First North American Records of the East Palearctic Seed Terrenus (Coleoptera: Chrysomelidae: Bruchinae), a Specialist on Mimosa (Albizia Julibrissin, ) Author(s): E. Richard Hoebeke, A. G. Wheeler, Jr., John M. Kingsolver and David L. Stephan Source: The Florida Entomologist, Vol. 92, No. 3 (Sep., 2009), pp. 434-440 Published by: Florida Entomological Society Stable URL: http://www.jstor.org/stable/40343681 . Accessed: 29/09/2014 17:35

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FIRST NORTH AMERICAN RECORDS OF THE EAST PALEARCTIC SEED BEETLE BRUCHIDIUS TERRENUS (COLEOPTERA: CHRYSOMELIDAE: BRUCHINAE), A SPECIALIST ON MIMOSA (ALBIZIA JULIBRISSIN, FABACEAE)

E. Richard Hoebeke1,A. G. Wheeler, Jr.2,John M. Kingsolver3andDavid L. Stephan4 departmentof Entomology, Cornell University, Ithaca, NY 14853

departmentof Entomology, Soils, and Plant Sciences,Clemson University, Clemson, SC 29634

3FloridaState Collectionof , c/o P.O. Box 147100,Gainesville, FL 32614

departmentof Entomology, North Carolina State University,Raleigh, NC 27695

Abstract

The easternPalearctic bruchine seed beetleBruchidius terrenus (Sharp) (Coleoptera: Chry- somelidae:Bruchinae), a specialistseed predatorof mimosa or silk tree (Albizia julibrissin), is reportedfor the first time in NorthAmerica based on collectionsfrom seven southeastern states.This is the thirdAsian insectspecies recently reported from A. julibrissinin the UnitedStates. A diagnosis,description, photographs of the adult,a summaryof the known U.S. distribution,notes on seasonalhistory, and a revisionto an existingkey to North Amer- icanBruchidius are presented.

Key Words:Coleoptera, Chrysomelidae, Bruchinae, Bruchidius terrenus, adventive, inva- sivespecies, new records

RESUMEN

Se reportala presenciadel escarabajobruquido Palearctico oriental de la semilla,Bruchi- dius terrenus(Sharp) (Coleoptera:Chrysomelidae: Bruchinae), un depredadorespecialista en semillade mimosao el arbolde seda (Albiziajulibrissin), por primera vez en Norteame- ricabasado en coleccioneshechas en sieteestados de surestede los EstadosUnidos. Este es la terceraespecie de insectoasiatico reportado de A. julibrissinen los Estados Unidos.Se presentauna diagnosis,description, fotos del adulto,un resumende la distributionconocida en los EE.UU, notassobre su cicloestacional y una revisionde una clave existentede los Bruchidiusde Norteamerica.

