North American Allien Aphid Species

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30 June 2006PROC. ENTOMOL. SOC. WASH.108(3), 2006, pp . 583-610

ADVENTIVE APHIDS (HEMIPTERA: APHIDIDAE)O F AMERICA NORTH O F MEXICO

(RGF, EM) Agriculture and Agri-Food Canada, Canadian National Collectionof Insects, Ottawa, ON, K1A 0C6 Canada (e-mail: [email protected];[email protected]); (SEH) Florida Department of Agriculture and ConsumerServices, Division of Plant Industry, Gainesville, FL 32614, U.S.A. (e-mail:[email protected]); (GLM, LMR) Systematic Entomology Laboratory, PlantScience Institute, Agricultural Research Service, U.S. Department of Agriculture,Beltsville, MD 20705, U.S .A. (e-mail: [email protected])

Abstract.-We provide a compilation of 262 species of aphids that are consideredas adventive to North America north of Mexico. Included for each species, whereapplicable, is reference to: the location and date of introduction of the first NorthAmerican record; pest status in North America; principal economic hosts; andbiogeographical origin. Information is also provided for species whose presence inNorth America is considered erroneous or questionable and for those species that areconsidered Holarctic or Beringian.Key Words: Aphidoidea, plant lice, plant pathogens, invasive species, biological

control, quarantine, agriculture, forestry, horticulture, ornamentals,fruit trees

In an era of a global economy andcommerce, the increase of adventivespecies represents both domestic andinternational concerns. If non-indige-nous species become pests, the economicconsequences include loss of production,diminished quality, production cost in-creases, flexibility decreases in produc-tionlmanagement decisions, increasedrisk of human disease, and damagingenvironmental and aesthetic effects (Hu-ber et al. 2002). Various insects ofagricultural concern have been identifiedas potentially damaging if introduced or

1 I t is noteworrhy that Louise Russell, a pioneer womanenmrnologist an d special ist in the S~ ern orr h~ nch a ,ontinuesto publish at the age of 101.

reintroduced into the United States(Huber et al. 2002).

Many aphid species have been addedto the North American fauna sincecolonization by Europeans, and someof these have become serious pests. Inaddition to direct feeding damage,aphids constitute one of the largestgroups of known plant-virus vectors(Chan et al. 1991). Aphids are primarilycontrolled through pesticide applica-tions, adding to crop production costs,and having environmental implicationsassociated with increased pesticide use.The effects of some non-indigenousaphids in North America have beenstaggering. For example, by the early199OYs, he Russian wheat aphid, Diur-


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58 4 PROCEEDINGS OF TH E ENTOMOLOGICAL SOCIETY OF WASHINGTONaphis noxia (Kurdjumov), was responsi-ble for $500-900 million cumulativelosses in the western U.S. (Bernal et al.1993, Morrison and Peairs 1998). Theappearance of the brown citrus aphid,Toxoptera citricida (Kirkaldy), in Flor-ida in 1995 had a major impact on theU.S. citrus industry (Halbert and Brown1996). More recently, losses related tothe soybean aphid, Aphis glycines Mat-sumura, have been estimated at about$120 million in 2003 for Minnesota alone(Anonymous 2005).

The origins of this work actuallybegan in part in the 1970's when one ofus (LMR) contributed data on invasivespecies in the United States (e.g., Sailer1978). Nearly 30 years later, adventiveinsects are still a major concern foragriculture in North America. The ob-jectives of this paper are to: 1) list theadventive aphid species (excluding adel-gids and phylloxerans) of North Americaand record data on first detection date(along with validation source), principalhosts, and zoogeographic area of origin;2) analyze these data and determine ifpatterns exist; and 3) compare these datato other sternorrhynchan groups.

We use the term adventive species asdefined by D.R. Miller et al. (2002), i.e.,for those species that are non-native toNorth America regardless of economicharm. Other terms such as invasive,alien, exotic, and immigrant have some-times been used interchangeably (seeWheeler and Hoebeke (2001) and Milleret al. (2005) for discussion). Using Smithand Parron's (1978) list of North Amer-ican aphids as the base, we compileda new list for North American north ofMexico. We exclude species that havebeen intercepted at ports-of-entry ordiscovered in nurseries or greenhousesand subsequently eradicated. With theexception of Aphis chloris Koch, a bio-control agent for Hypericum perforatum

(Harris 2003), no aphid species has beenpurposely introduced.Geographic origins are given in gen-

eral terms only, relying heavily on Black-,

man and Eastop (1994, 2000), supple-mented by other references, as noted inTable 1, and by opinion of the authors.The origins of a few cosmopolitanspecies are unknown. It is not alwaysreadily apparent whether a species isadventive or native; some species wehave listed as adventive may in fact benaturally Holarctic in distribution andvice versa.For determining the earliest record ofthe North American adventive aphids,

we relied on published records and onspecimens examined in The CanadianNational Collection of Insects, Nema-todes and Arachnids (CNC) and theEdith Patch Collection (CNC-Patch)housed at CNC, Ottawa, ON, Canada;the MacGillivray Collection (ACNB),Agriculture and Agri-Food Canada,Fredericton, NB, Canada; The FloridaState Collection of Arthropods (FSCA),Gainesville, FL, U.S.A.; and The Na-tional Entomological Collection of theNational Museum of Natural History(USNM), Beltsville, MD, U.S.A. Inaddition, Asa Fitch's aphid species notesthat are currently housed with theUSNM aphid collection at Beltsville,MD, were also examined. This collectionof written records and observations fromthe 1840's-1870's provide some of ourearliest glimpses into the North Ameri-can aphid fauna. While there are somequestions concerning Fitch's "speciesconcept" versus current thinking forsome of his included species, there islittle confusion given the morphologyand associated host plants for others. Hisreference to the occurrence in 179 1 of thecabbage aphid, Brevicoryne brassicae (L.)is possibly the earliest record of anadventive aphid for North America.

Nomenclature followed herein reflectsthat of Remaudiere and Remaudiere


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VOLUME 108. NUM BER 3 585

Table 1. Introduced aphid species in Can ada and con tinental United States of America, giving earliestrecord with source of record, geographic origin, pest status, and economic host species. Aphid namesfollow Remaudikre and Remaudikre (1997). Earliest records are based on literature references and onspecimens in the following collections: USNM - National Museum of Natural History; CNC -Canad ian National Collection of Insects, Arachnids and Nematod es; FSCA- lorida State Collection ofArthropods; ACNB - MacGillivray collection, Agriculture and Agri-Food Canada, Fredericton, NB;CNC-Patch - dith Patch collection, housed at CNC; EMEC - Essig Museum of EntomologyCollection, University of California, Berkeley.

