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Coleoptera: Chrysomelidae

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Coleoptera: Chrysomelidae JapaneseJapaneseSociety Society of AppliedApplied Entomology and Zoology AppL Entomol. ZooL 42 (3): 337-346 (2007) http/]todokon.orgl Review Applied evolutionary ecology of insects 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 beetles 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 seed 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 Bruchus 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 seeds (Fabaceae). They are fi:equently damage in storage. `weevils' 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' `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 Acanthoscelides and Bruchidius 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 NII-Electronic Library Service JapaneseJapaneseSociety Society of AppliedApplied Entomology and Zoology 338 M.Tum 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 <Eubaptus) Kytorhinini-1 genus (19,torhintts) spinipes (Tbran, 1962), Caryedon fasciatus (Pre- - Pachymerini 12 genera (Cat:]Jedon) vett, 1966), Cai:yoborus serripes (Delobel et al., Rhaebini-1 genus(Rhaebus) 1995a) and Stator beali (Nilsson and Johnson, 1993; Fox and Mousseau, 1995), The host pods The genera in bold inc]ude pest species. and seeds of these beetles are hard and relatively large (Janzen, 1971; Moreno-Casasola et al., 1994; ranapanichpan, Szentesi and Jermy, unpublished). Delobel et al., 1995a; Delgado et al., 1997), In Ckel- These molecular studies also indicate potential losohruchus, such egg clustering had previously problems about relative phylogenetic positions not been observed but it was recently fbund in a among other recently dcscribcd genera. For exam- non-pest species, which develops in a large, hard ple, a recent molecular study found 7Liberculo- seeded legume (TUda and Buranapanichpan, un- bruchus to be polyphyletic (Kergoat, 2005b), indi- published). These examples imply that egg laying cating a plesiomorphic trait is used for the generic pattern is not phylogenetically constrained but may key and revision of the genus using genital mor- respond to selection impesed by host plant charac- phology may be necessary. Data on recently de- teristics (the size andfor hardness of pods and scribed species, synonymies and host plants have seeds), parasitoid pressure and abiotic factors been actively accumulated from different biogeo- (Janzen, 1971; Mitchell, 1977; Delobel et al., graphic regions of Asia and Africa that would con- 1995a). tribute to the understanding of evolutionary diver- [[Ypical hosts of bean beetles are legumes sity of the Old Wbrld (e,g., Arora, 1977, 1980; (Fabaceae) (84% of the known hosts; Johnson, Egorov and Tbr-Minassian, 1983; Morimoto, 1990; 1970) but the palm (Arecaceae), morning glory Anton et al,, 1997; Anton, 1999, 2000; Kingsolver, (Convolvulaceae), mallow (Malvaceae) and about 1999; Tuda, 20e3; Johnson et al., 2004; Tuda and 30 other families are also used as hosts (Southgate, rlUda Morimoto, 2004; et al., 2005) and the New 1979; Jehnson, 1981). Most bean beetles are either Wbrld (e.g., Johnson, 1970; Romero and Johnson, monephagous or oligophagous; their host range is 2000) bean beetles (see also Zacher, 1952; Udaya- limited to restricted plant taxa, usually subtribes giri and Wadhi, 1989 fbr overviews). and tribes at most (Jermy and Szentesi, 2003; De- lobel and Delobel, 2003; [IUda et al., 2005; Kergoat 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-insect 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 NII-Electronic Library Service JapaneseJapaneseSociety Society of AppliedApplied Entomology and Zoology 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- Bruchus pisorum, the pests of broad beans and draogo and Huignard 1981; Messina, 1984; Toque- peas, respectively
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