern Japan as well (Ishihara, unpub- wards stored bean pests (TUda et al., 2e06). In fu- lished). A molecular analysis of A. pallidipennis

Table 2. Bean bectlc invasion to Japan

Bean beetle Time of invasion Sourcepopulation

Bruchuspisorum" 1888192119501970s-1997 USAEngtand,'Europe BruchusrtLfimanus" Callosobruchus tnaculatttsb ?Korea

A['aiithoscetidespaUidipennist and China Acanthoscelidesmacrophthalmttsd reeent (-2000) ?

a b C d lybshida, 199e; Morimoto and Kiritani, Ttida et al., 2001; Tl]da, Tlateishi, Niyomdham, Morimoto, Chen, Zhu, Zhang,

Murugan, Chou and Johnson, unpublished.1995;

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from fbur States of the USA and from East Asia re- lotypes fbund in the Japanese populations are vealed that Japanese populatiens consist of several clearly different from those in European popula- unrelated haplotypes that are sharcd by Chinese tions, which indicates that Japanese B. toti is not and Korean populations and individuals intercepted introduced from Europe and likcly originated from by import inspection at Japanese seaport quaran- the Far East including Japan (Tuda, Ichita and Ker- tines (Tbda, Ishihara, Wasano, Morimoto, Paik, goat, unpublished). The genetic uniqueness of the Houck and Johnson, unpublished). This molecular Japancsc population indicates that B. Ioti should evidence supports our hypothesis (Tuda et al., perhaps be conserved in this country. The status of 2001) that A. pallidipennis establishcd in Japan Japanese Caltosobrt{chtts chinensis is also contro- was transported from the introduced populations in versial. Our molecular study shows that popula- nearby East Asian countries. Indeed from these re- tions of C. chinensis in agricultural habitats are ge-

the host seeds have been im- netically homogeneous, whereas those from natural gions, (A..fi'uticosa) ported for erosion eontrol since the 1970s (TUda et habitats are heterogeneous, indicating a bottleneck aL,2001). event caused by human agricultural practice and The other invader is Acanthoscetides macroph- transportation (Tuda et al., 2004). thalmus (TUda, Tateishi, Niyomdham, Morimoto, Chen, Zhu, Zhang, Murugan, Chou and Johnson, APPLICMION TO BIOLOGICAL CONTROL unpublished). It is a Neotropical species that feeds on Wild tamarind or Gin-nemu, Leucaena teuco- As natural enemy of conflict plants. A control cephala. Leucaena teucocephata is a fast-growing agent should ideally be a spccialist feeding only on

nitrogen-fixing leguminous tree that is cultivated targets of control and not harm beneficial organ- fbr fbdder (Elharith et al., 1980), green manure isms (Sweetman, 1936; Huffaker, 1964). Internal (Chagas, 1981), refbrestation, windbreak, fuel, feeders (`endophages') tcnd to have narrower diet pulp and erosion control (Kondo et al,, 1987; Sa- brcadth than external feeders (`exophages') take et al,, 1989). Because A. mactnphthalmus is (Lewinsohn, 1991; Frenzel and Brandl, 1998) and specialized to Leucaena species (e.g., Johnson, are prebably more suitable as biological control 1979; Hughes and Jehnson, 1996; Delobel and agents. Indeeq bean beetles have been used as bio- Johnson, 1998), L, leucocephala is the only Leu- logical cQntrol agents of weedy plants and proven caena 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 ([IUble 3; c.f, Julien, 1992). likely for bean beetles ([[Uda et al., 2001), there is I propose two bean beetle species that have been little doubt that A, macmphthalmus has been intro- accidentally introduced and can be used fbr biolog- duced with the host seeds. ical control of new leguminous weeds in Japan. Accelerated long-range human transportation The first is A. patlidipennis, The narrow diet range promotes unexpected dispersal of and ofA. pallidipennis and the absence of congeneric plants that eventually become pests in the areas relatives of lndigobush and naturally associated where they are introduced. Asa consequence, it is parasitoids ofthe beetle are preferable features as a sometimes dirncult to determine geographic ori- control agent, in contrast with the case ofBruchid- gins of widely distributed erganisms. Whether itts vitlosus that attacked not only the target but local populations are native or of relatively recent also non-target (but non-indigenous) congeners introduction from fbreign populations can be esti- after its introduction to New Zealand (Paynter et mated by statistics based on population genetics al., 2004). In this respect, A. paUidipennis is suit- theory if DNA sequence or restriction site data are able as an agent that controls escape ofAmorpha rlempleton available (e.g., et al., 1992; 1fempleton, .fruticosa from cultivation by seed dispersal, The 1998). Our recent findings regarding Japanese pop- suitability ofA. pallidipennis as a control agent has ulations of Bruchtts loti may be a good example, also been pointed out in the native distribution area Palcarctic fauna and fiora share many species in- of Indigobush (Rogers and Garrison, 1975). The cluding B. Ioti. The Japanese populations ofB. Ioti bean beetle would be even more suitable if it would arc currcntly listed as an alien species that would be applied fbr control in introduced areas where become a target of eradication. We fbund that hap- its naturally associated enemies (i.e,, parasitoids,

