POPULATION ECOLOGY Diaprepes abbreviatus (Coleoptera: Curculionidae): Host Plant Associations
S. E. SIMPSON,l H. N. NIGG, N. C. COILE,2 AND R. A. ADAIR:!
University of Florida, Institute of Food and Agricultural Sciences, Citrus Research and Education Center, 700 Experiment Station Road, Lake Alfred, FL .'3.'3850
Environ. Entollloi. 25(2): 333-349 (1996) ABSTRACT Diaprepes ahhreviatus (L.) is an insidious pest of citrus, sugarcane, and other economic crops of subtropical and tropical areas of the United States and several Caribbean island nations. Host plants associated with this pest, 157 genera, =270 species in 59 plant families, are listed. Plants that support egg deposition to adult include Citrus spp.; Arachis hypogal'a 1.., peanut; Sorghum hicolor L. (synonym: S. vulgare) sorghum; Sorghum hic%r L. ssp. him/or, guinea com; 7£a mays L., com; Eugenia uniflora L., Surinam-cherry; Dracaena draco (L.) L. dragon tree; Ipomoea hatata (L.) Lam., sweet potato; and Saccharum officinarl1lll L., sugarcane. More than 40 plant species, in 20 families, are associated with larval feeding. As a result of inadequate management strategies and a wide range of adult and larval food plants, D. abbreviatus can be considered a major long-term threat to the survival of several agronomic crops.
KEY WORDS Diaprepes ahhreviatlls, host plant, citrus, sugarcane, corn, potato, peanut
Diaprepes abbreviatlls (L.) (=the West Indian sug- ulatory and control measures to be effective, the arcane rootstalk borer weevil) has become an im- plant hosts of D. abbreviatlls must be identified. portant long-term pest of citrus and ornamental Otherwise, an overlooked hosl could allow endem- crops in Florida (Hall 1995).Since its introduction ic infestation. Consequently, we have compiled in 1964, D. abbreviatlls has spread from an Orange here the following 5 D. abbreviatus plant associa- County citrus nursery to =22,727 ha of commer- tions: (1) plants from which adults have been col- cial citms in 17 counties tmd 94 commercial citms lected; (2) plants upon which adults feed; (3) and ornamental plant nurseries (Woodruff 1968, plants upon which eggs have heen laid; (4) plants Beavers et a!. 1979, Hall 1995). To limit distribu- from which larvae have been collected; and, (5) tion of the weevil, strict quarantines are currently plants that support the entire life cycle. The ob- enforced in weevil-infested citrus and ornamental jectives of this study were to document the plants nurseries. No movement of contaminated host associated with D. abbreviatus in Florida and var- plant material is allowed. ious island nations of the Caribbean and to deter- S(weral strategies have been llsed for the control mine additional plants of the family Rutaceae lIpon of adult and immature D. abbreviatlls over the pasl which D. abbreviatus adults will feed. centnry. For example, weevil control was reported in Pm'lto Rico by handpicking adults from infested sugarcane, Saccharum officinarum L. (Wolcott Materials and Methods 1933a). Early eradication attempts in Florida Forced, no-choice feeding experiments lIsed 5- proved unsuccessful (Griffith 1975). In the past 60 10 adult, field-collected weevils caged (30.5 by yr, a few persistent and nonpersistent pesticides 30.5 em, aluminum, stocking front cages, BioQuip, have been used for adult and larval control; several Petaluma, CAl for 24 h with bouquets of the fU- persistent pesticides have provided some measure taceous plants listed in Table 1. Deionized, glass- of control (Wolcott 1954), but with environmental distilled water was provided in 25-ml Erlenmeyer hazards. Biological control methods have had lim- flasks with a cotton dental wick. Leaf bouquets in ited success (McCoy and Simpson 1994). For reg- deionized glass-distilled water in 2.5-mlErlenmey- er flasks were presented for 24 h. If after 24 h no IFlorida Departlllent of Agriculture and Consumer Services, Division of Plant Industry, 3027 Lake Alfred Road, Winter Haven, feeding had occurred, the bouquet was left for an VI. 331;1;1. additional 24 h. After 48 h of testing, weevils were 2Florida D•.partment of Agricnlture and Consumer Selvice.s, provided \vith immature, fully expanded red grape- Division of Plant Industry, P.O. Box 14700, Gainesville, 1'1, 32614- fruit leaves for 24 h to ensure adequate nutrition, 7l00. "TIl(>Kt'rr Center for Snstainable Agriculture, Vero Beach Re- and then testing was resumed with that cage. search Station, 7055 Cherry Lane, V•.ro Beach, FL 32966. Leaves were obtained from the Division of Plant
0046-225X/96/0333-34\l$02.00/0 © 1996 Entomological Sudety of America 334 ENVIRONMENTAL ENTOMOLOGY Vol. 25, no. 2
Table 1. Aduh D. abbreviatus forced, no-choice feeding on RutaceoU8 plants
