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Alate Aphid (Hemiptera: Aphididae) Species Composition and Richness in Northeastern Usa Snap Beans and an Update to Historical Lists

Alate Aphid (Hemiptera: Aphididae) Species Composition and Richness in Northeastern Usa Snap Beans and an Update to Historical Lists

Bachmann et al.: Alate on Snap Beans in NE USA 979

ALATE APHID (: ) SPECIES COMPOSITION AND RICHNESS IN NORTHEASTERN USA SNAP BEANS AND AN UPDATE TO HISTORICAL LISTS

1 2 3, Amanda C. Bachmann , Brian A. Nault and Shelby J. Fleischer * 1South Dakota State University Department of Plant Science, 412 W Missouri Ave, Pierre, SD 57501, USA

2Cornell University Department of Entomology, New York State Agricultural Experiment Station, 630 W. North Street, Geneva, NY 14456, USA

3Penn State Department of Entomology, 501 ASI, University Park, PA 16802, USA

*Corresponding author; E-mail: [email protected]

Abstract

Recent aphid-vectored in the northeastern U.S. led to extensive surveys of aphid (Hemiptera: Aphididae) species composition. We report the species composition and rich- ness of alate associated with processing snap bean (Phaseolus vulgaris L.; Fa- bales: Fabaceae) agroecosystems from field surveys conducted during 5 yr in New York and 3 yr in Pennsylvania. Rates of species accumulation were similar between the 2 states, and asymptotic, suggesting reasonably adequate sampling intensity. Our results suggest that about 95 to 100 aphid species are present as alates within these agroeco- systems, a surprisingly high percentage (~14 to 18%) of the total aphid richness. Host records suggest that 61% of the alate aphid species we collected from pan traps placed within snap bean fields were dispersing through this agroecosystem, originating from woody plants in the surrounding landscape. We compiled this information with a recent study of aphid species composition from orchards and an exhaustive inspection of museum samples, and present an updated list of the aphid species in Pennsylva- nia.

Key Words: Aphis glycines, host, pan traps, peach, Phaseolus vulgaris, -vec- tors

Resumen

Los virus recienes transmitidos por áfidos (pulgones) en el noreste de los EE.UU. resultó en un sondeo amplio de la composición de las especies de los áfidos (Hemiptera: Aphidi- dae). Se reporta la composición y riqueza de especies de áfidos alados asociados con el agroecosistema de la habichuela procesada (Phaseolus vulgaris L.; Fabales : Fabaceae) de estudios de campo realizados durante 5 años en Nueva York y durante 3 años en Pennsylvania. La tasa de acumulación de especies fue similar entre los 2 estados y asin- tótica, lo que sugiere la intensidad del muestreo fue razonablemente adecuada. Nuestros resultados sugieren que aproximadamente 95 a 100 especies de áfidos están presentes como alados dentro de estos agroecosistemas, un porcentaje sorprendentemente alto (~ 14 a 18 %) de la riqueza total de los áfidos. Los registros de los hospederos sugieren que el 61 % de las especies de áfidos alados que recogimos en las trampas de caída colocadas dentro de los campos de habichuela se dispersaba a través de este agroecosistema, pro- cedentes de plantas leñosas en el área alrededor. Hemos recopilado esta información con un reciente estudio sobre la composición de especies de áfidos en los huertos de durazno y una inspección exhaustiva de especimenes depositados en museos. Se incluye una lista actualizada de las especies de áfidos en Pennsylvania.

Palabras Clave: Aphis glycines, hospedero, trampas de caída, durazno, melocotón, Phaseolus vulgaris, vectores de virus

Aphids are a small, but diverse group of in- primarily feeders and when present sects with an origin in the and a total in high densities can damage their host plant. of 4800 species world-wide (Grimaldi & Engel Aphids excrete excess carbohydrates from their 2005; Dixon 1985a; Dixon 1985b). They are diet of phloem sap, providing a nutrient-rich 980 Florida Entomologist 97(3) September 2014 substrate for sooty mold fungi to grow. Sooty vania. The CMV epidemics coincided with the mold can be a major problem in a number of appearance of a newly invasive aphid, Aphis agricultural because the mold can either glycines Matusmura (Nault et al. 2009), and as render produce unmarketable or reduce plant was the case with stone fruit, the threat of viral quality of the commodity. Aphids are also im- epidemics led to extensive surveys of the alate portant vectors of viruses that can kill their aphids species composition in the affected . host plant or substantially reduce crop yield These recent surveys of aphid collected from and quality (Agrios 2005). Some viruses are snap bean fields in Pennsylvania and New York, transmitted by aphids in a non-persistent, sty- and peach orchards in Pennsylvania, were quite let-borne manner. They are obtained quickly by extensive. Also from Pennsylvania, J. O. Pepper their aphid vector during short tasting probes, specialized in aphid identification and actively adhere to the stylet lining by binding to helper collected them for most of the 20th century. component proteins or directly to the stylet, and His collections centered at his home in cen- remain there until they are flushed out during tral Pennsylvania (State College) and included another tasting probe (Ng & Falk 2006). Non- much of the surrounding forest and farmland. persistently transmitted viruses can be vectored The bulk of his collection is housed in the Frost by alates, sometimes by multiple aphid species Entomological Museum (University Park, Penn- (Gildow et al. 2008) regardless of whether or sylvania), and he also contributed slides to the not there is on the plant host, and United States National Collection (Beltsville, the epidemiology of these viruses can be influ- Maryland). Pepper (1965) reported 345 species enced heavily by the alate aphid community. in a published list of the aphids of Pennsylvania Several viruses of this type have been intro- and their host plants. To date, this is the most duced recently, or increased in frequency, in the comprehensive published list of aphids for the northeastern U.S. One, virus (PPV), state. However, since and systemat- threatened the stone fruit industry following ics are in flux, the names that Pepper published its arrival in the U.S. This virus causes sharka are currently out of date and in need of revision. disease in parts of Europe and South America The purpose of this study was to identify the where it is endemic (Roy and Smith 1984; Ro- species composition and estimate aphid spe- sales et al. 1998). Type D isolates were detected cies richness in snap bean agroecosystems in in the U.S. in Pennsylvania in 1999 (Damsteegt the northeastern states from field surveys, and et al. 2001), and surveillance and eradication generate a current list of aphid species in this efforts of this included destruc- region using field survey data, literature, and tion of approximately 23% of the non-cherry an examination of the J. O. Pepper aphid col- stone fruit orchards of Pennsylvania (Wallis et lection. al. 2005). As part of these efforts, studies were conducted to determine the potential aphid Materials and Methods species that might serve as reservoir or route of transmission in the region where this virus Detailed methods for alate aphid collection in was first detected (Wallis et al. 2005). Soon snap bean fields in Pennsylvania and New York thereafter, in the early 2000s, Northeastern were published in Nault et al. (2009). To sum- and Midwestern U.S., snap bean crops (Phaseo- marize, we used water pan traps baited with a lus vulgaris L; Fabales: Fabaceae.) had virus- green ceramic tile (Webb et al. 1994) and filled like symptoms ( mosaic and blistering, de- with a 20% propylene glycol solution in snap formed pods) and experienced dramatic yield bean fields in both states from 2002 – 2006 in loss (Larsen et al. 2002). Among the viruses NY and 2004 – 2006 in PA. Traps were installed detected were mosaic virus, bean com- in a total of 56 fields in western NY (12 each mon mosaic virus, bean pod mottle virus, bean yr, except for 2004 which had 8 fields) and 18 yellow mosaic virus, clover yellow mosaic virus, fields in Centre county PA (6 each yr). The traps clover yellow vein virus (ClYVV), cucumber mo- in Centre County formed an approximately 40 saic virus (CMV), tobacco streak virus and white mile transect in the southern portion of the clover mosaic virus (Grau et al. 2002; Larsen et county roughly following state routes 45 and al. 2002; Shah et al. 2006). CMV was the most 192. The traps were checked weekly for aphids prevalent virus detected in these snap bean from the early trifoliate stage (early to mid Jul) fields (Larsen et al. 2002; Shah et al. 2006). As until field harvest. Collection methods in the is the case with PPV, CMV is transmitted by peach ( persica (L.) Stokes; Rosales: Ro- aphids in a non-persistent, stylet-borne manner saceae) orchard are documented in Wallis et al (Nault 1997). CMV-infected plants were often (2005), and also used the water pan traps bait- found in clumps in snap bean fields, which were ed with a green tile. Trapping occurred during 2 consistent with aphid-initiated virus epidemics yr in 2 orchards in central Pennsylvania. (Shah et al. 2005). CMV epidemics also occurred For both studies, aphids were removed more frequently in New York than in Pennsyl- from pan traps and then stored in 70% etha- Bachmann et al.: Alate Aphid Species on Snap Beans in NE USA 981