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2003) Tbble3. Bean beetles used as biological control agents ofweeds (modified from Julien, 1992; ARC-PPRI,

Control agent Tlarget weed Locationofrelease

AcanthosceiidesmacFephthalmus Lettcaenaiet{cocqphala South Africa Acanthoscetidespttniceus Mimosapigtu Australia, Thailana Vietnam (Malaysia, Myanmar) Acanthoscelidesquadridentatus Mimosapigrn Australia, Thailanct Vietnam (Malaysia, Myanmar, USA)

Algaivbittsbottinieri Prosopisglandttlosa South At'rica

Aigarohiusproscpis Pivsopisvetutina South Africa Brttchidittsvillosus C),tisusscoparius New Zealand Biitchidiussahlbeigi Acacia nitotica AustraliaSouth

Sulcobruchussubsuttttutis Caesaipinia deeapetala Africa

Lecations in parentheses are due to natural spread ofre]eased agents.

predators, parasites, and pathogens) and competi- c.g., Southgate, l979; Steffan, 1981; Fojii and Wai, tors (e.g., Acanthoscelides submutieus) are rare or 1990; Mitsunaga and Fejii, 1999; Schmale et al., absent (IUda et al., 2001). Acanthoseeiides pal- 2001; Tuda et al,, 2001; Kobayashi et al., 2003; lidipennis feeds only on Amorpha and rclatcd gen- Jaloux et al., 2004; Tuda and Shimada, 2005; Wu era (Amorpheae: Erruzt{rizia and thrFlyella) 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, 1990) have been actively explored fbr bi- The second is A. macJQphthalnius. Leucaena olegical control (see Fojii et al., 199e; Huis, 1991 leucocephala, initially introduced as a beneficial for review). In addition to biological and botanical tree in the 19th centurM escaped from cultjvation control, controlled temperature (e.g., Rahman, by seed dispersal and has become weedy in tropical 1990), mechanical control (e,g., Quentin et al., regions of Japan and other introduced areas (e,g,,1991), controlled atmosphere (e.g., Oosthuzien and Smith, 1985; Henderson, 2001; Wu et al., 2003). Schmidth, 1942) and radiation (e.g., Kiyoku and Its seed predator, A. macmphthalmtts, satisfies the Tsukuda, 1968; Hossain et al., 1972; Reddy et al., above criteria as a control agent in Asia, i.e., nar- 2006) have been proposed as effbctive control row host range and scarcity of parasitoids. In fact, methodology (see also Highley et al., 1994). I sug- this bean beetle has already been deliberately intro- gest an endosymbiotic baeterium, Pfolbachia, from duced to South Africa for the control ofL. Ieuco- bean beetle pests and its functional relation with cephala (ARC-PPRI, 2003; Olckers, 2004; Table the host (Kondo et al,, 2002) may be applied to bi- 3), Nevertheless, there remains a risk that the con- ological control of stored product pests. When the trol agent couid easily accumulate local generalist above-mentioned multiple mcthods are used in parasitoids and in the long term the centrol effect 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), al,, 2003), 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. Hew- identification to species, Most recently male gcnital cver, methyl bromide, used for fumigation of grain traits fbr Cbllosobruchus were compared and sum- legumes (YOneda et al., 1990) has been recognized marized by [fuda 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 anq hence, was called fbr its wcll studied because they are not readily available controlled use by the Montreal Protocol (UNER (but see Pfaffenberger and Johnson, 1976; Arora, 2000), 1978; Delobcl ct al., 1995b), which makes identifi- Alternatively, parasitoids (e.g., a trichogram- cation bascd on pre-adult stages diMcult. In con- matid egg parasitoid UScana, larval-pupal para- trast to the differential accessibility among devel- sitoids, pteromalid Dinarmus, Anisopteromalus, an opmental stages of morphological characters, mo- eupelmid Eupelmus, and a braconid Heterospilus; lccular characters can be mere stable and usefu1 as

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keys fbr identification. Using the PCR-RFLP tech- 441 63-74, Anton, K.-W. {1999) Revision of the Sttlcohntehus nique, which is relatively easy and inexpensive, genus Chojo 1937 and descriptien of llarasttteobruchtts nov. species of all developmental stages of important gen. (Co[eoptera, Bruchidae, Bruchinae). Linz BioL stored bean in the Callosobrttchtts have pests genus Beitr. 31: 629-650. succcssfu11y et al., been distinguished(TUda 1995). Anton, K.-W. (2000) Fivc ncw spccies ofthe C2iUosobruchus chinensis group from the Orienta] Region and Australia {Coleoptera: Bruchidae: Bruchinae), Genus 1I/ 13-28. CONCLUSION Anton, K.-W, J. Halperin ancl M. Calderon (1997) An anno- tated list of the Bruchidae (Co]eoptera) of Israel and aclj a- Bean beetles, as other insects, ex- phytophagous centareas. Is'rael J. EntomoJ. 31/ 59-96. hibit significant conservatism in host utilization. ARC-PPRI (2003) Releases qf Biologicat Control Agents Fceding on dried seeds not only enables repcated against Pftteds in South "ica. http:1115S.240,199.39f institutesfppri/mainldivisionslweed$divlreleases.htm. generations but also broadens the diet of some Arora, G. L.(1977) Taxonomy ofBruchidae CColeoptera) of bean beetle pests. Our study based on molecular Northwest India. Part I. Adults. Orient. Inseet SortzpL 7: indicates this to use phylogenetics preadaptation 1-132. dry, hard beans that human store ofbeans precedes Arora. G. L. (1978) Taxonomy ofBruchidae (Coleoptera) of has been selected fbr by arid habitat climate (Tuda Northwest India. PEirt II. Larvae. Orient. Insect Stmpl, et al,, 2006). Recent investigations of Asian bean 8: lg8.