Scientific name Common name Aegle mannelos (L.) Corr. Serr. Bael Aeglopsis chevalieri Swingle Chevalier's aeglopsis Afraegle gabonensis (Swingle) Eng!. Gabon powder flask Afraegle pan/G1llala (Shum.) Eng!. Nigerian powder flask Amyris balsamifera L. Sea amyris Atalantia sp. Malantia Gas/m/roa edulis Llave & Lex. White sapote Gas/m/roa tetrameria Millsp. Wooly white sapote Gilrops/s gilletim'" Swingle & M. Kellerman Gillet's cherry orange Gitms amhlycarpa Ochse Nasnaran mandarin Gitms aurantiifolia (Christm.) Swingle 'Oavao' Lime Gitrus aurantiifolia (Christm.) Swingle Persian lime Gitms aurantiifolia (Christm.) Swingle Giant Key lime Gitms auranti/folia (Christm.) Swingle Citrus of Moi lime Gitms aurantium L. 'Bouquet' Sour orange Gitms aurantium L. 'Chinotto' Sour orange GUms bergam/a Risso & Poit. 'Bergamont' Bergamot Gitms depressa Hay Shekwasha mandarin Gitms hystrix DC. Caffre lime Gitms ichangensis (Swing.) Ichang papedaa Gltms limettioides Tan. Sweet lime GUms limettloides Tan. 'Palestine' Sweet lime Gitms limon (L.) Burm. f. 'Baboon' Lemon Gitms limon (L.) Burm. f. 'Bearss' Lemon Gitms limon (L.) Burm. f. 'Cameron Highlands' Lemon Gitms limon (L.) Bum1. f. 'Ponderosa' Lemon Citms X limonia Osbeck 'Volkamer' Lemon Gitms 11Ulxillla(Burm.) Merr. Pummelo Citrus nUicroptera Montr. Melanesian papeda Gitms medica L. var. Sarcodactylis (Noot.) Swingle Buddha's hand Gitn,s medica L. 'Etrog' Etrog citron Gitms m€yeri Tan. Meyer lemon Gitn,s obovoidea Hart. ex Takahashi Kinkoji Gitms reticulata Blanco 'Silver Hill' Satsuma Gitn,s tachibana (Mak.) Tan. Taehibana mandarin Gitn,s tachibana (Mak.) Tan. Kinokuni mandarin Gitms X limonia Osbeck Rangpur lime Gitms X paradisi Madfad. 'McCarty' McCarty grapefruit Glausena lansium (Lour.) Skeels Wampee Eremocitms glauca (Lind!.) Swing. Lime, Australian desert" Eremocitms hybrid EremocitrIls X Skekwasha mandarin Fortunel/a crassifolia Swingle Meiwa kumquat Fortunel/a japonica (Thunh.) Swingle 'Marumi' Round kumquat Fortlmella margarita (Lour.) Swingle 'Nagami' Oval kumquat Fortunella /Illlrgarita (Lour.) Swingle 'Centennial' Oval kumquat Fortrmella polyandra (Rid!.) Tan. Tropical kumquat Glycosmis pentaphylla (Retz.) Correa Toothpick glycosmis Linwnia acidissima L. Wood apple Microcitms australasica (F.J. Muell.) Swingle Australian finger lime Microcitms australis (Planch.) Swingle Australian round lime Microcitms papuana H.F. Winters New Cuinea lime Murraya koenigii (L.) Sprengel Curry leaf Murraya paniG1llata (L.) Jack Orange jasmine Naring; crent/lata (Roxb.) Nicholson Hesperethusa Pambums missionis (Wight) Swingle Pamburas Severinia buxifo/ia (Poi ret) Ten. Chinese box orange Severinia disticha (Blanco) Swingle Philippine box orange Swillg/ea g/rltinosa (Blanco) Merr. Tabog Triphasia trifo/ia (Bunn. f.) P. Wilson Myrtle lim!" X Citroncin.lS Citrangequat hybrid X Citrofortlmella microcarpa (Bunge) Wijnands Calamondin Zanthoxylum fagara (L.) Sarg Lime, prickly ash or wild lime
This was a forced, no choice feeding experiment. a No feeding.
Industry (DPI) Citrus Arboretum, Florida Depart- ceous species. Weevil feeding was recorded as ei- ment of Agriculture and Consumer Services ther feeding or no feeding. (FDACS), Winter Haven, FL. This arboretum sup- Florida Department of Agriculhlre and Con- plies validated budwood and seed to the Florida sumer Services/Division of Plant Industry main- citrus industry and breeding stock for other ruta- tains the Florida State Collection of Arthropods. April 1996 SIMPSON ET AL.: D. abbreviatus HOST PLANTS 335
Insect specimens are collected from various loca- plant. Sugarcane, citrus, dragon tree, and sweet oems, submitted to the department for identifica- potato are hosts known to support the entire life tion and a detailed specimen identification report cycle. Corn, peanut, and sorghum have not been issued for each sample submitted (FDACS 1964- reported to support a generation of this weevil in 1994). The records for D. abbreviatus from 1964 the field. There were 59 plant families with which through 1995 were obtained and compiled by ac- D. abbreviatus has been associated by observation tivity-adult collection only, adult feeding, egg de- of adults, adult feeding, egg masses, or larvae on position, larval feeding, and all life stages present. the plant (Table 2). We also reviewed the scientific literature from Observations of host plants upon which adult 1898 to 1995 for reports of the same activities. weevils feed and lay eggs were more varied than plants which support larva] development (Tab]es 2 and 3) (Myers 1931), but this is perhaps only be- Results and Discussion cause of the ease of above ground observation as Sixty-two plants in the family Rutaceae were compared with the examination of roots. D. ab- tested