The Old Worldgenus Bruchidius, with about America,came to ourattention in Aug 2004.A ho- 300 describedspecies (Kingsolver 2004), is repre- meownerin ElizabethCity, North Carolina (Tony sentedin NorthAmerica by 2 adventivespecies, Barefoot),by happenstance,examined developing B. villosus(R) and B. cisti (R). Both species ap- podsof mimosa and discoverednearly all werein- parentlywere accidentally introduced with seeds festedwith small yellowish larvae. Wanting to know oftheir leguminous host plants (Bottimer 1968). moreabout the , he contactedan extensionen- TheEuropean B. villosus,first recorded in North tomologistat North Carolina State University Americafrom Massachusetts (Olsen 1918), has (StephenBambara). It was firstthought that the been reportedfrom Ontario and Quebec southto larvaemight represent a seed chalcid (Eurytomidae NorthCarolina (Chantal 1972; Redmon et al. 2000; or Torymidae).Additional infested pods ofA juli- Kingsolver2004). This specialistof genistoidle- brissin,collected on Sep 1,2004 (byD.L.S.), about 5 gumeshas beenreleased for the biological control of miles NNW of Raleigh (Wake Co., NC), were Scotchbroom, (L.) Link,in the Pa- broughtindoors for rearing. Adults of an unfamiliar cificNorthwest (Coombs et al. 2004;Hulting et al. bruchineseed beetleemerged in mid-to late Sep. 2008).The Eurasian speciesBruchidius cisti was Specimenseventually sent to E.R.H. in May 2007 firstfound in NorthAmerica in BritishColumbia in wereidentified as theeastern Palearctic Bruchidius 1922 at Nicolaand again in 1965 at Lillooet(Bot- terrenus(Sharp). The identificationwas confirmed timer1968) and has beenreported as a pestof the afterspecimens were compared with identified ma- foragelegume sanfoin, Onobrychis viciifolia Scopoli, terialhoused in the collectionof the National Mu- in Montana(Hewitt & Burleson1976). seumof Natural History (Washington, DC). An unidentifiedbeetle, which would become the In thispaper, we givethe first North American thirdadventive species of Bruchidiusin North recordsof B. terrenus,an Asian seed specialistof

This content downloaded from 158.135.136.72 on Mon, 29 Sep 2014 17:35:33 PM All use subject to JSTOR Terms and Conditions Hoebekeet al.: FirstNorth American Records of Bruchidius terrenus 435 mimosaor silk tree (Albiziajulibrissin) that oc- testaceous,but somespecimens with apical seg- curs widely in the eastern Palearctic Region: mentsblack; foreand mid legs reddishorange; China,Taiwan, and Japan (Udayagiri& Wadhi metafemurusually withbasal 1/2black, apical 1989;Morimoto 1990; Hua 2002).We list and map 1/2testaceous. localityrecords for 7 southeasternstates (Ala- Vestiture.- Composed of fine silvery gray setae bama,Florida, Georgia, Mississippi, North Caro- evenlydistributed over body but withdark brown lina,South Carolina, and Tennessee);give a diag- circularpatches on either side of midline of pronotal nosis,description, and photographsof the adult to disk, each with conspicuouscentral gray spot facilitateits recognitionamong the U.S. bruchine (Fig.1); elytralinterstices with dark brown spots fauna; and provideobservations on its seasonal (Fig. 1); apices of elytrasometimes dark brown historyand habitsin the Southeast. (Fig.3); pygidium(Fig. 3) usuallyimmaculately sil- verygray, sometimes with faint darker clouding. Systematics Structure.- Head (Fig. 4) triangular,eyes pro- Bruchidius Terrenus (Sharp) tuberant,ocular sinus 3/5length of eye, postocu- (Figs. 1-6) lar lobe narrow;frons slightly convex, in some withbrief frontal disk Bruchusterrenus 1886:35. specimens carina;pronotal Sharp, semicircularin outline,lateral marginsgently Bruchidiusterrenus: Chujo, 1937a: 194; 1937b:61; Na- curvate;disk evenlyconvex with slightdepres- kane,1963: 319; Tan et al., 1980:38; Morimoto,1984: sions at caudal angles,densely, evenly punctate; 266; 1990:135. scutellumbroad, bilobed; elytral striae shallow butdistinct, finely punctate; 3rd and 4thstriae orig- Bruchidiusnotatus Chujo, 1937a: 196; 1937b: 64; Tan et inatingon marginaltubercle; interstices alternat- al., 1980:39; Morimoto,1990: 136. ing in width,densely setose; pygidium shallowly Acanthoscelidesterrenus: 1952: convex,uniformly, densely punctulate; fore and Zacher, 465; Udayagiri mid not metacoxae and Wadhi,1989:66. legs modified; denselypunctu- late, lateral 1/2of face denselysetose; metatibia Diagnosis.Members of the Old Worldgenus with lateral carina completeand ending in a Bruchidiusare characterizedby the presenceof a short,acute tooth, ventral carina complete ending singleminute acute tooth on theventromesal mar- in shortmucro, mesal carina complete;basal ab- ginof the metafemur at theapical 1/4;the absence dominalsternite with mesal pore;terminal stern- of marginalteeth or denticleson the pronotum; ite emarginatedto fitpygidial apex. pronotumconical or transverse, without lateral car- Male genitalia.- As in Fig. 5 and 6. Median ina; presenceof a well-developedslender mucro at lobe (Fig.5) 4x as longas its apical width;ventral theapex of the tibia ventrally; and themedian lobe valve subtriangularwith apex bluntlyrounded; ofthe male genitalia lacking "hinge sclerites" (Bot- internalsac denselylined with very fine spicules; timer1968; Borowiec 1987; Kingsolver 2004). apical valve circular;lateral lobes (Fig. 6) sepa- Bruchidiusterrenus can be easily separated ratedby deep cleft, apices spatulate and inwardly fromB. villosusand B. cistiby the charactersin curvate.The male genitaliawere illustratedby thefollowing description and revisedkey. Morimoto(1990: p. 135). Description(Figs. 1-4).Male: Color.- Integu- Female.- Similarin all respectsto male, ex- ment of head usually black, sometimestesta- ceptbasal sternitelacking medial pore; terminal ceous; pronotum,elytra, and pygidiumblack, sterniteevenly rounded, not emarginate. sometimestestaceous (among specimensexam- Size.- Variableamong specimens examined in = ined);ventral areas black exceptabdomen some- 31);body length ranging from 2.46 to 4.07mm; ma- times testaceous; antenna generally entirely jorityof specimens measured 3.69 to 3.82mm.

