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Home , Aphid, Aphididae, Aphidinae, Aphidini, Aphis nasturtii, Atheroides

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

ADVENTIVE (: ) OF AMERICA NORTH OF MEXICO

(RGF, EM) and Agri-Food Canada, Canadian National Collection of , Ottawa, ON, K1A 0C6 Canada (e-mail: [email protected]; [email protected]); (SEH) Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Gainesville, FL 32614, U.S.A. (e-mail: [email protected]); (GLM, LMR) Systematic Laboratory, Plant Science 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 of aphids that are considered as adventive to North America north of Mexico. Included for each species, where applicable, is reference to: the location and date of introduction of the first North American record; status in North America; principal economic hosts; and biogeographical origin. Information is also provided for species whose presence in North America is considered erroneous or questionable and for those species that are considered Holarctic or Beringian. Key Words: Aphidoidea, plant lice, plant pathogens, , biological control, quarantine, agriculture, , horticulture, ornamentals, fruit trees

In an era of a global economy and reintroduced into the United States commerce, the increase of adventive (Huber et al. 2002). species represents both domestic and Many species have been added international concerns. If non-indige- to the North American fauna since nous species become pests, the economic colonization by Europeans, and some consequences include loss of production, of these have become serious pests. In diminished quality, production cost in- addition to direct feeding damage, creases, flexibility decreases in produc- aphids constitute one of the largest tionlmanagement decisions, increased groups of known plant- vectors risk of human disease, and damaging (Chan et al. 1991). Aphids are primarily environmental and aesthetic effects (Hu- controlled through pesticide applica- ber et al. 2002). Various insects of tions, adding to production costs, agricultural concern have been identified and having environmental implications as potentially damaging if introduced or associated with increased pesticide use. The effects of some non-indigenous aphids in North America have been 1 It is noteworrhy that Louise Russell, a pioneer woman enmrnologist and specialist in the S~ernorrh~ncha,continues staggering. For example, by the early to publish at the age of 101. 199OYs,the Russian aphid, Diur- 584 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON aphis noxia (Kurdjumov), was responsi- (Harris 2003), no aphid species has been ble for $500-900 million cumulative purposely introduced. losses in the western U.S. (Bernal et al. Geographic origins are given in gen- , 1993, Morrison and Peairs 1998). The eral terms only, relying heavily on Black- appearance of the brown aphid, man and Eastop (1994, 2000), supple- citricida (Kirkaldy), in Flor- mented by other references, as noted in ida in 1995 had a major impact on the Table 1, and by opinion of the authors. U.S. citrus industry (Halbert and Brown The origins of a few cosmopolitan 1996). More recently, losses related to species are unknown. It is not always the , Aphis glycines Mat- readily apparent whether a species is sumura, have been estimated at about adventive or native; some species we $120 million in 2003 for Minnesota alone have listed as adventive may in fact be (Anonymous 2005). naturally Holarctic in distribution and The origins of this work actually vice versa. began in part in the 1970's when one of For determining the earliest record of us (LMR) contributed data on invasive the North American adventive aphids, species in the United States (e.g., Sailer we relied on published records and on 1978). Nearly 30 later, adventive specimens examined in The Canadian insects are still a major concern for National Collection of Insects, Nema- agriculture in North America. The ob- todes and Arachnids (CNC) and the jectives of this paper are to: 1) list the Edith Patch Collection (CNC-Patch) adventive aphid species (excluding adel- housed at CNC, Ottawa, ON, Canada; gids and phylloxerans) of North America the MacGillivray Collection (ACNB), and record data on first detection date Agriculture and Agri-Food Canada, (along with validation source), principal Fredericton, NB, Canada; The Florida hosts, and zoogeographic area of origin; State Collection of (FSCA), 2) analyze these data and determine if Gainesville, FL, U.S.A.; and The Na- patterns exist; and 3) compare these data tional Entomological Collection of the to other sternorrhynchan groups. National Museum of (USNM), Beltsville, MD, U.S.A. In addition, Asa Fitch's aphid species notes We use the term adventive species as that are currently housed with the defined by D.R. Miller et al. (2002), i.e., USNM aphid collection at Beltsville, for those species that are non-native to MD, were also examined. This collection North America regardless of economic of written records and observations from harm. Other terms such as invasive, the 1840's-1870's provide some of our alien, exotic, and immigrant have some- earliest glimpses into the North Ameri- times been used interchangeably (see can aphid fauna. While there are some Wheeler and Hoebeke (2001) and Miller questions concerning Fitch's "species et al. (2005) for discussion). Using Smith concept" versus current thinking for and Parron's (1978) list of North Amer- some of his included species, there is ican aphids as the base, we compiled little confusion given the morphology a new list for North American north of and associated host plants for others. His Mexico. We exclude species that have reference to the occurrence in 179 1 of the been intercepted at ports-of-entry or aphid, brassicae (L.) discovered in nurseries or is possibly the earliest record of an and subsequently eradicated. With the adventive aphid for North America. exception of Aphis chloris Koch, a bio- Nomenclature followed herein reflects control agent for perforatum that of Remaudiere and Remaudiere VOLUME 108. NUMBER 3 585