Table 3. New aphid records from Pennsylvania re- Consulting, Frenchtown, New Jersey), whereas ported in Nault et al. (2009) and/or Wal- those from Pennsylvania were identified by W. lis et al. (2005), but not found in pepper Sackett and A. Bachmann using keys by Smith (1965). et al. (1992) and Blackman & Eastop (2000). Voucher specimens are located at the New York Species Nault et al. Wallis et al. State Agricultural Experiment Station in Gene- kondoi • va, New York, and the Department of Entomol- Aphis glycines • ogy, Pennsylvania State University, University Park, Pennsylvania. Aphis lugentis • Species rarefaction curves were calculated Aphis (Protaphis) middletonii • • for the Pennsylvania and New York collections Aphis pulchella • individually and for both states combined using Nearctaphis clydesmithi • EstimateS (Colwell 2005). Tetraneura nigriabdominalis • A complete list of aphids from Pennsylvania was compiled using the J. O. Pepper Aphid Slide Collection, which is housed at the Frost Ento- mological Museum (University Park, Pennsyl- nol (EtOH), then transferred to potassium vania), as well as species recorded in Pepper hydroxide and heated for 1 h or until clear. (1965). We searched the slide collection in ad- Cleared aphids were rinsed for 10 min each in dition to using Pepper (1965) because Pepper a sequence of 95% EtOH, absolute EtOH, and continued to collect aphids and make slides into clove oil. Once rinsed, each aphid was placed on the late 1980s, but did not publish any updates a drop of Canada balsam on a glass slide and to his original 1965 paper. Because the collec- positioned to expose diagnostic features before tion and Pepper (1965) contained aphid species a coverslip was placed on top. Aphids collected names from the early 20th century, we consult- in New York were identified by R. Eckel (RVWE ed 2 online aphid databases to ensure that the

Table 4. Species in six subfamilies of the family aphididae occurring in Pennsylvania.

Subfamily Tribe Species Anoeciinae Anoecia corni Anoecia cornicola Anoecia oenotherae Anoecia setariae Gillette & Palmer Cerataphidini Cerataphis lataniae (Boisduval) Hormaphidini Hamamelistes spinosus Shimer Hormaphis hamamelidis Fitch Mindarus abietinus Koch Phyllaphidinae fagi (L.) Stegophylla quercicola (Monell) Stegophylla quercifoliae (Gillette) Stegophylla quercina Quednau Pterocommatinae Fullawaya terricola (Hottes & Frison) Plocamaphis flocculosa Weed Pterocomma bicolor Pterocomma medium Baker Pterocomma populifoliae (Fitch) Pterocomma smithiae Saltusaphidini Iziphya flabella (Sanborn) Iziphya vittata Richards Strenaphis elongate (Baker) Thripsaphidini Allaphis verrucosa (Gillette) Subsaltusaphis virginica (Baker) Thripsaphis ballii (Gillette) 982 Florida Entomologist 97(3) September 2014 adianti (Oestlund) Macrosiphum californicum (Clarke) Macrosiphum (Neocorylobium) carpinicolens Patch Macrosiphum (Neocorylobium) coryli Davis Macrosiphum cystopteris Robinson Macrosiphum gaurae (Williams) Macrosiphum gei (Koch) Macrosiphum geranii (Oestlund) Macrosiphum lilii (Monell) Macrosiphum pallidum (Oestlund) Macrosiphum (Neocorylobium) pseudocoryli Macrosiphum ptericolens Patch Macrosiphum tiliae (Monell) Mastopoda pteridis Oestlund (Walker) Microlophium sibiricum (Mordvilko) Microparsus desmodiorum Smith & Tuatay M icroparsus olivei Smith & Tuatay Microparsus singularis (Hottes & Frison) Muscaphis music Börner Myzaphis rosarum (Kaltenbach) Myzodium modestum (Hottes) cerasi (Fabricius) Myzus formosanus Takahashi Myzus lythri (Schrank) Liang Myzus (Nectarosiphon) persicae Nasonovia (Kakima) aquilegiae (Essig) Nasonovia compositellae (Theobald) Nasonovia (Kakima) cynosbati (Oestlund) Nasonovia (Kakima) heucherae (Thomas) Nasonovia (Ranakimia) purpurascens (Oestlund) (Mosley) Nearctaphis bakeri Nearctaphis clydesmithi Nearctaphis crataegifoliae (Buckton) formosana (Takahashi) Neotoxoptera violae (Pergande) Ovatus crataegarius . a ni enn s ylv P in