for adult feeding (Tab]e 1). Of these 62, D. breviatus females deposit eggs between leaf sur- ahhreviatlls would not feed on 2 plants-Citrus faces and glue the leaf surfaces together with a ichangensis (Swing.) and Eremocitrus glauca colorless, adhesive substance. Egg masses have (Lind!.) Swing. E. glauca is a desert plant and has been observed at the base ofleaves and at different small, narrow, leathery leaves. C. ichangensis has locations on the leaf tips, which makes locating leaves similar in size to commercial citrus varieties. them difficult (Jones 1915). Adult feeding is usu- Of 430 FDACS/DPI specimen identification re- ally limited to the leaves of host plants, but adult cords of D. abbreviatus, 378 were observations of feeding on icaco fruit, Chrysobalanus icaco 1", im- adults collected by hand or dislodged from foliage. mature citrus fruit (unpublished data), date palm There were 22 reports of adult feeding, 20 of ]ar- fruit, Phoenix dactylifera L., corn seed, and flowers vae in the soil, 6 of egg masses, and 4 reports of a of certain host plants, Phoenix spp., has been re- host with all life stages of D. abbreviatus. Plants ported (Jones 1915, Wolcott 1936, Jackson 1963). with all life stages present at the same time were Watson (1904) reported sugarcane and sweet Citms spp. (3 reports) and 1 report from Surinam- potato [Iporrwea batatas (L.) Lam.] as supporting cherry, Eugenia uniflora L., a common ornamental larva to adult development, while peanut and sor- in Florida and the Caribbean. Egg masses were ghum served as larval hosts with a life cycle from found on Dracaena (3 reports), citrus (2 reports), egg to adult of 357 d. He also reported adult feed- and 1 questionable report of egg masses on Bra- ing on young coffee, Coffea arabica L., and guava, zilian pepper, Schinus terebinthifolius Raddi. Lar- Psidium guajava L., plants. Pierce (1915) reported vae were found on citrus (13 reports), Dracaena (2 rearing larvae to adults on sugarcane, citrus, corn, reports), Schefflera (2 reports), Acacia (1 report), sweet potato, Guinea corn, sorghum, and lime, Veitchia (a palm) (1 report), and sugarcane (1 re- Citrus aurantiifolia Christm. A variety of other port). From the FDACSIDPI specimen reports, it plant species was reported as adult feeding hosts: appt'ars that under Florida conditions Citrus spp., mango, Mangifera indica L., avocado, Persea amer- Dracaena spp., and Surinam-cherry can support icana Mill., coffee, and grasses, family Gramineae. the entire life cycle. Also based on these reports, Jones (1915) reported adults feeding on the fol- Acacia spp., Schefflera spp., and Veitchia spp. may lowing hosts: jobo tree, Spondias rrwmbin L., icaco, support the development of larva to adult. citrus, mango, sicklepod, Cassia tora [current The association of D. abbreviatus with plants fell name of sicklepod is Senna obtusifolia (L.) Irwin into the following 5 categories: (1) plants from & Barneby], sugarcane, and corn. Jones (1915) re- which adults were collected; (2) plants upon which ported that the adults preferred citrus, aguacate the adults were observed feeding; (3) plants from (avocado), icaco, sicklepod, and particularly jobo. which larvae were collected; (4) plants upon which Wolcott (1933a) reported sword beans, Canavalia D. abbreviatus eggs were laid; and, (5) plants that gladiata (Jacq.) DC, and seed corn as supporting support all stages of the life cycle. larva to adult development in the laboratory. Wol- Using all of Table 2 data (FDACS/DPI reports, cott (1933b) also reported adult feeding prefer- scientific literature, personal observations), peanut, ence for bucare, Enjthrina berteroana Urban, gal- Arachis hypogaea L., sorghum, Sorghum bicolor L. lito, several species of Erythrina, including E. ssp. bicolor, citrus, corn, Zea maize L., sugarcane folkersii Krukoff & Moldenke, jobo, moca, Andira and sweet potato, Iporrwea batata (L.) Lam., are inermis (W Wright) Kunth ex DC, icaco, wild fig, crops which support all stages of the life cycle of Ficus sp., guava, gmicima, Guazuma 1.llmifolia D. abbreviatus and may serve as population res- Lam., tamarind, Tamarindus indica L., quenepa, ervoirs (Tables 2 and 3). Surinam-cherry and drag- Meliococcus bijugatus Jacq., willow,Salix humbold- on tree, Dracaena draco (L.) L. are common trop- tiana L., castor bean, Ricinus communis L., and ical and subtropical ornamentals which support the gandu], pigeon pea, Cajan1.lscajan (L.) Ruth. Wol- entire life cycle (Tables 2 and 3). These plants cott (1936) reported adult defoliation of moca and were classified as life-cycle hosts based on collec- adult feeding on young leaves of ceiba, Ceiba pen- tion of all life stages of D. abbreviatus from that tandra (L.) Gaertn., and oviposition on older 336 ENVIRONMENTAL ENTOMOLOGY Vol. 25, no. 2