Earliest N.A. record (I) Geographic Origin/ Slatus io North America1 North AmericanIntroduced Species (Collection or Rererence) Rercrcnce (2 ) Reference (2) Economic Hosl(s)Acyrthosiphon caraganae SK, 1940 (CNC) Palearctic pest Caraganrr

(Cholodkovsky) nrborescensAcyrthosipl~or~ondoi CA, 1974 (Kono Palearctic (e Asia) pest Halbert et al. Medicago

Shinji 1977) 2000 srrtivaAcyrthosiphon lactucae MD, 1943 (USNM) Palearctic (Europe) Lactuca sativa

(Passerini)Acyrthosipl~onmalvae NE, 1891 (USNM) Palearctic (Euro pe) Geranirrm,

(Mosle ~) 3) Pelargoi~iumAcyrthosipl~onpisum KS, 1877 (Harper Palearctic (Europe) pest Leguminosae,(Harri s) et al. 1978) incl. Pisum,

MedicagoAloephagus mnyersi CA, 1939 (Essig 1950) Afrotropical minor pest Essig Aloe

Essig 1950, regulatory (greenhouse)issue

Amnpl~orophorrr MN, 1887 (Oestland Palearctic (Europe)anzpullata Buckton 1887)

Anoecin corni IA, 1897 (USNM) Palearctic (Europe) roots of(Fabricius) Gramineae,

incl. cerealsAphidou~zg~risn r ~ l i NC, 1999 (CNC) Palearctic (e Asia)

Takahashi (4)Apl~i s inara~zthi FL, 1990 (Halbert Neotropical potential pest edible

Holman et al. 2000) (Cuba) Halbert et al. 2000 Ainara~~thusAplzis callzozae BC, 1974 (Forbes and Palearctic (Europe)

Theobald Chan 1976)Aphis clzloris Koch BC, 1980 (Harris Palearctic Released as weed Hypericr~m

2003) (Europe) biological control perforatumagent

Aphis craccae Linnaeus BC, NB, ME, 1965 Palearctic Vicia cracca(CNC, ACNB)

Aphis cr~accivorrr NE, 1890 (USNM) Palearctic polyphagous,Koch (5) with

preferencefor legumes

Aphis cytisorurn Hartig DC, 1906 (USNM) Palearctic (Europe) Cy isusscoparius

Aphis epilobii Kaltenbach CO, 1915 (USNM) Palearctic (Europe)Aplzis errpl~orbiae CA, 1933 (USNM) Palearctic (Europe)

Kaltenbach (6)Aplzis frrbae Scopoli (7) NY, 1869 (Fitch 1870 Palearctic (Europe) pest polyphagous

[as A. rumicis L.])Aphis ge~zistaeScopoli MA, 1921 (USNM) Palearctic (Europe)Aphis glycines IA, IL, IN, KY, MI, Oriental and major pest Ragsdale Glycine max

Matsumura MN, MO, OH, WI, Palearctic (East et al. 2004WV, 2000 (Voegtlin Asia)2000)

Aphis gossypii Glover NY, 1852 (Russell Palearctic major pest polyphagous1968)


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586 P R O C E E D IN G S O F T H E E N T O M O LO G I C A L S O C I E TY O F W A S H I N G T ON

Table 1. Continued.Earliest N. A. record (I ) Geographic Origin1 Status in Nortll Amerieal North AmericanInlrodueed Species (Collection or Rererence) Rererence (2) Rcrercnce (2) Economic Host@)

Aphis hederaeKaltenbachAphis idaei van derGoot

Aphis nasturtiiKaltenbach

Aphis nepetaeKaltenbach

Aphis nerii Boyer deFonscolombe

Aphis polygonata(Nevsky) (8)Aphis pomi De Geer

Aphis praeterita WalkerAphis rumicis Linnaeus

(7)Aphis salicariae KochAphis sambuci Linnaeus

Aphis sedi KaltenbachAphis spiraecola PatchAphis iaraxicola

(Borner) (9)Aphis ihaliciri KochAphis umbrella (Borner)

Aphis urricnta GmelinAphis vaccini (Borner)Aploneura lentisci

(Passerini)

Asiphonella daciyloniiTheobald

Aspidaphis adjuvans(Walker)

Ararsaphis ngrijoliae(Ferris)

Atheroides serrulniusHaliday

Aulacor ihumcircunzflexum(Buckton)

Aulacorihun7 solani(Kaltenbach)

Betulaphis brevipilosaBorner

CA , 19 10 (Essig1910)BC, 1951 (Forbeset al. 1973)

NE, 1894 (USNM )ID, 1957 (Gittins

et al. 1976)IL, 1879 (Thomas

1879)IL, 1977 (FSCA)PA, 1844 (Haldeman

1844)

Palearctic (Europe)Palearctic (Europe)PalearcticPalearcticPalearctic (e Asia)Palearctic (Central

Asia)Palearctic (Europe)

ME, 1918 (Patch Palearctic (Europe)1918)Unknown (7) Palearctic (Europe)

minor pest Hedera helix

pest

pest

pest

ON , 1952 (CNC ) Palearctic (Europe)NY, 1847 (Fitch Palearctic (Europe)

notes [as A.sambucifoliae Fitch])

ME, 1911 (Patch Palearctic (Europe)191 1)

CO, 1907 Palearctic or major pest(Gillette 19 10) Orien tal

QC, 1983 (Cloutier Palearcticet al. 1986)

BC, 1999 (CNC ) Palearctic (Europe)NE, SD, 1891 Palearctic (Europe)

(USNM [as A.maliae Koch])

PA, 19 13 (Olsen Palearctic (Europe)1921)

BC, 2000 (CNC ) Palearctic (Europe)IL, 1879 (Tho mas Palearctic

1879 [as (Mediterranean )Rhizobius poneThomas])

AZ, 1947 (US NM) Palearctic (e Asia) potential pestCO, 1907 (Gillette Palearctic (Europe)

1917b)CA, 1921 (Ferris Palearctic (e Asia)

1921)QC, 1966 (CNC) Palearctic minor pestCO, 1907 (Gillette Palearctic pest

1908) (e Asia) orOriental

NY, 1879 (Riley and Palearctic major pestMonell 1879)

BC, 1959 (Forb es Palearctic (Europe)et al. 1974)

ornamentals(flowers)

Malus pumila,Pyruscommunis,Cydoniaoblonga

polyphagous

Gramineae,includingTrir curn

Cynodondaciylon

Gramineae,incl. cereals

polyphagous

polyphagous


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VOLU ME 108, NUMB ER 3 587Table I. Continued .