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ac ro s iphini Decorosiphon corynothrix Börner Diuraphis (Holcaphis) holci (Hille Ris Lambers) Dysa - phis (Pompahis) plantaginea Dysaphis tulipae Ericaphis scammelli (Mason) Ericaphis wakibae (Hottes) atriplicis foeniculi Hyalomyzus collinsoniae (Pepper) Hyalomyzus eriobotryae (Tissot) Hyalomyzus mitchellensis Smith Hyalomyzus sensoriatus (Mason) Hyalopteroides humilis (Walker) lactucae Hyperomyzus (Neonasonovia) nabali (Oestlund) Hyperomyzus (Neonasonovia) picridis (Börner & Blunck) nephrelepidis Davis Illinoia azalea (Mason) Illinoia borealis (Mason) Illinoia canadensis (MacGillivray) Illinoia goldmaryae (Knowlton) Illinoia liriodendri Illinoia pepperi (MacGillivray) & Pepper) Illinoia (Masonaphis) rhokalaza (Tissot Illinoia richardsi (MacGillivray) Illinoia (Oestlundia) rubicola (Oestlund) Illinoia spiraecola (Patch) Linosiphon sanguinarium (Hottes & Frison) Liosomaphis berberidis (Kaltenbach) pseudobrassicae Longicaudus trirhodus (Walker) abrotani (Walker) Macrosiphoniella frigidicola Gillette & Palmer Macrosiphoniella leucanthemi (Ferrari) Macrosiphoniella ludovicianae Macrosiphoniella millefolii (De Geer) Macrosiphoniella (Phalangomyzus) pennsylvanica (Pepper) Macrosiphoniella sanborni Macrosiphoniella subterranean (Koch) Macrosiphoniella tanacetaria (Kaltenbach) Macrosiphoniella tapuskae (Hottes & Frison) M tri b e , phidin a e A mily s u bfa

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pe c ie s 5. S ab le T Abstrusomyzus phloxae (Sampson) Acuticauda solidaginifoliae (Williams) Acyrthosiphon lactucae (Passerini) Acyrthosiphon malvae (Mosley) Acyrthosiphon pseudodirhodum (Patch) agathonica Hottes Amphorophora ampullata Buckton Amphorophora rossi Hottes & Frison Amphorophora rubi Amphorophora sensoriata Mason (Prunaphis) cardui (L.) (Kaltenbach) Brachycaudus (Scrophulaphis)persicae group Brachycaudus (Brachycaudina) rociadae (Cockerell) Brachycaudus (Thuleaphis) rumexicolens (Patch) Brachycaudus (Appelia) schwartzi (Börner) Brachycorynella asparagi (Mordvilko) brassicae Cachryphora canadensis Hille Ris Lambers Cachryphora serotinae (Oestlund) carduinus (Walker) Capitophorus hippophaes Carolinaia caricis Wilson Carolinaia (Galbromyzus) howardii (Wilson) Carolinaia (Glabromyzus) rhois Catamergus Kickapoo (Hottes & Frison) Cavariella aegopodii (Scopoli) Cavariella cicutae (Koch) Cavariella hendersoni Knowlton & Smith Cavariella pastinacae (L.) Cavariella salicis (Monell) Cavariella theobaldi (Gillette & Bragg) Ceruraphis eriophori (Walker) Ceruraphis viburnicola (Gillette) (Pentatrichopus) fragaefolii (Cockerell) Chaetosiphon (Pentatrichopus) minor (Forbes) Chaetosiphon (Pentatrichopus) tetrarhodum (Walker) rufomaculata (Wilson) Cryptomyzus ribis (L.) Bachmann et al.: Alate Aphid Species on Snap Beans in NE USA 983

final list used the most current nomenclature (Aphid Species File – http://aphidspeciesfile. org, accessed 22-IV-2012 and Aphids on the World’s Plants - http://www.aphidsonworld- splants.info/, accessed 1-XI-2013). We combined our findings from the Pepper collection material with the results of our pan trapping study and Wallis et al. (2005) to create a more current list of the aphids of Pennsylvania (Tables 3–11).

Results

In snap bean fields in New York and Penn- sylvania, a total of 8,821 aphids were identified, with 7,484 from New York and 1,337 from Penn- sylvania. A total of 97 species were caught; 71 from New York and 41 from Pennsylvania. We were unable to identify only 254 (2.8%) of the aphids. Of the aphids captured, those species .

Uroleucon nigrotibium (Olive) nigrotuberculatum (Olive) Uroleucon obscuricaudatum (Olive) Uroleucon paucosensoriatum (Hille Ris Lambers) Uroleucon pepperi (Olive) Uroleucon pieloui (Richards) Uroleucon pseudambrosiae Uroleucon rudbeckiae (Fitch) Uroleucon (Uromelan) rurale (Hottes & Frison) Uroleucon russellae (Hille Ris Lambers) Uroleucon sonchellum (Monell) (L.) Uroleucon (Uromelan) taraxaci (Kaltenbach) Uroleucon (Uromelan) tardae (Hottes & Frison) Uroleucon (Uromelan) tuataiae Olive Utamphorophora crataegi Utamphorophora humboldti (Essig) representing 1% or greater of the total number caught in either state are listed in Table 1 (orig- a ni inally published in Nault et al. 2009) with their abundances. A comprehensive list of all aphid species found in Pennsylvania and New York enn s ylv snap bean fields is shown in Table 2 along with P

in their host associations based on Blackman & Eastop (1994, 2000, and 2006). From this host information we estimated that 61 percent of the species dispersing through snap bean fields in both states were most likely coming in from the o cc urring surrounding forests as their hosts are woody, not herbaceous, species (Fig. 1). Species accumulations followed asymptotic patterns (Fig. 2) suggesting reasonably ade- quate sampling of the aphid species present as ac ro s iphini alates in commercial snap bean fields. Overall, M there were fewer aphids collected in Pennsyl- vania, but based on the rarefaction curve there tri b e

, were a similar number of total species repre- sented in a sample of the same number of in- Uroleucon ambrosiae (Thomas) Uroleucon anomalae Uroleucon (Lambersius) caligatum (Richards) Uroleucon chrysanthemi (Oestlund) Uroleucon chrysopsidicola (Olive) Uroleucon (Lambersius) erigeronense (Thomas) Uroleucon eupatoricolens (Patch) Uroleucon (Uromelan) eupatorifoliae (Tissot) Uroleucon floricola Robinson Uroleucon (Lambersius) gravicorne (Patch) Uroleucon (Uromelan) helianthicola (Olive) Uroleucon (Uromelan) illini (Hottes & Frison) Uroleucon impatiensicolens (Patch) Uroleucon lanceolatum (Patch) Uroleucon leonardi (Olive) Uroleucon (Lambersius) luteolum (Williams) phidin a e A mily s u bfa