Arora, L. A Biolqgy and 1lrxonomy beetle fauna have revealed diversity ofthe species G. (1980) Stucij,qf'the of the Genus Bruchidius (Coteopte}u.' Bruchidoe) fiom and ecology of the seed predators. This also led to lndia. Flnat feck. Rept. (i974-1979) US, PL-480 Res. the finding of invasions of alien bean beetles (Tuda PJoj A7-EN7ii03. Department ofZoology, Puajab Uni- et al., 2001; TUda et al., unpublished). Invasion yersity, Chandigarh. 96 pp. routes estimated by our molecular approach indi- Balachowsky. A. S. (1962) Entomologie 42plieutie a iL4gri- cated the invasion of the bean beetle was associ- cttltura n)me I, Coltioptaies. Nlo1. 1. Masson et Cie Editeurs, Paris. 564 pp. ated with imported alien host seeds, I propose the Becerra, J. X. {2003) Synchronous coadaptation in an an- recent bean beetle invaders may be used fbr modest cientcase efherbivory, Proc, NatLAead. SZ]i. CLYd 100: control ofthe seed dispersal of beneficial but inva- 12804.12807. sive alien plants (`conflictplants' (Neser, 1994)). Boeke. S. J., A, A. C, Sinzogan, R. R de Almeida, R W M. de Bocr, G. Jeong, D. K. Kossou and J. J. A. van Loon ACKNOWLEDGEMENTS (2003) Side-effects ef cewpea treatment with botanical ins¢ cticides on two parasitoids of Callosobruchtts macu- I am gratefu1 to the Japanese Society ef Applied Entomol- tattts, EntomoL Eu]. A?7F)l. I08/ 43-51. ogy and Zoology for the honorable award and the opportunity Borowiec, L. The genera ofseed-beetles (Coleoptera, to write this review. Thanks are also due to A]ex Delobel and (l987) Bruchidae). R)isk. Pisino Entosnol. 57: 3--207. Steven Vamosi for their valuable coinments on the manuscript. Caswell, G. H. (1960) Observations on an abnormal forrn of This study was supported partly by the Filj'iwara Natural Callosobruchus maculatus (E). Butl. Entomoi, Res, 50: History Foundation, the Sumitomo Foundation, and Grant-in- 671-680. Aids lbr International Scientific Research (Ficld Research Center, T, D. and C. D. Johnson (1972) Comparative life his- 09041145) and for Scientific Research (A) (08304049, tories ofSennius. Environ. EntontoL 2: 669-672. 15208007), (B) (1440Se03, 17405005) and for lybung Scien- Center, T. D, and C. D, Johnson (1974) Coevolution ofsome tists (B) (15770011) from MEXT. This paper is dedicated to seed beetles (Coleoptera: Bruchidac) and their hosts. my collaborators, the late Prot Shwu-Bin IIorng for his work Ecotegy5S:1096-1103. on behaviera] eeology of Ckeilosohruchus and the late Dr. Chagas, J. M. (l981) Leuc,aena leucocephala as a green ma- Liang-Yih Chou for his contribution to applied entomology. nure fbr bean growing in cerrado soil. Rysq. Agn Btas. REFERENCES 16i 809-814. Delgado, C., G. Couturier and A. Delobel (1997) Oviposi- Abrahamson, IM G., C. R BIair, M. D, Eubanks and S, A, tion of sced-bcctlc CZityoborus serripes (Sturm) (Co- Morehead (2003) Sequential radiation of unrelated or- teoptera/ Bruchidae) on palm (AstrocaflJum cham- ganisms/ the gall fiy Eurosta solidaginis and the tumbting biia) fruits under natural conditions in Peru. Ann, Soc. flower beetle Moidellistena convicta. .1 EvoL BioL l6: EntomoL F',un. 33: 405-409. 781-789. Delobel, A. and B, Delobel (2003) Les plantes hotes des Alvarez, N., J. R. Napoles, K,-W Anton, B. Benrey and M. bruchcs (Coleoptera/ Bruchidae) de la faune dc France, Hossaert-McKey (2006) Phylogenetic relationships in une analyse critique. Bull. it4ens. Soc. Linn. L.von 72: the Neotropical bruchid genus Acanthosceiides (Bruchi- 199-221. nae, Bruchidae, Coleoptera). J Zool. S.vst, EvoL Res, Delobei, A. and C. D. Johnson (1998) First reeord ofaseed-

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