Earliest N.A. record ( I ) Geographic Origin/ Status in North America1 North AmcricanIntroduced Soecies ICollection or Reierence) Reference (2) Reference (2) Economic H o d s )Betulaphis I?elvetica Hille BC, 1926

Ris Lambers (Glendenning 1926)Betulaphis BC, 1946(ACNB)

quadrituberculata(Kaltenbach)

Bracl~ycaud us elichrysi OR, 1893 (USNM)(Kaltenbach)

Bracl~ycaud us ardui NY, 1887 (USNM)(Linnaeus)

Brachycatrdus persicae MD, DE, NJ, VA,(Passerini) MI, 1887 (Smith

1890)Brachycaudus CT 1913 (Patch

ruinexicolens (Patch) 1917)(10)Brachycaudus schlvartzi CA, 1975 (Eastop(Borner) 1975 )

Brachycaudus spiraeae ON, 1956 (CNC)(Borner)

Bracliycaudus WA, 1993 (FSCA)tragopogonis(Kaltenbach)

Brachycorynella asparagi NY, NJ, 1969(Mordvilko) (Leonard 1970,

Race 1970)Brevicoryne brassicae NY, 179 1 (Fitch

(Linnaeus) notes)Calaphis betulicola NY, 1854 (Fitch

(Kaltenbach) notes)Callipterir7ella calliptera OR, 1909 (USNM)

(Hartig)Callipterinella BC, 1975 (Forbes and

minulissima (Stroyan) Chan 1980)Capitophorus carduinus GA, 1948 (USNM)(Walker)

Capitophorus elaeagni SD, 1892 (USNM)(Del Guercio)

Capitophorus hippophaes DC, 1895 (USNM)(Walker)

Capitophorus pakansus IL, 1929 (Hottes andHottes and Frison Frison 1931)

Capitophorus horni ME, 1951 (Corpuz-Bomer Raros and Cook

1974)Cavariella aegopodii MN, 1903 (USNM)

(Scopoli)Cavariella archangelicae WA, 1953 (USNM)

(Scopoli)Cavariella pastinacae NY, 1855 (Fitch(Linnaeus) notes)

Cerataphis brasiliensis OH, 1917 (USNM)(Hempel)

Cerataphis lataniae NE, 1912 (USNM)(Boisduval)

Cerataphis orchidearum DC, 1923 (USNM)(Westwood)

Palearctic (Europe)Palearctic (Europe )

Palearctic major pest(Europe)

Palearctic pest(Europe)


Palearctic (Europe)


Palearctic

Prunus,Compositae

Prunus,Compositae

Prunus

Prunus persica

Palearctic

Palearctic locally serious pest Asparagus(Europe) Stoetzel 1990 officinalis

Palearctic pest Brassica cropsPalearctic (Europe)Palearctic minor pest Hajek Betula

(Europe) and Dahlsten 1986Palearctic minor pest Hajek Betula

(Europe) and Dahlsten 1987Palearctic potential weed Cirsum(Europe) biocontrol agent

Gassmann andKok 2002

Palearctic Potential virus(Europe) vector

Palearctic (Europe)Palearctic (Europe)Palearctic (Europe)


Palearctic (Europe,Iceland)Palearctic (Europe)

Daucus carota

Oriental pest palmsOriental pest palmsOriental pest greenhouse

orchids


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588 PROCEEDINGS O F THE ENTOMOLOGICAL SOCIETY O F WASHINGTON

Table 1. Continued.Earliest N. A. record (1) Geographic Origin1 Status in Nort h America1 North Americanlnlroduced Species (Collecti on or Reference) Reference (2) Refcrcnce (2) Econonlic Hos~(s)

Ceruropliis erioplzori UT, 1935 (Knowlton Palearctic (Europe)(Walker) and Smith 1936)C/znelosi[~horzoterztillrre NY, 1851 (Fitch Palearctic (Europe)(Walker) notes)

CO, 1916 (USNM)Clznetosiphon MN, 1907 (USNM) Palearctic (Europe)

tetrnrhodzrnz (Walker)Clznitophorz/s leztcor~zelcrs CA, QC, 1966 Palearctic (Europe)

Koch (USNM, Quednau1966)

Clzaitophorus popzrlinlbne ON, 1955 (CNC) Palearctic (Europe)(Boyer deFonsco lom be)

Clzrornnphis ugln~zdicoln CA, 1896 (USNM) Palearctic (Europe) pest Olsen 1974 Juglrrr?s regirr(Kaltenbach)

Cirmrrr cecli Mimeur NY 2004 (CNC, Palearctic (Europe) Cupressus,FSCA) Juriiperus

Cirznrn confinis (Koch) CO, 1920 (USNM) Palearctic AbiesCinrrro costnto ON, 1905 (USNM) Palearctic Picerr(Zetterstedt)

Cinara cupressi CA, 1916 Palearctic Cupressus,(Buckton) (Bartholomew Juniperus

1932)Cinarn jkesrri Blanchard FL, 1942 (Tissot unknown Cupress~~s,

1944 [as C. Jznziperuswacnsnssoe])

Cirzarn jur7ipn.i NY, 1902 (USNM) Palearctic (Europe) Juniperus(De Geer)

Ci~znrnoricis (Hartig) MA, 1921 (USNM) Palearctic (Europe) LorixCi~znrn ilicor~nis ME, 1906 (Patch Palearctic Picerr

(Hartig) 1912)Cinnrrr pirzea DC, 1919 (Wilson Palearctic Pirzzrs

(Mordvilko) 1919)Cirzarn preri~zosrr(Hartig) C O, 19 10 (Gillette Palearctic Picerr1917a [as Lochrzus

prrlr?ierae])Cinnrn lr~jcfilirin Del CA, 1914 (Davidson unknown

Guercio) 1914)Colornrlorr obrotarzi NB, 1964 (ACNB) Palearctic (Central

(Koch) Asia)Colororloo ~bsirzt/zii MB, BC, 1965 (CNC) Palearctic (Centra l

(Lichtenstein) Asia)Color~doorc/zillerre ON, 1962 (CNC) Palearctic (Central

Hille Ris Lambers Asia)Colorrrdorr rrrtenzisirre BC, 1965 (CNC) Palearctic (Centra l

(Del Guercio) Asia)Color~rrdorr zfor?zrrc~rlrrtn NE, 1890 (USN M) Palearctic (Central minor pest

(Wilson) Asia)Color~rrdorr rnzrrcetirzn RI , 1974 (Smith and Palearctic (Centra l

(Walker) Par ron 1978) Asia)Corylobizn?z avellrrnrre BC, 1983 (Forbes and Palearctic (Europe) pest(Schrank) Cha n 1984)

Cryptcryhis pone (Hardy) MB, 1965 (Robinson Palearctic (Europe)and Bradley 1965)

Cryptor?7yszr.r gnleopsidis ME, 1914 (Patch Palearctic (Europe)(Kaltenbach) 1914)

chrysanthemums

Corylusrrvellrrrzn


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Table 1 . Continued.Earliest N. A. record (I ) Geographic Origin1 Status in North America1 North AmericanIntroduced Species (Collection or Rererence) Reference (2) Rererenee (2) Economic Host(s)

