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Network Scan Data Selbyana 20(1): 40-48. 1999. BROMELIAD-EATING WEEVILS 1.R. FRANK Entomology and Nematology Department, University of Florida, Gainesville, FL 32611-0630 USA. E-mail: [email protected] ABSTRACT. All known bromeliad-eating weevils-24 species in four genera (Cactophagus, Cholus, Dias­ tethus, and Metamasius)-are native to the tropics and subtropics of the Americas. Several attack more than one bromeliad genus. Six are known to attack Ananas comosus. Four (Metamasius dimidiatipennis, M. callizona, M. quadrilineatus, and M. sellatus) have traveled far from their native ranges by hitchhiking in exported/imported bromeliads. Metamasius mosieri (a small weevil) attacks bromeliads of small stature, whereas M. callizona (a larger weevil) attacks bromeliads of larger stature. Tillandsia utriculata popUlations in Florida (USA) have suffered devastating attack by the immigrant weevil M. callizona (from Mexico), perhaps because these plants have evolved in isolation and have no method of resistance. A biological control campaign against M. callizona is in progress. RESUMEN. Todos los picudos comedores de bromelias conocidas, los cuales pertenecen a 24 especies en cuatro generos (Metamasius, Cactophagus, Diastethus y Cholus), son nativos de los tr6picos y subtr6picos de las Americas. Varios atacan mas de un genero de bromelias. Seis atacan Ananas comosus. Cuatro (Metamasius dimidiatipennis, M. callizona, M. quadrilineatus y M. sellatus) han viajado lejos de sus am­ bientes nativos como "polizones" en bromelias exportadas e importadas. Metamasius mosieri (un picudo pequeno) ataca bromelias de tamano pequeno, mientras M. callizona (un picudo mas grande) ataca bro­ melias de mayor tamano. Poblaciones de Tillandsia utriculata en la Florida (Estados Unidos) han sufrido ataques devastadores por el picudo inmigrante de Mexico M. callizona quizas porque dichas poblaciones han evolucionado aisladamente y no tienen ninglin mecanismo de resistencia. Una campana de control bio16gico en contra de M. callizona esta en progreso. Key words: Bromeliaceae, Curculionidae, Metamasius, Cholus, Cactophagus, bromeliaphagy INTRODUCTION The purposes of this paper-to review knowl­ edge of the hostplant relationships of weevils In a natural population of bromeliads, a large that feed on bromeliads, to consider the ecology proportion of dead and dying plants is an un­ of these weevils under natural and altered con­ common sight, at least when insects are causing ditions, and to document how some of them the damage. Although collectors of bromeliads have become pests-may stimulate additional may encounter numerous kinds of invertebrates observations and research by canopy biologists. living in leafaxils, few if any of these animals are detrimental to the plants. Even when lubber WEEVILS THAT EAT BROMELIADS grasshoppers, Romalea guttata (Houttuyn), swarm in the Florida Everglades, the damage Weevils (Curculionidae) are almost exclusive­ they do to the larger bromeliads seems to be ly phytophagous. The more than 40,000 de­ largely cosmetic; leaves are severely chewed, scribed species make them one of the largest but the plants are not killed. families of beetles. The diversity of weevil spe­ Such observations, coupled thus far with the cies often is attributed to their special adapta­ study of only a few insect species and those not tions to diverse hostplants. known to damage bromeliads grown as orna­ Adults and larvae of 24 species of weevils mental plants, may have given rise to the pop­ belonging to four genera (Cactophagus, Cholus, ular misconception among growers that brome­ Diastethus, and Metamasius) currently are liads lack consequential insect pests. Evidence known or suspected to be specialized feeders on in this paper suggests, to the contrary, that bro­ bromeliads (TABLE 1, with distribution records meliads in natural populations in the neotropics compiled from O'Brien & Wibmer 1982 and are attacked by phytophagous insects potentially Wibmer & O'Brien 1986). Each of these 24 spe­ lethal to the plants but that these insects gener­ cies is native to some restricted area of the trop­ ally have sparse populations. Among these in­ ics and subtropics of the Americas. Although sects are weevils (Curculionidae), which upon two of the genera (Cactophagus and Metama­ arrival in a previously unoccupied locality may sius) are closely related, the relationship of the cause severe losses to bromeliad populations, other two genera is remote, indicating that "bro­ whether the plants are growing under natural or meliaphagy" evolved independently in several cultivated conditions. ancestral lineages of weevils. 