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&:I ~< U z< E: ....l < S 8-. ~~_ ::.u.....< o~@'r::- 2 _<'l " u("":)~..s::. E§ :;..~.::..~ 8- <:J:J~ ~~ ~~~~ ~ E'" '" o" E" E o U 00 C) C) 'C'" 'C'" :;;::;::'" '" u C1 340 ENVIRONMENTAL ENTOMOLOGY Vol. 25, no. 2 E'" '" o" E" E o U .8 'I:'" <0.. o ..c: 2 o U April 1996 SIMPSON ET AL.: D. abbreviatus HOST PLANTS 341 ;;;- ...... l cl ~ ~ u r.5 ~ <00 ~<~ ;~::: ::§ ] ...,'" S'" ZZ ~'" c ~ bO " S ~~ C C 0 bOc'" c'"bO ~ E rl rl ::;" E ~ 0 0 0 ~ U E bO 1! G c .;S'" .~'" 2 0 i: - - ] 0 E" ..§ -s e OJ >-c ;:§;:§ ~ "C S ~ "C" "C-g --- '" ;;< 0 t e o c e :S:S ~ § § l ~ " :to ..c'" E .b" ~oo xx e" -s.. ~. f ..~ - 8(;; ~8 UU~o;:s!;:l (;i....• -€: € 8 ""~u ~~cor:- ~ ~~3~ §:<~< ~ur..U"'---- r.. Ii..t.. u r.. r..0u~ ...:1«< < «« «< "E leaves of mango. Total eggs laid by confincd fe- males ranged from 1,293 to 7,046. Many females laid =5,000 eggs over their lifetime (Wolcott 1936). According to Wolcott (1936), larvae feed on sug- arcane, grapefruit, C. X paradisi Macf., pepper, Capsicum annuum L., lima bean, Phaseolus luna- tus L., and yuca, Manihot esculenta Cranz. Al- though unsprouted kernels of corn comprised the normal larval laboratory diet, lima beans can also serve as a larval diet. Larvae were able to withstand long periods of fasting (Wolcott 1936). If larvae can survive long periods without food, this pest may be more difficult to manage, particularly in annual crops. Fennah (1942) listed citrus, sugarcane, string beans, Phaseolus vulgaris, velvet bean, Mucuna deeringiana (Bort.) Merr., lima bean, bonavist, La- blab purpureus L., pigeon pea, cassava, Manihot esculenta Crantz, mustard, Brassica spp., celery, Apium graveolens L., yam, Dioscorea batatas Dec- ne., although sometimes Ipomoea batatas is in er- ror called yam, banana, Musa sp., egg plant, So- "E '"c: lanum melongena L., Guinea corn, sorghum, sweet c: o potato, guava, cacao, Theobroma cacao L., cotton, E avocado, pear, Pyrus communis L., coffee, castor E o U bean, hog plum, Spondias purpurea L. or Ximenia americana L., icaco, physic nut, Jatropha curcas L., yellow calthrope, caltrop? Kallstroemia 1fwxima (L.) Hook. & Am., hibiscus, Hibiscus spp., and so forth, as adult feeding hosts. Larval hosts were Crotalaria spp., pigeon pea, sword bean, pepper, Gliricida spp., bread and cheese, also called cat- claw, Pithecellobium unguis-cali (L.) Benth., groundnut (peanut), sorghum, sugarcane, nut grass, Cyperus spp., cotton, cacao, sweet potato, citrus, guava, castor bean, cassava, rose, Rosa spp., ornamental palms, and Urena lobata L., caesar weed. A disturbing aspect of Fennah's (1942) re- port is cotton as a larval and adult host. We found no reports of egg deposition on cotton, although this is implied by the presence of larvae. There were no reports of life-cycle completion on cotton. Fennah (1942) also reported banana as an adult and larval feeding host. Banana is an especially im- portant tropical and subtropical crop. Taylor (1898) first described the economic dam- age to citrus, larval ravaging of orange tree roots growing in Jamaica. Watson (1904) reported eco- nomic damage to sweet potato and sugarcane. Skeete (1928) reported D. abbreviatus as a major pest of sugarcane. Table 2 lists other major and minor economic host crops of D. abbreviatus cul- tivated in the United States. These are tobacco, Nicotinia tabacum L., celery, potato, millet, beans, lima and string, Phaseolus vulgaris L., pecan, Car- ya illinoinensis (Wagenh.) K.. Koch, peach, Prunus persica (L.) Batsch, pear, blackberry, Rubus argu- tus Link (synonym: Rubus betulifolius), and avo- cado. Diaprepes abbreviatus should be considered a major long-term pest of many tropical and sub- 344 ENVIRONMENTAL ENTOMOLOGY Vol. 25, no. 2 Tahle 3. Life-stage ohservations for D. abbreviatus from the scientific literature and FDACS/DPI insect specimen reports (sec Tahle 2) FAMILY/Botanical name Common name Adult feeding, egg deposition, and larva CONVOLVULACEAE: [pomoea batatas (L.) Lam. Sweet potato GRAMINEAEIPOACEAE: Sacchanlnl officiuanlnt L. Sugarcane Sorghum bicolor (L.) Moench ssp. cultivars Guinea com Sorghum bicolor (L.) (synonym: S. vulgare) Sorghum Zea mays L. Com LEGUMINOSAE: Arachis hypogaea L. Peanut LILlACEAE: 1Jracaena draco (L.) L. Dragon tree MYRTACEAE: Eugenia unif/ora L. Surinam-cherry HUTACEACE: Citms (l/tr(mtiifolia (Christm.) Swingle Lime Citms X paradisi Macfad. cultivar Hed grapefruit Citms X paradisi Macfad. Grapefruit Adult feeding and larvae ANACAHDIACEAE: Spoudias mombin L. (synonym: Spondias luteal Yellow momhin, jobo ARALIACEAE: Scheff/era actinophylla (End].) Harms Scheffiera LEGUMINOSAE: Plwseolus vulgaris L. String bean, etc. MELIACEAE: Swietenia mahagoni Jacg. Mahogany SOLANACEAE: SOlllUlllU melougeull L. Eg!