Euthoracaphis CA, 1929 (Essig Palearctic (e Asia)umbellariae Essig 1932)

Fordaformicarir~ on CO, 1909 (USNM) Palearctic (Europe) roots ofHeyden Gramineae

Forda marginata Koch DC, 1885 (USNM) Palearctic (Europe) roots ofGramineae

Geoica setulosa ID, 1985 (FSCA) Palearctic (Europe) minor pest G.L. roots of(Passerini) Miller et al. 2002 Gramineae

Geoica utricularia IL, 1887 (USNM) Palearctic (Europe) roots of(Passerini) (1 5) Gramineae,

incl. cerealsGeopemphigusfloccosus FL, 1945 (USNM) Neotrop ical minor pest roots of

(Moreira) Zpomoeubututas

Glyphina betulue PA, 1945 (USNM) Palearctic (Europe)(Linnaeus)

Greenidea ficicola FL, 2002 (Halbert Oriental potential pest FicusTakahashi 2004)Greenidea psidii van der CA, 1998 (Gill Orienta l minor pest, MyrtusGoo t 1998a) regulatory pest

Hayhurstia atriplicis CA, 1887 (USNM) Palearctic (Europe) Chenopodium(Linnaeus) including

Ch. quinoaHyadaphis coriandri FL, 1997 (Halbert Palearctic (Centra l major pest Umbelliferae,

P a s ) et al. 2000) Asia) includingCoriundrum,Foeniculum

Hyadaphis foeniculi NY, 1879 (USNM) Palearctic (Central minor pest Lonicera ;(Passerini) Asia) Umbelliferae

Hyadaphis tartaricae IL, 1979 (Voegtlin Palearctic (Centra l major pest Voegtlin Lonicera(Ajzenberg) 1981) Asia) 1981 tatarica

Hyalopteroides humilis DC, MA, 1909 Palearctic(Walker) (Hayhurs t 1909)

Hyalopterus amygdali IL, NJ (date ?) (16) Palearctic (e Asia) pest Prunus (s.g.(Blanchard) (1 6) (Smith and Par ron Amydalus)1978)

Hyalopterus pruni IL , 1879 (Thomas Palearctic (Europe) pest Prunus (s.g.(Geoffroy) (16) 1879) Prunus)

Hyperomyzus FL, 1997 (Halbert Palearctic (e Asia)carduellinus et al. 2000)(Theobald)

Hyperomyzus lactucae IL, 1878 (Thomas Palearctic (Europe) minor pest Ribes(Linnaeus) 1879)

Hyperomyzus pallidus ME, 19 14 (Hille Ris Palearctic (Europe) RibesHille Ris Lambers Lambers 1949)

Hyperomyzus picridis PA, 1942 (USN M) Palearctic (Europe) potential pest Ribes(Borner)

Hyperomyzus rhinanthi ME, 1951 (CNC) Palearctic (Europe) Ribes(Schouteden) (1 7)

Zdiopterus nephrelepidis IL, 1907 (Davis 1909) Neotropical pest Stoetzel 1990 greenhouseDavis Stoetzel 1990 fernsJacksonia papillata ID, 1959 (USNM) Palearctic (Europe)

TheobaldLiosomaphis berberidis NY, 1846 (Fitch Palearctic (Europe)

(Kaltenbach) notes)Lipamyzodes matthiolae WA, 1976 (Fox 1977) Palearctic (Europe)

(Doncaster)


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VOLU ME 108, NUM BER 3 59 1Table I . Continued

Earliest N.A. rccord ( I ) Geographic Origin1 Status in North America1 North AmcricanIntroduced Species (Collection or Reference) Reference (2) Rcference (2) Economic Host(s)Lipaphis pseudobrassicae

(Davis) (1 8)Longicnudus trirlzodus(Walker)

Macrosiphoniellaabrotani (Walker)

Macrosiphoniellaabsinthii (Linnaeus)

Macrosiphoniellaarlemisiae (Boyer deFonscolombe)

Macrosiphoniellagrandicauda Takahashiand Moritsu

Macrosiphoniellaleucantlzemi(Ferrari)

Maoosiphoniellamillefolii (De Geer)Macrosiphoniella

sanborni (Gillette)

Macrosiphoniellasubterranea (Koch)

Macrosiphoniellatanace aria(Kaltenbach)

Mao.osiphrrm lilii(Monell) (19)

Macrosiphum rosae(Linnaeus)Mucrosiphurn stellariae

TheobaldMelanaphis bambusae

FullawayMelanaphis sacchari

(Zehntner)Melanaphis sorini

Halbert andRemaudikre

Metopolophiumdirhodum (Walker)

Metopolophium festucae(Theobald)Microlophium carnosum(Buckton) (20)

Microlophium sibiricurn(Mordvilko) (20)

DC, 1899 (USNM)MA, 1908 (USNM)IL, 1929 (USNM)IL , 1879 (Thomas

1879)NB, 1950 (ACNB)

MD, 1997 (USNM)

PA, 1948 (USNM)

CT, 1937 (EMEC)NE, 1891 (USNM)

PA, 1948 (USNM)

CT, 1909(CNC-Patch)

NY , 1878 (Comstock1880)

MA, 1841 (Harris1841)BC, 1980 (Forbes

and Chan 1983)GA, 1914 (USNM)FL, 1977 (FSCA)

FL via CA, 1996(Halbert andRemaudikre 2000)

UT, 1910 (USNM)

CA, 1970 (USNM,Eastop 1971)NE, 1889 (Williams1911)

PA, 1963 (USNM,Pepper 1965)

Palearctic (Europe) major pest Paddock Cruciferae1915Palearctic (Europe) Rosa

PalearcticPalearcticPalearctic

Palearctic

Palearctic

Palearctic

Leucanthemumvulgare

Palearctic (e Asia) pest Asteraceae(esp.chrysanthe-mums)

Palearctic cultivatedchrysanthe-mums

Palearctic minor pest cultivatedchrysanthe-mums(occa-sionally)

Palearctic? pest Cultivated lilies

Palearctic (Europe) pestPalearctic pestPalearctic (e Asia)

Rosaceae (esp.Rosa)Caryophyllaceae

(esp. Dianthus)bamboos

Oriental pest White et al. Gramineae2001 (esp. sugar-

cane)Oriental Halbert pest Halbert and Miscanthus

and Remaudikre Remau diire 20002000

Palearctic (Europe) major pest Gramineae,incl. cereals;Rosa

Palearctic (Europe) pest GramineaePalearctic (Europe) UrticaPalearctic (Europe) Urtica