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pe c ie s ) S

ontinued Fig. 1. Proportion of aphids from the pan trapping collection in Pennsylvania and New York that use her-

5. (C baceous plants, trees, or crops as primary hosts. Host associations for North America characterized from

ab le Blackman & Eastop (1994, 2000, 2006). T Papulaphis sleesmani (Pepper) Pleotrichophorus ambrosiae Hille Ris Lambers Pleotrichophorus asterifoliae (Strom) Pleotrichophorus glandulosus (Kaltenbach) Pleotrichophorus patonkus (Hottes & Frison) Pleotrichophorus wasatchii (Knowlton) Pseudacaudella rubida Börner porosum Rhopalosiphoninus latysiphon Rhopalosiphoninus (Myzosiphon) solani (Thomas) Rhopalosiphoninus staphyleae (Koch) Rhopalomyzus (Judenkoa) lonicerae (Siebold) Rhopalomyzus poae (Gillette) avenae 984 Florida Entomologist 97(3) September 2014 pruni setariae Pseudasiphonaphis corni (Tissot) cerasifoliae (Fitch) Rhopalosiphum enigma Hottes & Frison Rhopalosiphum musae (Schouteden) Rhopalosiphum nigrum Richards Rhopalosiphum oxyacanthae Rhopalosiphum parvae Hottes & Frison Rhopalosiphum rufiabdominale McVicar Baker Rhopalosiphum sanguinarium McVicar Sanbornia juniper Pergande ex Baker graminum Schizaphis nigra (Baker) . a ni enn s ylv P in

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Aphis (Protaphis) knowltoni Hottes & Frison Aphis lugentis Aphis maculatae Oestlund Aphis (Protaphis) middletonii Aphis neilliae Oestlund Boyer de Fonscolombe Aphis (Bursaphis) oenotherae Oestlund Aphis oenotherae sanborni Aphis oestlundi Aphis pawneepae Hottes Aphis pulchella Aphis rubicola Oestlund Aphis rubifolii Aphis rumicis Aphis sambuci (L.) Aphis spiraecola Aphis spiraephila Patch Aphis (Burspahis) varians Patch Aphis vernoniae Thomas Aphis viburniphila phidini A tri b e , phidin a e A mily s u bfa

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pe c ie s 6. S ab le T Aphis angelicae Lee & Seo Aphis asclepiadis Fitch Aphis caliginosa Hottes & Frison Aphis carduella Aphis cephalanthi Aphis coreopsidis (Thomas) Aphis cornifoliae Fitch Aphis debilicornis Gillette & Palmer Aphis decepta Hottes & Frison Aphis fabae Aphis farinosa Gmelin Aphis feminea Hottes Aphis folsomii Davis Aphis forbesi Weed Aphis frangulae Kaltenbach Aphis gerardiae (Thomas) Aphis glycines Aphis hamamelidis Pepper Aphis hederae Aphis illinoisensis Shimer Aphis impatientis Thomas Bachmann et al.: Alate Aphid Species on Snap Beans in NE USA 985 mucida (Fitch) (Zetterstedt) Hannabura alnosa (Pepper) (Lineomyzocallis) walshii (Monell) Neosymydobius albasiphus (Davis) Patchia virginiana Baker Protopterocallis fumipennella (Fitch) Protopterocallis gigantean Bissell Protopterocallis pergandei Bissell alnifoliae (Fitch) ononidis (Kaltenbach) Therioaphis riehmi Therioaphis trifolii forma maculate (Buckton) ulmifolii (Monell) Tinocallis punctatella Fitch Tuberculatus . a ni enn s ylv Calaphis (Cepegillettea) myricae (Patch) Callipterinella calliptera (Hartig) P Koch Euceraphis gillettei Davidson Euceraphis lineata Baker Myzocallis (Lineomyzocallis) bella (Walsh) Myzocallis (Castaneomyzocallis) castaneae (Fitch) Myzocallis (Agrioaphis) castanicola Baker Myzocallis coryli (Goetze) Myzocallis (Neomyzocallis) discolor (Monell) Myzocallis (Lineomyzocallis) exultans Boudreaux & Tissot Myzocallis (Lineomyzocallis) frisoni Boudreaux & Tissot Myzocallis (Lineomyzocallis) granovskyi Boudreaux & Tissot Myzocallis (Lineomyzocallis) longiunguis Boudreaux & TIssot Myzocallis (Lineomyzocallis) melanocera Boudreaux & Tissot Myzocallis (Lineomyzocallis) multisetis Boudreaux & Tissot Myzocallis (Neomyzocallis) punctata (Monell) Myzocallis (Lineomyzocallis) spinosa Boudreaux & Tissot Myzocallis (Neomyzocallis) tuberculata Richards in

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a l phidin e C mily s u bfa

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Species Betulaphis quadrituberculata (Kaltenbach) Calaphis alni Baker Calaphis betulaecolens (Fitch) Calaphis betulella Walsh Calaphis leonardi Quednau Chromaphis juglandicola (Kaltenbach) tiliae (L.) Hoplochaitophorus heterotrichus Quednau Hoplochaitophorus quercicola (Monell) Lachnochaitophorus obscurus (Tissot) Melanocallis caryaefoliae (Davis) caryella Monellia hispida Quednau Monellia microsetosa Richards Monelliopsis bisselli Quednau Monelliopsis caryae (Monell) Monelliopsis nigropunctata (Granovsky) Myzocallis alhambra Davidson Myzocallis (Neomyzocallis) asclepiadis (Monell) pe c ie s 7. S ab le T Tribe Calaphidini 986 Florida Entomologist 97(3) September 2014

dividuals (Figs. 2 and 3, at 1,250 individuals there would be 45 species sampled in Pennsyl- vania and 50 in New York). Based on the his- torical collections reported by Pepper, there are approximately 350 aphid species in Penn- sylvania. Historical reports in Leonard (1963) suggest that there are approximately 430 aphid species in New York. Combining the list of aphids collected from snap bean fields, peach orchards and those pub- lished by J. O. Pepper in 1965, we developed a new, more comprehensive list of the aphids present in Pennsylvania. We found 7 species present in our collections that were not present in the slide collection housed in the Frost Ento- mological Museum (University Park, Pennsyl- vania) or published in Pepper (1965) (Table 3). viminicola Hille Ris Lambers Periphyllus americanus Periphyllus californiensis (Shinji) Periphyllus lyropictus (Kessler) Periphyllus negundinis (Thomas) Sipha glyceriae One of these aphids, Aphis glycines Matsumura, was introduced to the US around the turn of the 21st century and is now widespread through- out the Midwest, Northeast and southeastern Canada (Ragsdale et al. 2011).