REVISED KEY TO NORTH AMERICANBRUCHIDIUS (ADAPTEDFROM BOTTIMER 1968 ANDKINGSOLVER 2004)

1. Bodyand appendagesentirely black (or onlyantennal segments 1-4 reddish brown); dorsal vestiture uniformly white 2 - . Bodyusually black,but appendagesnot entirelyblack; head, thorax,elytra, and ventralabdominal seg- mentssometimes testaceous; antennae and legs testaceousto reddishorange; dorsal vestiture predom- inantlysilvery gray with brown mottling, especially on elytralsurface (Figs. 1-2);southeastern United States terrenus(Sharp) 2. All antennalsegments black; 4th stria with prominent, subbasal denticle; mucro absent; western United States and westernCanada cisti(F.) - . Basal 4 antennalsegments reddish brown; striae lacking subbasal denticles; mucro one-eighth as longas basi- tarsus;eastern United States and easternCanada villosus(F.)

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Figs. 1-4.Bruchidius terrenus. 1, Adult,dorsal aspect. 2, Adult,lateral aspect.3, Pygidium,dorsal aspect. 4, Head,frontal aspect. Scale line = 2 mm.

Host Plant to China and Korea (Zhenget al. 2006). It was in- troducedinto the UnitedStates in 1785 (Spong- Mimosaor silk tree(Albizia julibrissin) is na- berg 1990). Withits attractivepink flowers and tiveto southernand easternAsia, occurringeast umbrella-likecanopy, it has been widelyculti-