(plant Adult feeding, egg deposition ANACAHDIACEAE: M(lIlgifera indica L. Mallgo SpOlulias mombin L. Jobo, yellow mombin CONVOLVULACEAE: Argyreia neroosa (Burm. f.) Cephalic vine Egg deposition and larvae LILIACEAE: Dracaena compacta Schott, Nyman & Kotschy Dracaena Dracaetla cincta Bak. Dracaena rainbow Egg deposition only GRAMINEAEfPOACEAE: Panicum maximum Jacg. Guineagrass LILIACEAE: Dracaena fragrans (L.) Ker-Gaw!. Com plant MORACEAE: Ficus citrifolia P. Miller Short-leaved fig FiclIs sp. Wild fig RUTACEAE: Citn.ls sp. Citrus Larvae only AHALIACEAE: ScheJJlera elegantissima (Veitch) Lowry & Frodin False aralia ASCLEPIADACEAE: Hoya camosa (L. f.) R.Br. Wax plant CUPRESACEAE: jllllipems conferta L. Shore juniper jllnipems virginiana L. Red-cedar CYPERACEAE: Cypems sp. Nut grass April 1996 SIMPSON ET AL.: D. abbreviatl.ls HOST PLANTS 345 Toble 3. Continued FAMILY/Botanical name Common name DIOSCOREACEAE: Dioseorea X cayene.lSis Lam. Yellow yam EBENACEAE: Diospyros virginiana L. Wild or common persimmon E UPHORBIACEAE: Manihot eseulenta Cranz Cassava, yucca GRAMINEAEIPOACEAE: Bothrioehloa penusa (L.) Pitted bluestem LEGUMINOSAE: Grotalaria sp. Hattlebox Phaseolus IWJatlls L. Lima bean Pitheel/obi ••m unguis-cali (L.) Benth. Bread and cheese, catclaw Pitheel/obium sp. LILIACEAE: Aloe vera (L.) Bunn. f. Aloe Dracaena sanderianfl Hart. Ribbon plant Uriope sp. Lilyturf MALVACEAE: Urena lobata L. Caesar weed MARANTACEAE: Maranta leneonenm E. Morr. Prayer plant MYRSINACEAE: Ardisia crenata Sims Coralbeny PALMAE/ARECACEAE: Veitchia merril/ii (Becc.) H.E. Moore Christmas palm PIPEHACEAE: Piper sp. Black pepper RUTACEACE: EI"ven g"nus & species. Citrus SOLANACEAE: Capsictlln allnum L. Pepper STERCULIACEAE: Theobroma cacao L. Cocoa Adult feeding only 89 species tropical crops. Worldwide, only 12 plant species (1992) defined a host for fruit fly quarantine pur- furnish 90% of the world's food (Thurston 1969). poses as a plant supporting larval development into These 12 are potato, sweet potato, cassavas (man- viable adults. Schroeder et al. (1979) took the po- iocs), banana, coconut, and grains: rice, wheat, sition that plants not supporting D. abbreviatus lar- maize (corn), sorghum, millet, rye, and barley. Of val development would not contribute to the these 12 plants, 7 are adult and larval D. abbrev- spread of this weevil. The synopsis in Table 3 of iatus hosts. More than half of the world's diet of the observations in Table 2 illustrate the problem fats and oils comes from oilseed plants: cotton, co- with D. abbreviatus. Eleven species were reported conut, oil palms, olives, peanut, rape, soybean, and with all life stages present though not necessarily sunflower (Guidry 1964). Cotton and palm(s) sup- at the same time (Table 3). Thus, sweet potato, port the larva and the adult. Peanut appears to be sugarcane, guinea corn, corn, dragon tree, (Dra- a life-cycle host. caena sp.), sorghum, peanut, Surinam-cherry, and From a regulatory standpoint, D. abbreviatus most (if not all) commercial citrus varieties can be presents exceptional problems. Forty-eight of the considered hosts as defined by Saxena and Pathak ""220 hosts in Table 2 were associated with the egg (1977), Schroeder et al. (1979), and Cowley et al. or larval stages, or a combination with the adult (1992). There were 40 additional species that had stage. Eighty-nine species were associated with larvae present, either alone or with feeding adults adult feeding (Table 2) plus 60 additional ruta- or with egg masses also present (Tables 2 and 3). ceous plants in Table 1. Saxena and Pathak (1977) Schroeder et al. (1979) presented data on larval defined a host as a plant supporting generation to survival for a few of these species; for the others, generation survival. Liquido et al. (1990) regarded we have no information. Adults fed on an addi- a nonhost for regulatory purposes as a plant un- tional 89 species (Tables 2 and 3) plus the 60 ru- suitable for support of the life cycle. Cowley et al. taceous species/cultivars in Table 1. If we confound 346 ENVIRONMENTAL ENTOMOLOGY Vol. 25, no. 2 Table 4. Host plant genera with reference to family from Tables I and 2 Abel1lwschus MALVACEAE Delonix LEGUMINOSAE Nicotianll SOLANACEAE Acacia LEGUMINOSAE Desmodium LEGUMINOSAE Pambunts RUTACEAE Acer ACERACEAE DiT/wcarpus SAPINDACEAE Panicum GRAMINAEAE Acnistus SOLANACEAE Dioscorea DIOSCOREACEAE Passiflora PASSIFLORACEAE Aegle RUTACEAE Diospyros EBENACAE Pe~ea LAURACEAE Aeglopsis RUTACEAE Dizygotheca (see SchejJlera) Phaseolus LEGUMINOSAE Afraegle RUTACEAE Dolichos (see l.ahlab) Phaseo/,1S (see Vigna) Albizia LEGUMINOSAE Dracaena