Discussion

Our passive trapping in snap bean fields alone yielded a surprisingly high percentage of the species present throughout Pennsylvania and New York (~14% and ~18% respectively). Our sampling method concentrated on only one . habitat (commercial snap bean fields), but did

a ni intercept aphids moving from the surround- ing forests and hedgerows. The high degree of landscape heterogeneity and in Chaitophorus populifolii Chaitophorus pusillus Hottes & Frison Chaitophorus saliniger Shinji Chaitophorus stevensis Sanborn Chaitophorus viminalis Monell the trapping areas includes plant species that enn s ylv

P serve as hosts for many of the aphid species in

that represented less than 1% of the total cap- ture (Pfleeger et al. 2006). These aphids were captured in very small numbers (mostly single- tons), and are not important contributors to the epidemics reported by Wallis et al. o cc urring (2005) and Nault et al. (2009). Of the aphids we captured, 2 species were especially notable; Therioaphis trifolii Monell, which comprised 31.8% of the identified aphids, and A. glycines which represented 18.2 % of the identified aphids. Both of these aphids were h a itophorin e C

Table 9. Species in the subfamily Species Chaitophorus longipesi Tissot Chaitophorus nigrae Oestlund Chaitophorus nigricentrus Richards Chaitophorus nudus Richards Chaitophorus populicola Thomas Chaitophorus populifoliae Oestlund Sipha (Rungsia) elegans del Guercio

mily s u bfa occurring in Pennsylvania.

the Drepanaphis Drepanaphis Drepanaphis in

acerifoliae nigricans spicata Smith Drepanaphis Drepanaphis Drepanosiphum carolinensis parva Smith platanoidis pe c ie s Drepanaphis Drepanaphis Shenahweum kanzensis Smith sabrinae Miller minutum (Davis) 8. S Drepanaphis Drepanaphis

ab le monelli Davis simpsoni Smith T Tribe Chaitophorini Siphini Bachmann et al.: Alate Aphid Species on Snap Beans in NE USA 987 Schizolachnus piniradiatae (Davidson) Schizolachnus parvus (Wilson) Eulachnus rileyi Eulachnus americanus Takahashi Eulachnus agilis (Kaltenbach) Essigella pini watsoni Tissot Cinara (Cupressobium) tujafilina (Del Guercio) Cinara taedae Tissot Eriosoma wilsoni Remaudière (Walsh & Riley) Kaltenbachiella ulmifusa (Walsh Tetraneura nigriabdominalis Tetraneura (L.) Tetraneura longianus Smith Prociphilus (Pulvius) probosceus (Sanborn) Prociphilus (Paraprociphilus) tessellatus (Fitch) Thecabius affinis (Kaltenbach) Thecabius (Parathecabius) gravicornis (Patch) Thecabius populimonilis (Riley) Species Species . . a ni Cinara strobe Fitch salignus (Gmelin) Tuberolachnus Cinara spiculosa Bradley Cinara pruinosa (Hartig) Cinara pinivora (Wilson) Cinara pinea (Mordvilko) Cinara pilicornis (Hartig) Cinara pergandei (Wilson) Cinara laricis (Hartig) Cinara laricifex (Fitch) Cinara juniperivora (Wilson) a ni enn s ylv P in enn s ylv

P Eriosoma (Mimaphidus) lanuginosum (Hartig) Eriosoma mimicum Hottes & Frison Eriosoma rileyi Thomas Smynthurodes betae Westwood Pemphigus populitransversus Pemphigus populivenae Prociphilus americanus (Walker) Prociphilus caryae (Fitch) Prociphilus caryae spp. fitchii Baker & Davidson Prociphilus (Neoparacletus) corrugatans (Sirrine) Prociphilus erigeronensis (Thomas) Prociphilus (Meliarhyzophagus) fraxinifolii in

o cc urring

o cc urring

tin a e rio s om a ac hnin a e E L mily s u bfa mily s u bfa

Cinara juniperi De Geer caryae (Harris) rara Mordvilko Lachnus allegheniensis McCook Cinara harmonia Hottes Cinara gracilis (Wilson) Cinara fornacula Hottes Cinara (Cupressobium) cupressi (Buckton) Cinara costata (Zetterstedt) Cinara canatra Hottes & Bradley Cinara braggii (Gillette) Cinara banksiana Pepper & Tissot Cinara atlantica the the

Colopha graminis (Monell) Colopha ulmicola (Fitch) Eriosoma americanum (Riley) Eriosoma crataegi (Oestlund) Geoica ultricularia group Forda marginata Koch Grylloprociphilus imbricator (Fitch) Mordwilkoja vagabunda (Walsh) Neoprociphilus aceris (Monell) Pachypappa pseudobyrsa (Walsh) Pemphigus bursarius (L.) Pemphigus monophagus Maxson Pemphigus nortonii Maxson Pemphigus populicaulis in in

pe c ie s pe c ie s 10. S S 11. ab le ab le Tramini Lachnini T Tribe Eriosomatini T Tribe Eulachnini Fordini Pemphigini 988 Florida Entomologist 97(3) September 2014

ew N

7.5 2.9 0.2 0.2 0.3 1.4 0.9 1.0 1.4 1.5 1.5 3.8 3.4 2.7 8.6 12.9 18.2 31.8 a nd 100.0 Percent of Total (2004–2006) Overall 17 15 29 77 86 a n 660 254 122 123 128 129 331 302 239 760 1134 1606 2809 8,821 Total enn s ylv P

in

f ield s

6.3 2.8 1.1 1.0 1.0 3.4 0.0 0.5 1.9 0.0 0.1 9.2 0.0 5.6 2.1 9.8 15.0 40.0 b e a n 100.0

s n a p

Percent of Total 0 7 0 1 0 84 38 15 14 14 45 26 75 28 201 123 131 535 Pennsylvania 1,337 c ommer i a l

in

Total tr a p s

a n p

ter a w

7.7 2.9 0.2 1.0 1.0 1.1 1.3 1.7 1.7 1.7 2.4 3.2 9.2 14.8 19.7 30.4 <0.1 <0.1 100.0 . (2009). f rom

a l

et

Percent of Total ca pture

a ult 2 1 New York N the 15 77 77 79 97

576 216 128 128 130 179 239 685 in 7,484 o f 1,106 1,475 2,274 1 Total ab le > 1 % 1 > T f rom

erived repre s enting

s pe c ie

a phid

(2002–2006). D te ork l a Y 1. A 1. *Published in Nault et al (2009) as (Kaltenbach). ab le Total Others Unknown Brachycaudus persicae (Passerini) Anoecia sp. Aphis fabae Scopoli Rhopalosiphum padi (L.) Aphis sp. Capitophorus eleagni (Del Guerica) (Sulzer) (Davis) * Hayhurstia atriplicis (L.) Aphis gossypii Glover Aphis craccivora Koch Pemphigus populicaulis Fitch Rhopalosiphum maidis (Fitch) Acyrthosiphon pisum (Harris) Aphis glycines Matsumura Therioaphis trifolii (Monell) T Species Bachmann et al.: Alate Aphid Species on Snap Beans in NE USA 989