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only,by A.G.W.)was on 15-IX-2007,in Emanuel Co., GA (Swainsboro).Mimosa trees flowerin Georgia fromMay throughAug and the fruits (pods)mature from Sep toNov (Pardini and Ham- rick2008). In TelfairCo., GA (McRae),on 20-VI- 2007,A.G.W. noted the presenceof ~5 inchpods. Ovipositionbegins when green pods are forming, probablyin earlyJul. Pods ripenfrom late Augto Nov and beginto disintegratesoon afterbut re- ill main on the treesinto winter. Females probably 0 on the ■ lay eggs individually(in clusters) young pods and cementthem in place. Eggs probably M, hatchin 1-2weeks, the larvae emergingfrom the undersideof the egg and tunnelinginto the devel- opingpod. A neonatelarva burrowsthrough the podwall intoa softgreen seed. In NorthCarolina, early-instarsinfested most seeds of pods exam- ined fromearly to mid-Julin ElizabethCity (T. Barefoot,in litt.) and the Raleigh area (D.L.S., personalobservation). Unhatched eggs werestill observed(by D.L.S.) on theoutside of pods in late Jul2004. mid- late instars male Me- By Aug (ll-VIII-2004), Figs.5-6. Bruchidius terrenus, genitalia.5, (-3-4 mm werefound in seeds (T. Barefoot, dianlobe. 6, Laterallobes. long) in litt.).During an examinationof several trees in Raleighin Sep 2004, D.L.S. foundseeds heavily infested(>90% examined),whereas seeds ofthese vated along roadsidesbecause it is fastgrowing same treesin Sep 2007 wereonly slightly infested and drought-tolerant;it is plantedin gardensas a (<5% examined).Although several larvae prob- highlyprized ornamental (Spongberg 1990; Co- ablydevelop in each pod,D.L.S. notedmost seeds thran2004). Albizia julibrissin has becomeinva- contained a single , occasionally2, and sive in the UnitedStates and has spread widely rarely3 larvae;it is unknownif more than a sin- fromsouthern New Englandwest to Missouri and gle larva can successfullydevelop in a seed. Pre- Illinoisand southto Floridaand Texas;it is culti- sumably there are 4 instars. Pupation occurs vatedin Californiaand Oregon,where it has not withina seed insidethe closedpod. The pupation becomeinvasive (Anonymous 2008). periodprobably takes from 10 to 20 d. New-gener- AlthoughB. terrenusappears to be a specialist ation adults chewthrough the seed coat (Fig. 7) on seeds ofA. julibrissin,it has been reported and then chew throughthe pod coat to escape. fromseeds ofblack locust, Robinia pseudoacacia Adultsemerged indoors in mid-to late Sep from (L.), andAcacia confusaMerr. (Morimoto 1990). It infestedpods collected near Raleighon l-IX-2004 is one of 75 species of arthropodsclosely associ- (D.L.S., personalobservation). Adult emergence atedwith Albizia spp. in Asia and is consideredan holes wereobserved (by A.G.W.) on old seed pods importantpest ofA. julibrissinin Taiwan and collectedin early Sep at Clemson,SC (Fig. 8). China(Zheng et al. 2006). New adults, afteremergence, probably feed on pollen in the fall if flowersare still available. Seasonal History and Habits Adults probablyoverwinter near host trees in plant litter.Based on collecting(by A.G.W.)in The seasonalhistory and habitsof B. terrenus 2007 and 2008,adults are foundon thehost from are based mainlyon observationsby the original earlyto late Junto mid-Sep.Bruchidius terrenus collector(T. Barefoot,Elizabeth City, NC), D.L.S. appearsto be univoltine. andA.G.W, supplemented by reference to thebiol- ogyof B. villosus(Redmon et al. 2000). New North AmericanRecords Overwintered,sexually immature adults most likelyemerge in the Southeastin late springand Distributionaldata forB. terrenusin North disperseto mimosawhere they probably attain America(see materialexamined below and Fig.9) sexual maturityby feedingon pollen.In 2008 at are based primarilyon collectionsfrom mimosa Clemson, South Carolina, A.G.W. first found (A.julibrissin) by A.G.W.from Jun to Sep 2007, adults on 10-VI (none were foundduring sam- unless noted otherwise.Parenthetical numbers pling of the same trees on 2-VI) and observed referto adults collected.Voucher specimens are adultsdeep in flowersat Clemsonand elsewhere depositedin theCornell University Insect Collec- in the Southeastwhile surveyingfor the beetle. tion(Ithaca, NY), the NationalMuseum of Natu- The latestobservation of an adult in the field( 1 ral History,Smithsonian Institution (Washing-