LILIACEAE Phoenix PALMAE Aloe LILIACEAE Eriobotrya ROSACEAE PiT/lenta MYRTACEAE Ambrosia COMPOSITAE EremocitrusRUTACEAE Piper PIPERACEAE Amomis (see Pimental Erythrina LEGUMINOSAE Pisonia (see Gual'ira) Amyris RUTACEAE Eugenia MYRTACEAE Pitheel/obium LEGUMINOSAE Andira LEGUMINOSAE Euphoria (see DinwcaryJUs) Pittosporum PITTOSPORACEAE Andml'0gon (see Bothriochloa) Feijoa MYRTACEAE Platycl{l(hts CUPRESSACEAE Apium UMBELLIFERAE Ficus MORACEAE Poncints RUTACEAE Arachis LEGUMINOSAE Fortunella RUTACEAE Pongamia LEGUMINOSAE Ardisia MYRSINACEAE Gliricidia LEGUMINOSAE Prunus ROSACEAE Argyreia CONVOLVULACEAE Glycosmis RUTACEAE Psidi"m MYRTACEAE Atalantia RUTACEAE Gossypium MALVACEAE Pueraria LEGUMINOSAE Aulllirachta MELIACEAE Guaiacum ZYGOPHYLLACEAE Pynts ROSACEAE Baccharis COMPOSITAE Guapira NYCTAGINACEAE QuemlS FAGACEAE Bauhinia LEGUMINOSAE Guarea MELIACEAE Rhizophora RHIZOPHORACEAE Bischojia EUPHORBIACEAE GuazurruJ STERCUIACEAE Rhus ANACARDIACEAE Bothriochloa GRAMINEAE Haematoxylum LEGUMINOSAE Ricinus EUPHORBIACEAE Brassien CRUCIFERAE Hemerocallis LlLIACEAE Rosa ROSACEAE Brysonima MALPIGHIACEAE Hibiscus (see Abe/moschus) Roystonea PALMAE Btlcida COMBRETACEAE Hoya ASCLEPIADACEAE RubllS ROSACEAE Bursera BURSERACEAE llex AQUIFOLIACEAE Saccharum GRAMINEAE Caesalpinia LEGUMINOSAE Illicium ILLICIACEAE Salix SALICACEAE CajllllllS LEGUMINOSAE lndigofera LEGUMINOSAE SchejJlera ARALIACEAE Callistemon MYRTACEAE Jnga LEGUMINOSAE Schinus ANACARDIACEAE Calophyllum GUTTIERAE Jl'01lwea CONVOLVULACEAE Schrankia LEGUMINOSAE Canavalia LEGUMINOSAE Jatropha EUPHORBIACEAE Simna LEGUMINOSAE Capsicum SOLANACEAE JunipenlS CUPRESSACEAE S''lJerenia RUTACEAE Can}a JUGLANDACEAE Ka/Istroetnia ZYGOPHYLLACEAE Solanum SOLANACEAE Casimiroa RUTACEAE Kaelreuteria SAPINDACEAE Sorghum GRAMINEAE Cassia (see Senna) Lab/ab LEGUMINOSAE Spondias ANACARDIACEAE Cassia LEGUMINOSAE Lage~rro~rnaLYTHRACEAE Swietenia MELIACEAE Cedrela MELIACEAE Leptoglottis (see Schrankia) Swinglea RUTACEAE Ceiba BOMBACACEAE Linwnia RUTACEAE Syzygium MYRTACEAE Celtis ULMACEAE Liriope LILIACEAE Tabebuia BIGNONIACEAE Centrosemll LEGUMINOSAE Lanchocarpus LEGUMINOSAE Tamarindus LEGUMINOSAE Cestnm. SOLANACEAE Magnolia MAGNOLIACEAE TI1Jhrosia LEGUMINOSAE ChnJSalidocaryJlIS PALMAE Mangifera ANACARDIACEAE Tenninalia COMBRETACEAE ChrysobalaenrlS CHRYSOBALANACEAE Manihot EUPHORBIACEAE Theobroma STERCULIACEAE ChnJSophylium SAPOTACEAE Manilkara SAPOTACEAE Thuja (see Platycladus) Citropsis RUTACEAE Maranta MARA:'IlTACEAE Tormbia (see Guapira) GitnlS RUTACEAE Melia MELIACEAE Triphasia RUTACEAE Clausena RUTACEAE Melicoccus SAPINDACEAE Triplaris POLYGONACEAE C/erodendrum VERBENACEAE Microcitrus RUTACEAE UlmllS ULMACEAE Cocc%ba POLYGONACEAE Minwsa LEGUMINOSAE Umna MALVACEAE Goffea RUBIACEAE Montezuma MALVACEAE Veitchia PALMAE Conocarptts COMBRETACEAE Mucuna LEGUI\IINOSAE Vigna LEGUMINOSAE Cordia BORAGINACEAE Murraya RUTACEAE Vigna LEGUMINOSAE Cmta/aria LEGUMINOSAE Musa MUSACEAE Ximenia OLACACEAE Cupaniopsis SAPINDACEAE Myrcianthes MYRTACEAE Zanthoxylum RUTACEAE CupresSlIS CUPRESSACEAE Myrica MYRICACEAE Zea GRAMINEAE CY)'ems CYPERACEAE Naringi RUTACEAE the observations here with host factors such as ovi- feeder and will oviposit on anything that has two position site selection (Boetel et al. 1992), water surfaces close together, including the calendar in stress (Hoffman and Hogg 1992), and hosts and the office" (W. Schroeder, USDA, Orlando, per- seasonality (Jones et al. 1992, Doederlein and Sites sonal communication). Yet in a choice experiment 1993, Diawara et al. 1994, Zalom et al. 1995), the with ornamental plants, oviposition site prefer- number and expense of host experiments for D. ences were shown between plant species; waxed abbreviatus is daunting. paper was preferred over all plant species tested No regulation of this weevil would allow adult (Abreu-Rodriguez and Escobar 1983). The detec- movement, therefore all plants associated with the tion of larvae is not convenient. The survival po- adult are suspect. The egg stage is difficult to de- tential of D. abbreviatus for most larval hosts is tect and D. abbreviatus appears to show little pref- not known. The larval development period has erence for oviposition sites. "The insect is a general been reported: 180-240 d, laboratory (w. Schroe- April 1996 SIMPSON ET AL.: D. abbreviatus HOST PLANTS 347 del', USDA, Orlando, FL, personal communica- Based on the information available on D. ab- tion), 48-443 d (Wolcott 1934, 1936), 300-312 d breviatus, we define a regulatory host for this in- (Watson 1904), 240-300 d (Wolcott 1933c), and sect as any plant associated with any life stage. It =1 yr in the field (Jones 1915; W. Schroeder, is unfortunate that D. abbreviatus appears to be USDA, Orlando, personal