a nd ho s t

all HPS HPS HPS HPS HPS HPS HPS HPS HPS HPS HPS HPS AWT AWT AWT AWT AWT AWT AWT AWC AWC AWC AWC AWC AWC AWC AWC AWC Source s e c ond a ry (2004–2006)

a nd a ni

rim a ry enn s ylv P P

in

Secondary Host for Aphid Secondary Host for . 2005). hps ]). a l Gramineae Gramineae Oenothera biennis Umbelliferae polyphagous Glycine max polyphagous, , cucurbits wide range Scrophulariaceae tubuliferous Compositae Polygonum spp., Persicaria spp. f ield s

et

b e a n 2006 [

a lli s a nd W s n a p

], (2003–2004, c ommer i a l ], 2000 [ awc

Cuscuta in

a ni awt tr a p s

a n enn s ylv (1994 [ p P

Primary Host for Aphid Primary Host for

ter a as top w

c entr a l & E

in Leguminosae, Trifoleae, Loteae Leguminosae, Trifoleae, Fabae, Leguminosae, Genistae, Trifoleae, Hedysareae Rubus spp. Cornus sanguinea Cornus spp. Cornus spp. Cornus stolonifera Cephalanthus occidentalis polyphagous, Leguminosae opulus Euonymous europaeus, Viburnum Fragaria spp. spp. Catalpa, Hibiscus, Celastrus, Rhamnus, Punica Hedera helix , Araliaceae, Senecio spp., Erigeron sp. Compositae, Cruciferae, Umbelliferae, Grami - nae Rhamnus cathartica , R. alnifolia Oenothera biennis Pyroidea Euphorbia Rubus spp. Rumex spp., Rheum spp. , Spiraea spp., polyphagous spp. Viburnum Polyphagous armeniaca Prunus persica , P. Cruciferae Elaeagnus spp. Elaeagnaceae f rom

l ac km a n • • • • • • • • • • • • • B PA Peach or c h a rd s

c olle ted , f rom

• • • • • • • • • • • • • • • • • • • • • • • NY pe ac h

tion s a ken in

t Crop Sampled

• • • • • • • • • • • • • • • • PA Snap bean ass o c i a tr a p s

meri ca A ho s t

s imil a r

orth with

N f rom

f or

a phid s a nd

te tion s a l

o f

ass o c i a

(2002–2006), pe c ie s N y pl a nt 2. S 2. ab le Aphid Species T Acyrthosiphon kondoi Shinji Acyrthosiphon pisum Harris Amphorophora rubi Kaltenbach Anoecia corni (Fabricius) Anoecia cornicola (Walsh) Anoecia oenotherae Wilson Aphis carduella Walsh Aphis cephalanthi Thomas Aphis craccivora Koch Aphis fabae Scopoli Aphis forbesi Weed Aphis glycines Matsumura Aphis gossypii Glover Aphis hederae Kaltenbach Aphis lugentis Williams Aphis (Protaphis) middletonii Thomas Aphis nasturtii Kaltenbach Aphis oestlundi Gillette Aphis pomi DeGeer Aphis pulchella Hottes & Frison Aphis rubifolii (Thomas) Aphis rumicis L. Aphis spiraecola Patch Aphis viburniphila Patch Aulacorthum solani (Kaltenbach) Brachycaudus (Scrophulaphis) persicae group (L.) Capitophorus elaeagni (del Guercio) Capitophorus hippophaes (Walker) 990 Florida Entomologist 97(3) September 2014

a nd

a ni HPS HPS AWT AWT AWT AWT AWT AWT AWT AWT AWT AWT AWT AWT AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC Source rim a ry enn s ylv P P

in

. 2005). hps ]). a l

f ield s

et

2006 [ b e a n

a lli s a nd W ], s n a p

Secondary Host for Aphid Secondary Host for Labiatae Graminieae highly polyphagous, Solanaceae spp. communis, Arundo communis, Phragmites donax Umbelliferae spp. Plantago spp. Gramineae (2003–2004, ], 2000 [ awc c ommer i a l

in a ni

awt tr a p s

enn s ylv (1994 [ a n p P

ter as top Chenopodium spp. a w

& E c entr a l

in

f rom

l ac km a n B Primary Host for Aphid Primary Host for or c h a rd s

c olle ted f rom ,

pe ac h

Rhus glabra , R. typhina Cruciferae Liriodendron tulipifera Ulmus rubra Artemisia ludoviciana, A. vulgaris Rosaceae , Rosa spp. Prunus domestica Dendranthema indicum, morifolium, frutes - cens , Compositae Rosa spp Ribes spp. Chenopodiaceae, Atriplex, Prunus domestica, P. armeniaca Prunus domestica, P. Pinus spp. Lonicera spp. many monocots Pyroidea, , Crataegus, Coloneaster Pinus spp. Malus spp., Pyrus Acer pseudoplatanus, spp., sycamore Acer saccharum Acer rubrum Acer saccharum, A. rubrum Acer saccharinum, A. rubrum, saccharum Pinus spp. spp. a ken tion s in t

• • • • • • • • • • tr a p s PA

ass o c i a

meri ca Peach A ho s t

• • • • • • • • • • • • • • • • s imil a r NY

orth N with Crop Sampled

• • • • • • • • • • PA f rom

Snap bean f or

a nd a phid s

tion s te a l

ass o c i a o f

(2002–2006), pl a nt N y

pe c ie s a nd ) 1 S ho s t

ontinued (2004–2006) s e c ond a ry 2. (C ab le (Walsh & Riley) Kaltenbachiella ulmifusa (Walsh Lipaphis pseudobrassicae (Davis) Macrosiphoniella ludovicianae (Oestlund) Macrosiphoniella sanborni (Gillette) Illinoia liriodendri (Monell) Macrosiphum pallidum (Oestlund) Hysteroneura setariae (Thomas) Macrosiphum euphorbiae (Thomas) (L.) Geoica squamosa Hart Hayhurstia atriplicis (L.) Hyadaphis foeniculi (Passerini) (Geoffroy) Eulachnus rileyi (Williams) Dysaphis tulipae (Boyer de Fonscolombe) Eriosoma lanigerum (Hausmann) Essigella pini (Wilson) Dysaphis (Pomaphis) plantaginea (Passerini) Drepanosiphum platanoidis (Schrank) Drepanaphis sabrinae Miller Drepanaphis nigricans Smith Drepanaphis carolinensis Smith Aphid Species Cinara atlantica (Wilson) Drepanaphis acerifoliae (Thomas) T Carolinaia (Glabromyzus) rhois (Monell) Chaitophorus populifolii Essig Bachmann et al.: Alate Aphid Species on Snap Beans in NE USA 991