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592 P R O C E E D I N G S O F T H E E N T O MO L O GI C A L S O C I ET Y O F W A S H I N G T O N

Table 1. Contin ued.Earliest N.A. rccard ( 1 ) G e a g r a p l ~ i cOrigin/ S ta tu s in N o r t h Amcrical N o r t h Amcric;tnl~~ l roducedpccies (Collection or Rcfcrcncc) Reference ( 2 ) Refercncc (2) Econonl ic H o s l ( s )

M i r ~ c I r ~ ~ s~bietiizzr171Koch (21)

M o r ~ ~ ~ h i sriterzrza ta(Kaltenbach)

Mir.scrrphis I I I L I S C ~BornerMyznplzis bzrckf017i

JacobMyzaphis rosrrr~rri7

(Kaltenbach)Myzocrrllis crrrpi/?i

(Koch)MyzoctrNis castnr?icolc~

BakerMyzocrrllis cor.yli

(Goeze)Myzus ~nti1.1.17i17i(Macchiati)Myru s ascalonicus

DoncasterMyrus certrsi PasseriniMyrirs cerrus (Walker)Myzirs he177eroccrllis

(Schrank)Myzirs lig~rstriMosley)

Myzirs lythri (Schrank)

Myrirs persicoe (Sulzer)Myrirs ~~ctrinnsavidsonNrrsor?ovicz ribisnigri

(Mosley)Neophyllt~l)lri.s rrr~rcrrrine

TakahashiNeoplq~ll~~phisoclocoipi

TakahashiNeotoxo~terafvr.r7zosar?o

(Takahashi)Neotoxoptero oliveri

(Essig)0vatil.s cratoegnr.irrs

(Walker)Pnracolophn 1norrisor7i

(Baker)Prrr7trphis jugltrizdis

(Goeze)

IL 1879 (Th oma s 1879 Palearctic (Europe)[as Sclrizo~zerrropirzicolrr])

ON, 1996 (CN C) Palearctic (Europe)NC, 1962 (Smith and Palearctic (Euro pe)

Knowlton 1975)ME, 1959 (USNM) Palearctic (Europe) RosnNY, 1906 (USN M) Palearctic (Euro pe) pest Ro.mBC, 1964 (Forb es Palearctic (Euro pe) Crr~pirzrrs

et al. 1973) berir1ir.rCA, 1914 (USN M) Palearctic (Europe) pest Crrsm~zen nd

Q i ~ e r c ~ ~ sCA, 1903 (Clarke Palearctic (Europe) pest Corylrls

1903) ~r~~ellnrzrrMD, 1996 (USN M) Palearctic (Europe) polyphagousNB, 1954

(MacGillivray1954b)

NY 1855 (Thomas1879)

NB, 1952 (MacGil-livray 1954a)

FL, LA, 1958(USNM)

NY, 1915 (USNM)

DC , 191 6 (U SNM )

CA, 1936 (Essig1938)

MA. 1841 (Harris1841)

CA 191 1 (Davidson1912)

IL, I879 (Thomas1879)

FL, 1963 (Woodruff1963)

CA, 1954 (Russell1967)

CA, 1927 (Essig1935)

CA, 1927 (Essig1935)

OR, 1893 (USNM)M D 19 16 (Baker

1919)OR, 1928 (USNM)

unknown Blackman pestand Eastop 2000

polyphagous

Palearctic (Europe) pest Pru~zlrs s.g.Cerrrslls)

Palearctic (Euro pe) minor pest Caryophyllaceae,incl. Dirrnthus

Palearctic (e Asia) pest He177er~octrllisPalearctic ( E L I I . ~ ~ ~ )ccasionally severe Lig~rsrrirn7 p.;

pest Mason 1940 E L I ~ ~ ~ ~ I ~ Z L I SSP.Palearctic (Euro pe) potential weed Lytl7ru177

biocontrol agent .rtrlict~r.iaeVoegtlin 1995b

Palearctic pest polyphagousPalearctic (e Asia) major pest polyphagousPalearc tic (e Asia) pest Cle17?ati.rPalearctic (Euro pe) major pest; L~ct lrca otivn

regulatoryconcern

Austra lasian pest Russell 1967 Arnitctrr.itr(Australia) heierophylln

Austra lasian pest Russell 1967, Porlocrrry~iu(Australia) Denma rk 1969

Oriental pest Alli~r177Oriental pest ANiin7zPalearctic (Euro pe) pestPalearctic (e Asia)

Merzrl?~~Crntaegils

bamboosPalearctic (e Asia) pest Jirgln~zs egin


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VOLUM E 108. NUM BER 3

Table 1. Continued.Earlicsl N.A. rccord (1 ) Gcogrilphic O~.igin/ Slatrls in Norlh Amcr.lca/ North AnicricanIn~roduccd pecics (Collccrion or Rcfcrcncc) Reference ( 2 ) Reference ( 2 ) Economlc Hosr(s)

Pen7phigrts brtrsctrirts NY, 1872 (Fitch Palearctic (Europ e) pest Populrts riigrn;(Linnaeus) 1872) Lnctrrcn

s a t i ~ ~ aPernplzigr~s /~y~'othee~~fC, 1972 (Alleyne and Palearctic (Euro pe) Poprrlrls nigrn

Passerini Morriso n 1974)Pe/7tn/or7irr tzigroner~~osrr DC , 1909 (US NM) Unknown (either pest Mrlsn

Coquerel Neotropical or (banana);Oriental) ornamental

Musaceae,Araceae,Zingiberaceae

Per.i/~Iiy/hrs alifor~tiierisis WA, 1903 (Essig and Palearctic (e Asia) Acer(Shinji) Aberna thy 1952) Essig and

Abernathy 1952PerG~Iz~Nr1s)iro~~ictr~s VT, 1847 (Fitch notes Palearctic (Europ e) Acer

(Kessler) [as Apliis nceris L.]) plntarioidesPeriplzyllrls BC, 1924 (Essig and Palearctic (Europe) Acertestudinaceus Abernathy 1952)(Fernie)

Phloeoil7yzus passeri r~i NB? 1950, ME 1970 Palearctic(Signoret) (22) (Smith 1974a [as P.

rlfar,borr~i mith])P/7orodo17 11~1112~11i NY, 1863 (Walsh Palearctic (Europe) pest P ~ L I ~ L I S ;

(Schrank) 1863) Hrrr?iulrlsPIi~~llcrp/2isi ~ g i IL, 1879 (Thoma s Palearctic (Europ e) pest Fag~ls

(Linnaeus) 1879)Pleotrichopltor~~a CA, 1910 (USN M) Palearctic (Asia?) minor pest chrysanthelnums

rhrysnt~t/~er?~i(Theobald)