communication). The nonselective in oviposition sites because this means larval period may include up to 16 instal's and a a host could be any plant in an infested location. diapause period of up to 388 d (Wolcott 1933c, Within the United States, there are in excess of 1934, 1936). This diapause period may not be true 350 million cultivated acres of land (USDA 1994). diapause. It is marked by no feeding and quies- Approximately 70% of this acreage, with yields val- cence in a soil chamber (Jones 1915; Wolcott ued in excess of 36 billion dollars, contains D. ab- 1933c, 1934). The larval development period may breviatus 1000wn host plants. United States direct vary within these ranges for the larvae from a sin- and indirect economic losses to Diaprepes root gle egg mass (Wolcott 1934, 1936). Adult D. ab- weevil could be substantial. Additional studies are breviatlls were obtained from 132 to 297 dafter needed to determine the impact this pest may have infesting potted citrus seedlings with neonate lar- on agronomic crops growing in temperate regions vae obtained from field-collected females (Beavers of the United States. and Sdheime 1975). From the data in Tables 2 and 3, eggs may be Acknowledgments present without feeding adults and larvae may be present without evidence of oviposition. The nat- This study was supported by a Florida citms grower ural spread and dispersion of D. abbreviatus are box tax for research. Funds for this project were made not known. We suggest that D. abbreviatlls in one available from the Citrus Production Research Marketing Order by the Division of Marketing and Development, stage or another could be spread by movement of Florida Department of Agriculture and Consumer Ser- any of the plant species listed in Tables 1-3. Once vices. Florida Agricultural Experiment Station Joumal moved, however, the potential for survival and de- Series No. R-04636. velopment in temperate climates and annual crops is in question. Wolcott (1934) reported a larval pe- riod of 48 d for 1 grub fed only corn; 2 others References Cited developed in 53 and 58 d. There is a variable qui- Abreu-Rodriguez, E., and M. P. Escobar. 1983. Ovi- esct'nt or diapause period; the pupal period is = 15 position preference of Diaprepes abbreviatus (Cole- d; egg hatch is =7 d. The record for development optera: Curculionidae) on various omamental foliage from egg to adult in the tropics is =6 mo (Wolcott plants. J. Agric. Univ. PH. 67: 117-120. 1934), a record broken by =50 d (Beavers and Sel- Anonymous. 1908. Cacao and gmbs. J. Jamaica Agric. heime 1975). One year is usual for the general life Soc. 12: 367-368. 1909. Cocoa trees and grubs. J. Jamaica Agric. Soc. 13: cycle in the Caribbean (Watson 1904, Jones 1915). 125-127. However, a tropical and subtropical origin does not 1933. Service and regulatory announcements, July- preclude development in a temperate climate. For Sept. 1933. USDA Bureau of Plant Quarantine, Ser- example, the tropical Mediterranean fruit fly now vice and Regulatory Announcements 116: 197-243. infests many crops in several temperate zones Washin!,rton,DC. (Metcalf 1995). Overall development time and the Ballou, H. A. 1912. Insect pests of the Lesser Antilles. occurrence and the length of the larval diapause Imperial Department of Agriculture Barbados, West Indies Pamphlet Series 71: 66--69. period may be influenced by the female (Mous- Beavers, J. B., and A. C. SeUleime. 1975. Develop- seau and Dingle 1991). Diapause, which occurs in ment of Diaprepes abbreviatus on potted citrus seed- the same percentage in larvae in the tropics (32%) lings. Fla. Entomol. 58; 271-273. as in temperate zones (36%) and is equal in time, Beavers, J. B., R. E. Woodruff, S. A. Loveslrand, and may be used to pass the winter (Denlinger 1986). W. J. Schroeder. 1979. Bibliography of the sugar- Watson (1904) noted that D. abbreviatlls was not cane rootstalk borer weevil, Diaprepes abbreviatus. noticed in sugarcane and sweet potato before Entomol. Soc. Am. Bull. 25: 25-29. 1901, probably because of small numbers. This in- Boelel, M. A., D. D. Walgenbach, G. L. Hem, B. W. Fuller, and M. E. Gray. 1992. Oviposition site se- festation pattern was repeated in citrus (Woodruff lection of the northern com rootworm (Coleoptera: 1968). This pest is liable to be unobserved when Chrysomelidae). J. Econ. Entomol. 85: 246-249. it first attacks a crop. The first symptom of larval Cassani, J. R. 1986. Arthropods on Brazilian pepper- feeding, for instance, is wilting of the plant (Wat- tree, Schinus terebinthifolius (Anacardiaceae), in son 1904). With a female reproductive potential of south Florida. Fla. Entomol. 69: 184-196. 5,000 eggs, a fairly long life cycle, and hidden eggs Cowley, J. M., R. T. Baker, and D. S. Harte. 