Table 1. Introduced aphid species in Canada and continental United States of America, giving earliest record with source of record, geographic origin, pest status, and economic host species. Aphid names follow Remaudikre and Remaudikre (1997). Earliest records are based on literature references and on specimens in the following collections: USNM - National Museum of Natural History; CNC - Canadian National Collection of Insects, Arachnids and Nematodes; FSCA - Florida State Collection of Arthropods; ACNB - MacGillivray collection, Agriculture and Agri-Food Canada, Fredericton, NB; CNC-Patch - Edith Patch collection, housed at CNC; EMEC - Essig Museum of Entomology Collection, University of California, Berkeley.

Earliest N.A. record (I) Geographic Origin/ Slatus io North America1 North American Introduced Species (Collection or Rererence) Rercrcnce (2) Reference (2) Economic Hosl(s) caraganae SK, 1940 (CNC) Palearctic pest Caraganrr (Cholodkovsky) nrborescens Acyrthosipl~or~kondoi CA, 1974 (Kono Palearctic (e Asia) pest Halbert et al. Medicago Shinji 1977) 2000 srrtiva Acyrthosiphon lactucae MD, 1943 (USNM) Palearctic (Europe) Lactuca sativa ( Passerini) Acyrthosipl~onmalvae NE, 189 1 (USNM) Palearctic (Europe) Geranirrm, (Mosle~)(3) Pelargoi~ium Acyrthosipl~onpisum KS, 1877 (Harper Palearctic (Europe) pest Leguminosae, (Harris) et al. 1978) incl. Pisum, Medicago Aloephagus mnyersi CA, 1939 (Essig 1950) Afrotropical minor pest Essig Aloe Essig 1950, regulatory () issue Amnpl~orophorrr MN, 1887 (Oestland Palearctic (Europe) anzpullata Buckton 1887) Anoecin corni IA, 1897 (USNM) Palearctic (Europe) roots of (Fabricius) Gramineae, incl. Aphidou~zg~ris~nr~li NC, 1999 (CNC) Palearctic (e Asia) Takahashi (4) Apl~isainara~zthi FL, 1990 (Halbert Neotropical potential pest edible Holman et al. 2000) (Cuba) Halbert et al. 2000 Ainara~~thus Aplzis 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 perforatum agent Aphis craccae Linnaeus BC, NB, ME, 1965 Palearctic Vicia cracca (CNC, ACNB) Aphis cr~accivorrr NE, 1890 (USNM) Palearctic polyphagous, Koch (5) with preference for legumes Aphis cytisorurn Hartig DC, 1906 (USNM) Palearctic (Europe) Cyt isus scoparius 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. 2004 WV, 2000 (Voegtlin Asia) 2000) Glover NY, 1852 (Russell Palearctic major pest polyphagous 1968) 586 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Continued.