Pleotrichophor~ts CO, 191 5 (USN M) Palearctic (Europe) A177br,osicr,glnr~du/osus Arternisicr(Kaltenbach)Procij) /~i l~~sylostei ME , 1914 (Patch 1917) Palearctic Lor~iceru; oots(De Geer) (23) of Piceo

Pse~~rlocn~~dellcr~rbiclrr NC, TN, 1973 (Smith Palearctic (Europ e)(Bomer) and Knowlton 1975)

Pserrdoregti~n or~icolrr FL, 1928 (FSAC) Oriental(Takahashi)

PterocaNis uh ~ i IL, 1910 (Davis 1910) Palearctic (Europ e)(De Geer)

Pterocon?nicr pilos r~ri~ UT, 1958 (Knowlto n PalearcticBuckton 1958)

Pter.ocot~~rnaO ~ J L I ~ ~ L ~ ~ I IT, 1904 (USN M) Palearctic (Europe)(Kaltenbach)

Pteroror~~r~zasrrlici.s MN , 1886 (Oestlund Palearctic(Linnaeus) 1886)

R I ~ o ~ J N I o ~ ~ ~ ~ ~ z L I sor~icerrre VA, 1912 (USN M) Palearctic (Europe) minor pest Lor~icera(Siebold)R / I O I J ~ ~ ~ S ~ J / I O ~ I ~ ~ L I . SC, 1975 (Forbe s and Palearctic (e Asia)hydrarzgeoe Chan 1976)(Matsumura)

R / I O ] J ~ ~ O S ~ J ~ O I I ~ ~ L I SA, 19 12 (USN M) Palearctic minor pest Bulbs; Solnrzr~mlatysip/~ori Davidson) tuberosro?~


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VOLUME 108, NUMB ER 3 595Table 1. Contin ued.

Earliest N.A. record ( I ) Geograph ic Origin1 Status in North America1 North AmericanIntroduce d Species (Collection or Reference) Reference (2) Reference (2) Economic Host(s)Therioaphis riehmi

(Borner)Therioaphis trifoli i(Monell) (29)

Tinocalliskahawaluokalani(Kirkaldy)

Tinocallis platani(Kaltenbach)

Tinocallis sultans(Nevsky)

TinocaNis talcachihoensisHiguchi

Tinocallis ulmiparvifoliaeMatsumura

TinocaNis zelkorvae(Takahashi)

Toxoptera aurantii(Boyer deFonscolombe)

Toxoptera citricida(Kirkaldy)

Trama rara MordvilkoTrichosiphonaphis

polygoni (van derGoot)

Tuberculatus annulatus(Hartig) (30)

Tuberculatus querccus(Kaltenbach)

Tuberolachnus salignus(Gmelin)

Uroleucon achilleae(Koch)

Uroleucon cirsii(Linnaeus)

Uroleucon sonchi(Linnaeus)

Uroleucon taraxaci(Kaltenbach)

Vesiculaphis caricis(Fullaway)

Wahlgreniella vaccinii(Theobald)

KS, WA, 1948(USNM)NY, 1879 (USNM)

LA, 1925 (USNM)

BC, 1927(Glendenning 1929)

ID, 1986 (Halbertand Pike 1990)

MD, 1996 (USNM)CA,FL, 1998 (Gill

1998b, Halbert1998)

CA, 1983 (Kon o1983)

CA, 1907 (USNM)

FL, 1995 (Halbertand Brown 1996)

CO, 1929 (USNM)FL, 1974 (Smith and

Denmark 1982)WA, 1951 (USNM)BC, 1999 (CNC)IL, 1872 (Le Baron

1872)CA, 1966 (Hille Ris

Lambers 1966)ON, 1956 (CNC)DC, 1892 (USNM)NY, 1902 (USNM)VA, 1916 (USNM)IL, 1929 (USNM)

Palearctic (Europe)Palearctic (Europe) pest

Oriental pest Mizell et al.2002

Palearctic (Europe)Palearctic (e Asia)Palearctic (e Asia)Palearctic (e Asia) nuisance pest

Palearctic (e Asia)Palearctic (e Asia) major pest

or OrientalPalearctic (e Asia) major pest Halbert

or Oriental and Brown 1996Palearctic (Europe)Palearctic (e Asia)

Palearctic (Europe)Palearctic (Europe)Palearctic (Europe) minor pestPalearctic (Europe)Palearctic (Europe)Palearctic (Europe)Palearctic (Europe)Palearctic (e Asia)Palearctic (Europe)

TrifoliumLeguminosae,

esp. Trifoliumand Medicago

Lagerslroemiaindica

UlmusUlmusUlmus

polyphagous

Citrusroots of

CompositaePolygonum

punctatum

SalixAchilleaCirsiumSonchusTaraxacum

1. Names of Canadi an provinces and U.S. states indicated using standard two-letter postal abbre-viations (Canada Post 2005, United States Postal Service 1998).2. Area of origin and pest status without explicit references are based o n Blackman an d Eastop (1994,2000) supplemented by opinion of the authors.3. Acyrthosiphon malvae: Also known as A. pelargonii. This name applies to a complex of entities withvarious host associations. We have not distinguished records for the various forms.

4. Aphidounguis mali: Single collection from botanical garden.5. Aphis craccivora: Previously infrequent on alfalfa, the recent upsurge of this species on alfalfa may

represent a separate introduction of an alfalfa-feeding population from Asia comparable to thesituation in Therioaphis trifolii (A . craccivora is known as the alfalfa aphid in China).


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VOLUME 108, NUM BER 3 597

Table 2. Species previously reported a s introduced in North America, but now considered t o beerroneous or dubious records.

Species Reason for Consideration as Erroneous or DubiousAnzphoropkora itlbi

(Kaltenbach)

Anurr!phis fr~rfrirae(Koch)Anuraphis stlbterranea

(Walker)Aphis frangulae Kaltenbach

Aplzis acetosae LinnaeusAphis evonyrni Fabricius

Aphis nzedicriginis KochAphis grossulnrine

KaltenbachAplzis ilicis KaltenbachAplzis hieracii SchrankAplzis nepetae KaltenbachAphis plantaginis GoezeAplzis ubnariae Schrank

Aplzis viburni Scopoli

Aplzis vitis ScopoliBrackycaudus prunicola(Kaltenbach)

-May no t occur in North America. References to this species on raspberry

in North America are mostly misidentifications of Amplzorophoraagatlzonica Hottes (Blackman and Eastop 2000). However, Hille RisLambers believed A. rubi was present in northwestern North America(Hille Ris Lambers letter to L. Russell, 13 Nov . 1972, USN M letterfile). Note that the European aphid on raspberry is actually A. idaei(Borner) while A. rubi is restricted to blackberry (Blackman et al.1977).

Listed in Bodenheimer an d Swirski (1957) as occurring in North Americabut no supporting specimens located.