1992. and larvae, D. abbreviatus will probably be well Definition and determination of host status for mul- tivoltine fruit fly (Diptera: Tephritidae) species. J. established before it is detected. Perennial crops Econ. Entomol. 85: 312-317. are most at risk. For most annual plants associated DenIinger, D. L. 1986. Dormancy in tropical insects. with D. abbreviatus, the life-cycle time may pre- Annu. Rev. Entomol. 31: 239-264. clude a completed generation, but this is not Diawara, M. M., J. T. Trumble, C. F. Quiros, K. K. known and has not been systematically studied. White, and C. Adams. 1994. Plant age and scason- 348 ENVlHONMENTAL ENTOMOLOGY Vol. 25, no. 2 aI variations in genotypic resistance of celery to beet prepes abbreviatus (L.) in Florida citrus, pp. 247-256. armyworm (Lepidoptera: Noctuidae). J. Econ. Ento- Proceedings, Caribbean Food Crops Society 30th An- mol. 87: 514-522. nual Meeting, St. Thomas, U.S. Virgin Islands, 31 Doederlein, T. A., and R. W. Sites. 1993. Host plant July-5 August 1994. preferences of Frankliniella occidentalis and Thrips Metcalf, R. L. 1995. A biography of the medfly. Citro- tabaci (Thysanoptera: Thripidae) for onions and as- graph July: 10-12. sociated weeds on the southem high plains. J. Econ. Mousseau, K. T., and H. Dingle. 1991. Matemal ef- Entomol. 86: 1706-1713. fects in insect life histories. Annu. Rev. Entomol. 36: Earle, F. S. 1928. Insect and other pests of sugar cane, 511-534. pp. 162-188. In Sugarcane and its culture. Wiley, New Myers, J. G. 1931. A preliminary report on an inves- York. tigation into the biological control of West Indian in- FClmah, R. G. 1938. Citrus pests investigations. Rep. sect pests. Empire Marketing Board 42: 173. Agric. Dep. Dominica 1937: 27-28. Nowell, W. 1912. Plant pests and diseases. Rep. Local 194.2. The citrus pests investigation in the Windward Dep. Agric. Barbados 1911-1912: 50-51. and Leeward Islands. British West Indies. Agricultural Oramas, D., J. Rodriguez, and A. L. Gonzalez. 1990. Advisory Department, Imperial College of Tropical Effect on yam (Dioscorea rotundata Poir) of soil spray Agriculture. Trinidad, British West Indies. and seed treatment with the nematicide-insecticide 1947. 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Graphic Summary of World Ag- rieties to the brown planthopper, pp. 303-317. III riculture. U.S. Dep. Agric. Misc. Publ. 705. Brown planthopper: threat to rice production in Asia. Hall, D. 1995. A revision of the bibliography of the Intemational Rice Research Instihlte, Los Baros, Phil- sugarcane rootstalk borer weevil, Diaprepes abbrev- ippines. iatus. Fla. Entomol. 78: 364-377. Schroeder, W. J., R. A. Hamlen, and J. B. Beavers. Herbaugh, L. L., and E. C. Mosteller. 1980. Ovicidal 1979. Survival of Diaprepes abbreviatus larvae on capacity of various chemicals for control of '"VestIn- selected native and omamental Florida plants. Fla. dian sugarcane root borer on Dracaena marginata, Entomol. 62: 309-312. Plymouth, FL, 1979 Diaprepes abbreviatus. Insectic. Shapiro, J. P., and T. R. Gottwald. 1995. Resistance Acaricide Tests 5: 170-172. of eight cultivars of citrus rootstock to a larval root Hoffman, G. D., nnd D. B. Hogg. 1992. Effect of weevil, Diaprepes abbreviatus L. 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Cunningham, and S. Nakagawa. 1933c. The larval period of Diaprepes abbreviatus L. 1990. Host plants of Mediterranean fruit fly (Dip- J. Agric. Univ. P.R. 17: 257-264. tera: Tephritidae) on the Island of Hawaii (1949-1985 1934. The diapause portion of the larval period of Dia- survey). J. Econ. Entomol. 83: 1863-1878. 7Jrepesabbreviatus L. J. Agric. Univ. P.H. 18: 417-428. Martorell, L. F. 1945. A survey of the forest insects of 1936. The life history of Diaprepes abbreviatus at Hio Puerto Rico. Part n. J. Agric. Univ. P.R. 29: 457-481. Piedras, P.R. J. Agric. Univ. P.H. 20: 883-914. McCoy, C. W., and S. E. Simpson. 1994. Past and 1952. Control of the soil inhabiting grubs of Puerto current IPM strategies to combat the spread of Dia- Rico. J. Agric. Univ. P.R. 34: 333-337. April 1996 SIMPSON ET AL.: D. abbreviatus HOST PLANTS 349 1954. Hesidual effectiveness of insecticides against soil Zalom, F. G., C. Castane, ami R. Gabarra. 1995. Se- inhabiting insects. J. Agric. Univ. PH. 38: 108-114. lection of some winter-spring vegetable crop hosts by Woodruff, R. E. 1968. The present status of a West Bemisia argeTitifolii (Homoptera: Aleyrodidae). J. Indian weevil (Diaprepes abbreviatus (L.)) in Florida Econ. Entomol. 88: 70-76. (Coleoptera: Curculionidae). Florida Department of Agriculture Division of Plant Industry Entomology 77. Received for publication 1 August 1995; accepted 8 No- Gainesville, FL. vemher 1995.