Earliest N.A. record (I) Geographic Origin1 Status in Nortll Amerieal North American Inlrodueed Species (Collection or Rererence) Rererence (2) Rcrercnce (2) Economic Host@) Aphis hederae CA, 19 10 (Essig Palearctic (Europe) minor pest Hedera helix Kaltenbach 1910) Aphis idaei van der BC, 1951 (Forbes Palearctic (Europe) Goot et al. 1973) NE, 1894 (USNM) Palearctic pest Kaltenbach Aphis nepetae ID, 1957 (Gittins Palearctic Kaltenbach et al. 1976) Boyer de IL, 1879 (Thomas Palearctic (e Asia) pest ornamentals Fonscolombe 1879) (flowers) Aphis polygonata IL, 1977 (FSCA) Palearctic (Central (Nevsky) (8) Asia) De Geer PA, 1844 (Haldeman Palearctic (Europe) pest Malus pumila, 1844) Pyrus communis, Cydonia oblonga Aphis praeterita Walker ME, 1918 (Patch Palearctic (Europe) 1918) Aphis rumicis Linnaeus Unknown (7) Palearctic (Europe) (7) Aphis salicariae Koch ON, 1952 (CNC) Palearctic (Europe) Aphis sambuci Linnaeus NY, 1847 (Fitch Palearctic (Europe) notes [as A. sambucifoliae Fitch]) Aphis sedi Kaltenbach ME, 1911 (Patch Palearctic (Europe) 191 1) Patch CO, 1907 Palearctic or major pest polyphagous (Gillette 19 10) Oriental Aphis iaraxicola QC, 1983 (Cloutier Palearctic (Borner) (9) et al. 1986) Aphis ihaliciri Koch BC, 1999 (CNC) Palearctic (Europe) Aphis umbrella (Borner) NE, SD, 1891 Palearctic (Europe) (USNM [as A. maliae Koch]) Aphis urricnta Gmelin PA, 19 13 (Olsen Palearctic (Europe) 1921) Aphis vaccini (Borner) BC, 2000(CNC) Palearctic (Europe) Aploneura lentisci IL, 1879 (Thomas Palearctic Gramineae, (Passerini) 1879 [as (Mediterranean) including Rhizobius pone Trir icurn Thomas]) Asiphonella daciylonii AZ, 1947 (USNM) Palearctic (e Asia) potential pest Cynodon Theobald daciylon Aspidaphis adjuvans CO, 1907 (Gillette Palearctic (Europe) (Walker) 1917b) Ararsaphis ngrijoliae CA, 1921 (Ferris Palearctic (e Asia) (Ferris) 1921) serrulnius QC, 1966 (CNC) Palearctic minor pest Gramineae, Haliday incl. cereals Aulacor ihum CO, 1907 (Gillette Palearctic pest polyphagous circunzflexum 1908) (e Asia) or (Buckton) Oriental Aulacorihun7 solani NY, 1879 (Riley and Palearctic major pest polyphagous (Kaltenbach) Monell 1879) Betulaphis brevipilosa BC, 1959 (Forbes Palearctic (Europe) Borner et al. 1974) VOLUME 108, NUMBER 3 587

Table I. Continued.