Cited in Jones (1929) from Colorado. No specimens are present inUSNM.

North American records probably refer to Aphis nast~irtii.Aphisfiangulae is a member of a complex of taxa including Aphis gossypiiGlover (Smith and Parron 1978, Stroyan 1984, Blackman and Eastop2000). The overwintering host for A. fangulae sensu strict0 is Fiangulr~alnus (=Rhamn us frangula). A . nasturtii (as A. rlzamni Boyer deFonscolombe, see note below for Macchiatella rhamni) is the onlyAplzis reported on Rhamntls in Illinois by Hot tes and Frisson (193 1).

Listed in Borner and Heinze (1957) as occurring in North America, butno specimens have been located

There is considerable confusion in the application of the name A.evonyrni (or euonymi). It is probable that most, if not all, of the NorthAmerican records of A. evonynli (or A. euonymi) refer to othermembers of the Aphis fabae complex. Eastop and Hille Ris Lambers(1976) state tha t A. evonymi of authors prior to 1950 is a synonym ofApki sfab ae solanella Theobald. Eastop and Hille Ris Larnbers (1976)listed A. euonymi Fabricius 1775 as a valid species, and Remaudiereand Remauditre (1997) treat it as a subspecies of A. fabae.

North American records of this species are misidentifications of Aphiscraccivora.

Listed in Wilson (1913) as found in Oregon. N o specimens are present inUSNM.

Reported from California and British Columbia in Smith & Parron(1978). No supporting specimens have been located.

Listed in Rafinesque (1818). No North American specimens have beenlocated.

Listed in Smith and Parron (1978) as occurring in Idaho and Minnesota.There are no Nor th American records in the CNC, FSCA, or U SNM.

Listed in Smith and Parron (1978) as occurring in New York. No NorthAmerican specimens have been located

Listed in Smith and Parron (1978) from various states. Specimens fromColorado in USNM were actually A. spiraecola. Record by Thomas(1879) is A. pisum.

Recorded in Illinois as early as 1879 (Thomas 1879). However, recordsare likely the result of confusion with A. fabae an d A. viburniplzilaPatch.

Listed in Walsh (1863) from Illinois and Missouri. A single slide of poorquality found in USN M identified as A. vitis is likely Aphis illinoisensis.

Listed in Hunter (1901) as occurring in North America. There are noNorth American records for this species in the CNC, FSCA, orUS NM. Blackman and E astop (2000) state that this species is "not yetrecorded outside Europe."


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17/28

VOLUME 108, NUMBER 3Table 2. Continued

Species Reason for Consideration as Erroneous or DubiousMetopolophium frisicum Hille Listed in Smith and Parron (1978), citing Borner (1952). However, no

Ris Lambers specimens of this species have been located in No rt h Americancollections.Myzu s ajugae Schouteden Smith and Parron (1978) considered the Leonard (1968) reference to

"?My zus ajugae" suspect, believing "the determination was madebecause of the host association."

Myzus myosotidis (Borner) Smith and Parron (1978) cite Hille Ris Lambers (1966) as the basis forthe occurrence of M. myosofidis in California. However, Hille RisLambers (1966) mentions this species only in a key. No specimens arepresent in USNM. Halbert et al. (1995) considered North Americandistribution doubtful.

Nasonovia nigra (Hille Ris Recorded by Maw et al. (2000) in British Columbia and New Brunswick,Lambers) based on mididentified specimens of N. ribisnigrae in ACNB.

Pic turaphis brasiliensis Cited in Smith and Parron (1978) as occurring in Florida. Examination(Morei ra) of specimens in US NM showed the collection to be from "Florida,

P.R." (Puerto Rico), not Florida, U.S.A.Pachypappa tremulae Nor th American records probably refer to the native counterpart, P.(Linnaeus) rosettei (Maxson).Pemplzigus borealis Tullgren A single record from Idaho in 1952 (Gittins et al. 1976).Periphyllus aceris (Linnaeus) Although stated by Blackman and Eastop (1994) as introduced to North

America, and recorded by various other authors, all North Americancollections in USNM are interceptions. Essig and Abernathy (1952)believed that reports of P. ceris were referable to other species and P.aceris had not been recorded in the Western Hemisphere.

Prociphilus bumeliae (Schrank) According to Smith and Par ron (1978), probably misidentifications of P.americanus.

Saltusaphis scirpus Theobald A single record from Ohio, 1938 (Smith 1940). No Nor th Americanspecimens in USNM, CNC or FSCA.

Schoutedenia ralumensis A single specimen, cited in Palmer (1952) (as Setaphis viridis van derRiibsaamen Goo t) in Colorado. Not established.

Semiaphis dauci (Fabricius) Records from District of Columbia and Maryla nd (Leona rd and Bissell1970, Smith and Parr on 1978) may have been misidentification ofDysaphis crataegi.

Sitobion luteum (Buckton) Cited in Smith and Parr on (1978) as occurring in New Mexico and NewYork. All North American records in USN M are port interceptionsand do not represent established populations.

Thecabius affinis (Kaltenbach) Thecabius affinis is an Old World species, and Th. populiconduplifoliusNorth American. They are morphologically and biologicallyindistinguishable, but Th. afjinis has 2n = 38, and Th .populiconduplifolius has 2n = 28 chromosomes (Blackman and Eastop1994). It is unknown whether Th. affinis occurs in North America.

Thelaxes dryophila (Schrank) References to this species probab ly represent misidentification ofThelaxes calijbrnica (Davidson).

Therioaphis ononidis Records of T. ononidis from North America and Japan are(Kaltenbach) misidentifications of T. trifolii (Monell) (Quednau 2003).

Thripsaphis caricis (Mordvilko) North American records refer to Thripsaphis producta Gillette (Hille RisLambers 1974).

Titarzosiphon artemisiae (Koch) Questionably recorded for New York by Smith and Parron (1978), citingHunter (1901) who lists "Nectarphora arternisiae? Koch." Probablya Macrosiphoniella species.

Uroleucon composilae Listed in Hille Ris Lamb ers (1963) as present in "America." Although(Theobald) introduced into the Caribbea n, this species has not yet been found in

North America.


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600 PROCEEDIN GS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTONTable 2. Continued.

Species Reason for Consideration as Erroneou s or DubiousUroleucon nzuralis (Buckton) In Nebraska, according to in Williams (191 1). However, Rob inson

(1985) examined specimens from the US NM and determined that theywere Uroleucon sonchellui7~ Monell). He believed U. rnuralis does notoccur in North America.

Uroleucon solidaginis All known North American specimens identified as this species are in fact(Fabricius) various native species of Uroleucoii. Robinson (1985) stated that he

had "not seen authentic material of solidoginis from Nor th America."Uroleucon tanaceti (Linnaeus) Cited in Smith and Parron (1978) as occurring in several localities.