a nd a ni

AWT AWT AWT AWT AWT AWT AWT AWT AWT AWT AWT AWT AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC AWC Source rim a ry enn s ylv P P

in

]). . 2005). hp s a l

f ield s

et

2006 [ b e a n

a lli s a nd W s n a p ],

Secondary Host for Aphid Secondary Host for Dipsacaceae mosses polyphagous, over 40 families Leguminosae unknown Labiatae esp spp. Trifolium unknown Cruciferae Chenopodiaceae grasses Gramineae, Cyperaceae, Solanaceae water plants Gramineae Humulus lupulus (hops) grasses (2003–2004, ], 2000 [ awc c ommer i a l

in a ni

awt tr a p s

enn s ylv (1994 [ a n P p

ter as top a w

c entr a l & E

in

Primary Host for Aphid Primary Host for f rom

l ac km a n Ostrya virginiana, Corylus spp. Rosa spp. Rhus spp. ( glabra, typhina ) Fagaceae Carya spp. Prunus persica, spp. Crataegus, Cydonia, Malus, Pyrus Crataegus Crateagus spp. Crataegus spp tremuloides Populus deltoides , P. Populus spp. Populus spp. Alus, Pyrus, Cotoneaster, Crataegus, Alus, Pyrus, Cotoneaster, Sorbus Prunus spp. Gramineae Acer spp. Gramineae Prunus spp. Prunus virginiana Gramineae Ace r spp., Aesculus spp. Prunus spp. Gladiolus), runners bulbs (Tulipa, Fraxinus spp. Populus spp., Salix spp. Populus spp., Salix spp. Lonicera alpigena B or c h a rd s

c olle ted f rom ,

• • • • • • • • • • • • • • • • • PA pe ac h

Peach a ken tion s in

t

• • • • • • • • • • • • • • • • • • • • • • • NY tr a p s

ass o c i a

meri ca Crop Sampled • • • • • • • • • • • A PA Snap bean ho s t

s imil a r

orth N with

f rom

f or

a nd a phid s

tion s te a l

ass o c i a o f

(2002–2006), pl a nt N y

pe c ie s a nd ) 2 S ho s t

ontinued s e c ond a ry (2004–2006) 2. (C ab le T Aphid Species Macrosiphum (Neocorylobium) pseudocoryli (Patch) Macrosiphum rosae (L.) Melaphis rhois (Fitch) Monellia caryella (Fitch) Myzus (Nectarosiphon) persicae (Sulzer) Myzocallis sp. Nearctaphis bakeri (Cowen) Nearctaphis clydesmithi Hille Ris Lambers Nearctaphis crataegifoliae (Fitch) Ovatus crataegarius (Walker) Pemphigus populicaulis Fitch Pemphigus populitransversus Riley Pemphigus populivenae Fitch Periphyllus americanus Baker Rhopalosiphum maidis (Fitch) Rhopalosiphum nymphaeae (L.) Rhopalosiphum oxyacanthae (Schrank) Rhopalosiphum padi (L.) Rhopalosiphum rufiabdominale (Sasaki) (Rondani) Sipha flava (Forbes) Periphyllus testudinaceus (Ferni) Phorodon humuli (Schrank) Prociphilus (Meliarhizophagus) fraxinifolii (Riley) Pterocomma bicolor (Oestlund) Pterocomma smithiae (Monell) (Sanderson) Rhopalomyzus poae (Gillette) Rhopalosiphoninus latysiphon (Davidson) 992 Florida Entomologist 97(3) September 2014

a nd

a ni HPS HPS HPS HPS AWT AWC AWC AWC Source rim a ry enn s ylv P P

in

]). . 2005). . hp s a l

f ield s

et

2006 [ b e a n Cyperus spp. Gramineae

a lli s a nd W ], s n a p

Secondary Host for Aphid Secondary Host for (2003–2004, ], 2000 [ awc c ommer i a l

in

a ni awt tr a p s

enn s ylv (1994 [ a n P p

ter as top a w

& E c entr a l

Rhododendron spp. Crataegus spp. Leguminoseae Compositae, Lactuca spp. Aster novaeangliae Gramineae Ulmus spp. Melilotus spp. in

f rom Primary Host for Aphid Primary Host for

l ac km a n B or c h a rd s

c olle ted f rom ,

• • PA pe ac h

Peach a ken tion s in t

• • • • • • • NY tr a p s

ass o c i a

meri ca Crop Sampled • • • A PA Snap bean ho s t

s imil a r

orth N with

f rom

f or

a nd a phid s

tion s te a l

o f ass o c i a

(2002–2006), pl a nt N y pe c ie s

a nd ) 3 S ho s t

ontinued s e c ond a ry (2004–2006) 2. (C ab le Vesiculaphis caricis (Fullaway) Vesiculaphis Utamphorophora crataegi (Monell) Therioaphis (Pterocallidium) trifolii (Monell) Uroleucon (Lambersius) anomalae (Hottes & Frison) Uroleucon pseudambrosiae (Olive) (Fabricius) nigriabdominalis (Sasaki) Tetraneura Therioaphis (Rhizoberlesia) riehmi (Börner) T Aphid Species Bachmann et al.: Alate Aphid Species on Snap Beans in NE USA 993

may be transient. If this is the case, migrating aphids may be scrubbed of virions when they land in one of the many bordering forests con- taining many non-host plants. The Pepper (1965) aphid list in addition to the Pepper slide collection allowed us to com- pile a comprehensive list of the aphids present in Pennsylvania, but the nomenclature was in need of updating. Our efforts to update the no- menclature, and incorporate our more recent sampling efforts, resulted in a modern list of aphids of Pennsylvania that includes recently introduced species.