40 FRANK: BROMELIAD-EATING WEEVILS 41 TABLE 1. Weevils known to eat bromeliads, or that are suspected of doing so, with their distribution and hostplants in their native areas. Weevil Distribution Hostplants Cactophagus lojanus (Heller) Bolivia, Brazil, Ecuador, Peru ? Cactophagus miniatopunctatus Mexico, Belize, Guatemala, Hon­ Ananas comosus Chevrolat duras, Nicaragua, Costa Rica Cactophagus sanguinolentus Mexico, Belize, Guatemala, Hon­ Aechmea bracteata (Olivier) duras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador Cactophagus transatlanticus Costa Rica, Colombia, Ecuador, ? (Kirsch) Venezuela Cactophagus validirostris Mexico, Guatemala, Honduras, Aechmea bracteata, Vriesea sp. (Gyllenhal) Panama, Colombia, Ecuador Cholus spinipes (Fabricius) Grenada Ananas comosus Cholus vaurieae O'Brien Venezuela Ananas comosus Diastethus bromeliarum Costa Rica ? Champion Metamasius alveolus Vaurie Costa Rica ? Metamasius bromeliadicola Costa Rica ? Champion Metamasius callizona (Chevrolat) Mexico, Guatemala, probably not Tillandsia deppeana, T. ehlersi­ Panama despite old record ana, T. heterophylla, T. limba­ ta, T. roland-gosselinii, T. streptophylla, T. utriculata, Aechmea mexicana, Catopsis sp. (see also text) Metamasius ciliatus (Champion) Mexico Aechmea bracteata Metamasius cincinnatus Nicaragua, Costa Rica, Panama, Werauhia werckleana, Guzmania Champion Ecuador circinnata Metamasius dimidiatipennis Mexico, Guatemala, Nicaragua, Ananas comosus (Jekel) Costa Rica, Panama, Colombia, Venezuela, French Guiana, Ecuador, Peru, Brazil Metamasius fasciatus (Olivier) Costa Rica, Panama, Venezuela, ? Ecuador Metamasius jlavopictus Mexico, Guatemala Tillandsia velickiana (Champion) Metamasius hebetatus (Gyllenhal) Nicaragua, Costa Rica, Panama, ? Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia Metamasius mosieri Barber Cuba, Dominican Republic, USA Tillandsia balbisiana (see also (Florida) text) Metamasius nudiventris Champion Mexico, Nicaragua, Costa Rica, ? Panama Metamasius quadrilineatus Mexico, Guatemala, EI Salvador, Tillandsia guatemalensis, T. oro­ Champion Honduras genes, T. ponderosa, T. stand­ leyi, T. yunckeri, Racinaea spi­ culosa, Werauhia nephrolepis, W. pectinata, Catopsis hahnii, C. morreniana, Billbergia sp. Metamasius quadrisignatus Montserrat, Guadeloupe, Domini­ ? (Gyllenhal) ca, Martinique, Panama Metamasius ritchiei Marshall Cuba, Jamaica Ananas comosus Metamasius rugipectus Mexico, Costa Rica, Panama ? (Champion) Metamasius sellatus Champion Mexico, Belize, Guatemala, Nica­ Tillandsia guatemalensis ragua, Panama 42 SELBYANA Volume 20(1) 1999 TABLE 2. Weevils found attacking bromeliads in other-than-natural environmental conditions. Weevil Country Plants attacked Reference Cactophagus miniatopunctatus Mexico' Ananas comosus Vaurie 1967 Cholus spinipes Grenada' Ananas comosus Marshall 1922 Cholus vauriae Venezuela' Ananas comosus Salas & O'Brien 1997 Metamasius callizona Mexico2•3 Tillandsia spp. Frank & Thomas 1994b Mexico' Ananas comosus This paper USA (FL),,4 13 genera Frank & Thomas 1994b USA (FL)5 Native Tillandsia Frank & Thomas 1994b USA (FL)' Ananas comosus Frank & Thomas 1994b Metamasius cincinnatus Panama2 Werauhia werckleana This paper Metamasius dimidiatipennis Venezuela1.6 Ananas comosus Salas et al. 1996 Indonesia (Java)4 Imported bromeliads Vaurie 1966 Metamasius mosieri USA (FL)3 Imported Tillandsia This paper Metamasius ritchiei Jamaica' Ananas comosus Marshall 1916 , Cultivated pineapples in beds or fields. 2 Native bromeliads in habitats disturbed by humans (e.g., coffee plantations). 3 Cultivated bromeliads in shade-houses or greenhouses or gardens. 4 Imported bromeliads attacked by an adventive weevil species. 5 Native bromeliads in native habitats (attacked by an adventive weevil species). 6 Also in Costa Rica, Panama, and Ecuador (Vaurie 1966, Frank & Thomas 1994b. Weevil larvae mine inside the stem, flowers­ here, organized under the names of the weevils. talk, or (in plants such as Ananas) the fruit. Al­ Information on natural habitats and hosts pre­ though the presence of these larvae demon­ cedes information on weevils in altered habitats, strates bromeliaphagy by a weevil species, iden­ and the latter is summarized in TABLE 2. tification keys for weevils depend on characters provided by adults. Entomologists surveying the Metamasius callizona insect fauna of a region seldom bother to collect larvae, much less take time to rear them to the Vaurie (1966) examined one museum speci­ adult stage for identification. Adults feed exter­ men of M. callizona labeled "Panama, Potreri­ nally on leaves, flowerstalks, and flowers. After llos, Dec., 1935." For the greater part of three developing through immature stages inside a weeks in June 1994, part of the time accompa­ plant, resultant adults disperse to detect other nied by A. Muzzell, the author searched unsuc­ plants suitable for attack. During their dispersal, cessfully for
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