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Figs.7-8. Reproductive structures of Albizia julibrissin infested by B. terrenus.7, Seeds. 8, Seeds and podcoat. Arrowsdenote exit holes by newly emerged adults. Scale linein millimeters.

ton,DC), NorthCarolina State Universityinsect AL) and AdrieanJ. Mayor(Great SmokyMoun- collection(Raleigh), the Florida State Collection tains NationalPark, Gatlinburg, TN). ofArthropods (Gainesville, FL), and the personal Material examined:UNITED STATES: ALA- collectionsof Robert H. Turnbow,Jr. (Enterprise, BAMA:Baldwin Co., Loxley,30-VI-2007 (3). Bui-

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VT-2007(8). WilkinsonCo., Mclntyre,28-VI-2007 (17). MISSISSIPPI: KemperCo., De Kalb, 7-VII- 2007 (11). Lee Co.,Tupelo, 7-VII-2007 (1). Noxu- bee Co., E of Shuqualak, 7-VII-2007 (24). Lowndes Co., W of Columbus,7-VII-2007 (19). MonroeCo., W ofAmory, 7-VII-2007 (12). NORTH CAROLINA:Carteret Co., Ft. Macon St. Pk., 10- IV-2006,R. Newman(1, FSCA coll.).Polk Co., Co- lumbus,24-VI-2007 (3). Wake Co., 5 mi. NNW of Raleigh, l-IX-2004,D.L. Stephan (4). SOUTH CAROLINA:Anderson Co., Powdersville,24- VI- 2007 (1). GreenvilleCo., Greenville,24-VI-2007 (1). Oconee Co., SE of Newry,21-VI-2007 (2); Westminster,20-VI-2007 (4); Rt. 123,E ofChauga River,23-VI-2007 (4). PickensCo., Lake Hartwell, Clemson,20 & 21-VI-2007(5) & 10-VI-2008(2); SpartanburgCo., Inman,24-VI-2007 (24). TEN- NESSEE: BradleyCo., Cleveland,8-VII-2007 (1). HamiltonCo., E of Chattanooga,8-VII-2007 (1). Sevier Co., Old GatlinburgLand Fill offGnatty Branch, 5-X-2004,M. Tomkasky(1, Adriean Mayorcoll.).