Acknowledgements Fig. 2. Individual-based rarefaction curves showing aphid species accumulation in Pennsylvania and New The authors would like to thank Fred Gildow, York. William Sackett and Dana Roberts. Funding for this work was provided by the Pennsylvania Vegetable Growers Association. introduced to North America (A. glycines from Asia and T. trifolii from Europe) and were quite References Cited destructive to crops immediately after their in- troduction (in soybean and alfalfa, respective- Agrios, G. N. 2005. , 5th ed. Elsevier ly). Aphis glycines continues to cause signifi- Academic Press. Burlington, MA. 922 pp. cant economic damage in soybean (Ragsdale et Blackman, R. L., and Eastop, V. F. 1994. Aphids on al. 2011). While not known to colonize Phaseo- the World’s Trees: An Identification and Informa- tion Guide. J. Wiley & Sons. Chichester, UK. 987 lus spp., both species are competent vectors of pp. the legume strain of CMV (Gildow et al. 2008). Blackman, R. L., and Eastop, V. F. 2000 Aphids on The intermittent appearance of CMV in cen- the World’s Crops: An Identification and Informa- tral Pennsylvania snap bean crops could be tion Guide, 2nd ed. J. Wiley & Sons. Chichester, influenced by a unique agricultural landscape. UK. 250 pp. Agricultural fields are located in valleys bor- Blackman, R. L., and Eastop, V. F. 2006. Aphids on dered by the low, but steep, forested ridges of the World’s Herbaceous Plants and Shrubs (Vol- the Appalachian Mountains. The ridge and val- ume 1 Host Lists and Keys; Volume 2, The Aphids). ley system might be acting like a barrier, keep- J. Wiley & Sons. Chichester, UK. 1439 pp. Blackman, R. L., and Eastop, V. F. Aphids on the ing CMV out for most of the season. We did not World’s Plants. Available from http://www.aphid- search for a CMV reservoir outside of testing sonworldsplants.info/. a few alfalfa fields, which were also negative Colwell, R. K. 2005. EstimateS: Statistical estima- for CMV. It is possible, that much like our A. tion of species richness and shared species from glycines population, legume strains of CMV samples Version 8.2.0. Available from http://purl. oclc.org/estimates. Damsteegt, V., Stone, A. L., Luster, D. G., Gildow, F. E., Levy, L., and Welliver, R. 2001. Prelimi- nary characterization of a North American isolate of plum pox virus from naturally occurring peach and plum orchards in Pennsylvania, USA. Acta Hort. 550: 145-152. Dixon, A. F. G. 1985a. Aphid . Blackie & Son Ltd. London, UK. 157 pp. Dixon, A. F. G. 1985b. Structure of aphid popula- tions. Annu. Rev. Entomol. 30(1):155-174. Dixon, A. F. G., Kindlmann, P., Leps, J., and Hol- man, J. 1987. Why are there so few species of aphids, especially in the tropics. American Nat. 129:580-592. Favret, C. Aphid Species File Version 1.0/4.1. Avail- able from http://aphid.speciesfile.org. Gildow, F. E., Shah, D. A., Sackett, W. M., But- Fig. 3. Individual-based rarefaction curve showing zler, T., Nault, B. A., and Fleischer, S. J. 2008. aphid species accumulation from the combining of sam- Transmission efficiency of ples from PA and NY (solid line) and the 95% confidence by aphids associated with virus epidemics in snap intervals for the curve (dashed lines). bean. Phytopathology 98: 1233-1241. 994 Florida Entomologist 97(3) September 2014

Gray, S. M., and Banerjee, N. 1999. Mechanisms of Pfleeger, T. G., Olszyk, D., Burdick, C. A. King, transmission of plant and viruses. G., Kern, J., and Fletcher, J. 2006. Using a geo- Microbiol. Mol. Biol. Rev. 63:128-48. graphic information system to identify areas with Grimaldi, D., and Engel, M. 2005. The Evolution of the potential for off-target pesticide exposure. Envi- . Cambridge University Press. Cambridge, UK. ron. Toxicol. Chem. 25(8): 2250-2259. 772 pp. Smith, C., Eckel, R., and Lampert, E. 1992. A key to Jones, R. A. C., Coutts, B. A., Latham, L. J., and Mck- many of the common alate aphids of North Caroli- irdy, S. J. 2008. Cucumber mosaic virus infection of na (Aphididae: ). North Carolina Agric. chickpea stands: temporal and spatial patterns of Res. Serv. Tech. Bull. 299. spread and yield-limiting potential. Plant Pathol. 57: Ragsdale, D. W., Landis, D. A., Brodeur, J., Heim- 842-853. pel, G. E., and Desneux, N. 2011. Ecology and Larsen, R. C., Miklas, P. N., Eastwell, K. C., Grau, C. Management of the in North Amer- R., and Mondjana, A. 2002. A virus disease complex ica. Annu. Rev. Entomol. 56: 375-399. devastating late season snap bean production in the Ragsdale, D. W., Voegtlin, D. J., and O’neil, R. J. Midwest. Annual Report of the Bean Improvement 2004. Soybean aphid in North America. Coop. 45: 36-37. Ann. Entomol. Soc. America 97(2): 204-208. Leonard, M. D. 1963. A list of the aphids of New York. Rosales, M., Hinrichsen, P., and Herrera G. 1998. Proc. Rochester Acad. Sci. 10: 289-428. Molecular characterization of plum pox virus iso- Miller, G. United States National Collection of Aphi- lated from , plums, and in Chile. didae. Available from http://www.sel.barc.usda.gov/ Acta Hort. 472: 401-405. aphid/aphframe.htm. Roy, S., and Smith, I. M. 1994. Plum pox situation in Nault, B. A., Shah, D. A., Dillard, H. R., and Mcfaul, Europe. European Plant Prot. Org. (EPPO) Bull. A. C. 2004 Seasonal and spatial dynamics of alate 24: 515-523. aphid dispersal in snap bean fields in proximity to al- Wallis, C. M., Fleischer, S. J., Luster, D., and falfa and implications for virus management. Environ. Gildow, F. E. 2005. Aphid (Hemiptera: Aphididae) Entomol. 33:1593-1601. species composition and potential aphid vectors of Nault, B. A., Shah, D. A., Straight, K. E., Bachmann, A. plum pox virus in Pennsylvania peach orchards. J. C., Sackett, W. M., Dillard, H. R., Fleischer, S. J., Econ. Entomol. 98: 1441-50. and Gildow, F. E. 2009. Modeling temporal trends in Webb, S. E., Kok-Yokomi, M. L., and Voegtlin, D. J. aphid vector dispersal and cucumber mosaic virus epi- 1994. Effect of trap color on species composition of demics in snap bean. Environ. Entomol. 38:1347-59. alate aphids (Homoptera: Aphididae) caught over Ng, J. C. K., and Falk, B. W. 2006. Virus-vector inter- plants. Florida Entomol. 77: 146-154. actions mediating nonpersistent and semipersistent Zitter T. A., and Murphy, J. F. 2009. Cucumber transmission of plant viruses. Annu. Rev. Phytopathol. mosaic virus. The Plant Health Instructor. Avail- 44: 183-212. able from http://www.apsnet.org/edcenter/intropp/ Pepper, J. O. 1965. A list of the Pennsylvania Aphididae lessons/viruses/Pages/Cucumbermosaic.aspx. Ac- and their host plants (Homoptera). Trans. American cessed 28-V-2012. Entomol. Soc. 91: 181-231. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

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