However, Robinson (1985) stated that he had "not seen authenticmaterial of this species from North America."

Table 3 . Aphid species considered to have a natural Holarctic or Beringian distribution.

Aphis equiseticola Ossiannilson

Aphis farinosa GmelinBoernerina alni Takahashi

Cavariella aquatica (Gillette and Bragg)Cavariella konoi Takahashi

Clzaitoplzorus horii Takahashi

Gllaresia poluniizi StroyanMaculolachnus sijpkensis Hille RisLambersMindarus obliquus (Cholodkovsky)

Muscaphis escherisclzi (Borner)

Myzodiuni modestum (Hottes)Thripsaphis cyperi (Walker)

Uroleucon simile (Hille Ris Lambers)

The single previous North American record from Vancouver,BC suggested an introduction of this species otherwiseknown f rom Scandinavia. However, a collection (in CN C) oft h s species in 2004 from north ern Qukbec indicates a possiblecircumboreal distribution.

Widespread in both Nearctic and Palearctic.Subspecies alaskensis Hille Ris Lambers and Hottes (originally

described as a subspecies of B. variabilis Richards),widespread in Alaska and arctic Canada, is the NorthAmerican segregate of this species otherwise known fromeast Asia.

Widespread in both Nearctic and Palearctic, but infrequentlycollected.

According to Stroyan (1969), differences in host range amongOld and New World populations indicate the possibility thatthis name covers a complex of taxa.

Specimens from Northwest Territories match the nominateAsian subspecies, rather than the European subspecies(beuthani Borner), and probably represent natural migrationacross the Bering Strait.

Known from mountains of western North America andHimalayas.

Distributed across North America and in Mongolia (Blackmanand Eastop 2000).

Present across boreal forest zone of North America, northernEurope, and northern Russia, but apparently not recorded ineastern Siberia.

According to Stekol'shtshikov and Shaposhnikov (1993),Toxopterella rirepanosiphoides MacGillivrary and Bradley(known from across Canada, and in northern Europe assubspecies irae Shaposhnikov) is the primary host form of M.escherisclzi.

Widespread in both Nearctic and Palearctic.Although first Nort h American records of this species in Alaska(USNM) and O ntario (CN C) date from 1961, it is distributedacross boreal and subarctic Canada and is likely naturallycircumboreal.

Widespread in Palearctic. North American records (Alaskaonly) likely represent a natural expansion across BeringStrait.


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VOLUME 108, NUM BER 3 60 1

Before 1840- 1870- 1900- 1930- 1960- 1990-1840 1869 1899 1929 1959 1989 present

30-year inter valsFig. 1. Adventive aphid s. Number of adventive aphids in Nort h America north of Mexico recorded at

30-year intervals. Two species not included due to confusion with related species.

(1997), except for changes noted byEastop and Blackman (2005). The taxo-nomic status and biogeographic originsof some aphid species are difficult todetermine conclusively due to ongoingtaxonomic problems. We have summa-rized relevant information in footnotesto Table 1.

Parameters for determining a pest areas defined in D. R. Miller et al. (2002),Miller and Miller (2003), and Miller et al.(2005). That is, if statements are madethat a species is a pest, causes harm,requires control measures, is economi-cally important, or reflects some compa-rable statement or inference, then weconsider that species a pest. Pest statusentries in Table 1 without reference toother authors are either from Blackmanand Eastop (1994, 2000) or our personalexperience. The principal hosts listed arethose of the greatest economic impact inNorth America (including crops, orna-mental plants, trees and weeds). Species

causing damage to several plant familiesare listed as polyphagous.

A summary of species previously listedfor North America, but for which weconsider their occurrence doubtful orerroneous, and an explanation for theirexclusion is provided in Table 2. Table 3contains a compilation of those species,some of which were previously suspectedas adventive to North America, whichwe consider to be most probablyeither Holarctic or Beringian in distribu-tion.

RESULTSBased on our findings there are 1,416

aphid species in North America north ofMexico, which comprises about 30% ofthe world fauna of approximately 4,700species (Remaudiere and Remaudiere1997). Table 1 provides information on262 aphid species (1890 of the NorthAmerican and approximately 6% of the


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(Miller et al. 2005), most adventiveaphids are indigenous to the PalearcticRegion (especially Europe). This is to beexpected, given the immigration historyof the continent and the associatedtransport of crops and their weeds.However, European introductions havebecome relatively few (Fig. 2, and seenas a leveling off of the Europeancomponent of the cumulative curve inFig. 3). Asian species now account fora major proportion of new introduc-tions, although the total number issimilar to that in the early part of the20th century (Fig. 2). Because the Asianfauna was poorly known at the time,several of these early oriental introduc-tions were first described from popula-tions in the U.S.A. (Aphis spiraecolaPatch, Atarsaphis agrifoliae Ferris, Eu-thoracaphis umbellariae Essig, Neotoxop-tera olivieri (Essig), Paracolopha morri-soni Baker, Periphyllus californicus(Shinji), Takecallis arundicolens (Clarke),T. arundinariae (Essig)) as was theAfrican aphid Aloephagus myersi Essig.

The proportion of pests among theadventive species is much greater thanthe proportion among the native species.This to be expected, given that most ofthese introductions were enabled byhuman activity, and hence associatedwith economically important plants. Amore detailed analysis of the economicimpact of adventive and native species isin preparation.

ACKNOWLEDGMENTSWe thank D. Creel and J. Eibl

(USDA, ARS, Systematic EntomologyLaboratory, Beltsville, MD) for theirassistance with reference verification forthis paper. We also thank P. Mason(AAFC, Ottawa), E. Grissell (USDA,ARS, Systematic Entomology Laborato-ry, Washington, DC) and D. Nickle(USDA, ARS, Systematic EntomologyLaboratory, Beltsville, MD) for theirmanuscript reviews.

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Anonymous. 2005. Soybean aphid in Minnesota.http://www.soybeans.umn.edu~crop/insects/aphid/aphid.htm [accessed 4 May 20051.

Archibald, K. D. 1957. Report of the forestAphidae (Homo ptera ) of Nova Scotia togetherwith species and host plant lists. Annals of theEntomological Society of America 50: 3842.

Baker, A. C. 1919. An undescribed species ofDryopen injurious to Plzyllostaclzys. (Aphidi-dae-Hom.). Proceedings of the EntomologicalSociety of Washington 21: 106106.Baker, A. C. and W. M. Davidson. 1916. Woolypear aphis. Journal of Agriculture Research 6:351-360.

Bartholomew, P. S. 1932. Six new species of aphids,with records of other species new to California.Annals of the Entomological Society of Amer-ica 25(4): 713-729.

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North American Allien Aphid Species

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