Discussion Fig.9. Knowndistribution of Bruchidius terrenus in the southeasternUnited States (Alabama, Florida, Two otherAsian mimosa-associatedinsects Georgia,Mississippi, North Carolina, South Carolina, have been detectedrecently in NorthAmerica. and Tennessee). The buprestidbeetle Agrilus subrobustus Saun- ders,whose native range includes China, Japan, Northand SouthKorea, and Taiwan,was discov- lockCo., UnionSprings, l-VII-2007 (2). Calhoun ered in northernGeorgia in 2006. In Japan,the Co., Oxford,6-VII-2007 (3). ConecuhCo., SW of onlyknown host of this little-studiedspecies is Evergreen,30-VI-2007 (1). Conecuh Co., Ever- mimosa(Westcott 2007). Acizzia jamatonica (Ku- green,30-VI-2007 (4). CovingtonCo., W ofRiver wayama),a psyllidnative to east Asia, was re- Falls,30-VI-2007 (3). CrenshawCo., S ofLuverne, portedfrom 2 localitiesin Georgiain 2006 (Hal- l-VII-2007 (4). JeffersonCo., Hueytown,6-VII- bert 2007; Ulyshen& Miller 2007). Surveysof 2007 (6). Lee Co.,Auburn, l-VII-2007 (2). Macon thismimosa specialist in thesoutheastern United Co., Tuskegee,l-VII-2007 (7). Pike Co., Troy,1- States in 2007 and 2008 yieldedrecords from 5 VII-2007 (5). St. Clair Co., Cook Springs,6-VII- new states and additionalcounties in Georgia. 2007 (3). SumterCo., York, 6-VII-2007 (3). Tusca- Acizzia jamatonica was detectedin Europe in loosa Co., Tuscaloosa,6-VII-2007 (3). FLORIDA: 2001,with establishment probably resulting from Alachua Co., W of Hawthorne,29-VI-2007 (2); shipmentsof mimosafrom Asia as prizedorna- Gainesville,24-IV-2007, M. C. Thomas,beating mentals(Wheeler & Hoebeke2009). flowersof Hydrangea quercifolia and Cornusfoe- Bruchidiusterrenus might have beensimilarly mina (1, FSCA coll.). Gadsden Co., S of Quincy, introducedto the southeasternstates with mi- 30-VI-2007(1). JacksonCo., S ofCottondale, 30- mosa nurserystock. This seed predatormight be- VI-2007(8). JeffersonCo., S ofMonticello, 29-VI- comea pestof mimosa in landscapeplantings and 2007 (9). OkaloosaCo., S ofCrestview, 30-VI-2007 couldeven be considereda beneficialaddition to (4). Putnam Co., E of Palatka, 29-VI-2007(8). our faunaby thosewho regard mimosa as an in- Santa Rosa Co., N ofAvalon Beach, 30-VI-2007 vasive species and, therefore,an undesirable (3). WaltonCo., De Funiak Springs,30-VI-2007 plant. (4). GEORGIA: CoffeeCo., Douglas, 28-VT-2007 (1); Pridgen, 28-VI-2007 (4). Emanuel Co., Acknowledgments Swainsboro,15-IX-2007 (1). Lanier Co., Stockton, We thank Barefoot for 28-VI-2007(62). Madison Co., 72 & Tony (ElizabethCity, NC) jet. Hwys. bringingthe discoveryof an Asian seed beetlein mi- 172, l-VIII-2004,R. Turnbow(1, Turnbowcoll.). mosa seeds to ourattention Extension Mitchell initiallythrough Co.,S ofCamilla, 9-VIII-2008 (5). Oconee EntomologistStephen Bambara (Departmentof Ento- Co.,N ofBishop, 28-VI-2007 (1). PutnamCo., N of mology,North Carolina State University,Raleigh) in Eatonton,28-VI-2007 (5); Stephens Co., Toccoa August2004. We are gratefulto Paul Skelley(Florida Falls College,l-VIII-2004, R. Turnbow,on Sol- State Collectionof Arthropods, Gainesville) for provid- idago (10, Turnbowcoll.). Telfair Co., McRae,28- ingthe photographicimages of B. terrenusin Figs. 1-4,

This content downloaded from 158.135.136.72 on Mon, 29 Sep 2014 17:35:33 PM All use subject to JSTOR Terms and Conditions 440 Florida Entomologist92(3) September2009 to Kent Loeffler(Department of Plant Pathology,Cor- MORIMOTO,K. 1984. Bruchidae, pp. 225-226, pl. 44 In nell University,Ithaca, NY) forphotographing infested M. Hayashi, K. Morimoto,and S. Kimoto [eds.], Co- seeds and pods ofAlbizia julibrissin, and to 2 anony- leoptera of Japan in Color,Vol. 4. Hoikusha, Osaka, mousreviewers for their helpful comments on a draftof Japan (in Japanese). themanuscript. MORIMOTO,K. 1990. A synopsis of the bruchid fauna of Thisresearch was supportedby the Cornell Univer- Japan, pp. 131-140 In K. Fujii, A. M. R. Gatehouse, sityAgricultural Experiment Station federal formula C. D. Johnson, R. Mitchel, and T. Yoshida [eds.], funds,Project No. NYC-139404to ERH, receivedfrom Bruchids and Legumes, Economics,Ecology and Co- CooperativeState Research,Education, and Extension evolution. Proc. 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