Earliest N.A. record (I) Geographic Origin/ Status in North America1 North Amcrican Introduced Soecies ICollection or Reierence) Reference (2) Reference (2) Economic Hods) Betulaphis I?elvetica Hille BC, 1926 Palearctic (Europe) Ris Lambers (Glendenning 1926) Betulaphis BC, 1946(ACNB) Palearctic (Europe) quadrituberculata (Kaltenbach) Bracl~ycaudushelichrysi OR, 1893 (USNM) Palearctic major pest Prunus, (Kaltenbach) (Europe) Compositae Bracl~ycauduscardui NY, 1887 (USNM) Palearctic pest Prunus, (Linnaeus) (Europe) Compositae Brachycatrdus persicae MD, DE, NJ, VA, Palearctic pest Prunus (Passerini) MI, 1887 (Smith (Europe) 1890) CT 1913 (Patch Palearctic (Europe) ruinexicolens (Patch) 19 17) (10) Brachycaudus schlvartzi CA, 1975 (Eastop Palearctic pest Prunus persica (Borner) 1975 ) (Europe) Brachycaudus spiraeae ON, 1956 (CNC) Palearctic (Borner) Bracliycaudus WA, 1993 (FSCA) Palearctic tragopogonis (Kaltenbach) Brachycorynella asparagi NY, NJ, 1969 Palearctic locally serious pest Asparagus (Mordvilko) (Leonard 1970, (Europe) Stoetzel 1990 officinalis Race 1970) NY, 179 1 (Fitch Palearctic pest Brassica (Linnaeus) notes) Calaphis betulicola NY, 1854 (Fitch Palearctic (Europe) (Kaltenbach) notes) Callipterir7ella calliptera OR, 1909 (USNM) Palearctic minor pest Hajek Betula (Hartig) (Europe) and Dahlsten 1986 Callipterinella BC, 1975 (Forbes and Palearctic minor pest Hajek Betula minulissima (Stroyan) Chan 1980) (Europe) and Dahlsten 1987 carduinus GA, 1948 (USNM) Palearctic potential weed Cirsum (Walker) (Europe) biocontrol agent Gassmann and Kok 2002 SD, 1892 (USNM) Palearctic Potential virus (Del Guercio) (Europe) Capitophorus hippophaes DC, 1895 (USNM) Palearctic (Europe) (Walker) Capitophorus pakansus IL, 1929 (Hottes and Palearctic (Europe) Hottes and Frison Frison 1931) Capitophorus horni ME, 1951 (Corpuz- Palearctic (Europe) Bomer Raros and Cook 1974) Cavariella aegopodii MN, 1903 (USNM) Palearctic pest (Scopoli) (Europe) Cavariella archangelicae WA, 1953 (USNM) Palearctic (Europe, (Scopoli) Iceland) Cavariella pastinacae NY, 1855 (Fitch Palearctic (Europe) (Linnaeus) notes) brasiliensis OH, 1917 (USNM) Oriental pest palms (Hempel) Cerataphis lataniae NE, 1912 (USNM) Oriental pest palms (Boisduval) DC, 1923 (USNM) Oriental pest greenhouse (Westwood) orchids 588 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Continued.

Earliest N.A. record (1) Geographic Origin1 Status in North America1 North American lnlroduced Species (Collection or Reference) Reference (2) Refcrcnce (2) Econonlic Hos~(s) Ceruropliis erioplzori UT, 1935 (Knowlton Palearctic (Europe) (Walker) and Smith 1936) C/znelosi[~horzpoterztillrre NY, 185 1 (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, Quednau 1966) Clzaitophorus popzrlinlbne ON, 1955 (CNC) Palearctic (Europe) (Boyer de Fonscolom be) Clzrornnphis jugln~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 Abies Cinrrro costnto ON, 1905 (USNM) Palearctic Picerr (Zetterstedt) cupressi CA, 1916 Palearctic Cupressus, (Buckton) (Bartholomew Juniperus 1932) Cinarn jkesrri Blanchard FL, 1942 (Tissot unknown Cupress~~s, 1944 [as C. Jznziperus wacnsnssoe]) Cirzarn jur7ipn.i NY, 1902 (USNM) Palearctic (Europe) Juniperus (De Geer) Ci~znrnloricis (Hartig) MA, 1921 (USNM) Palearctic (Europe) Lorix Ci~znrnpilicor~nis ME, 1906 (Patch Palearctic Picerr (Hartig) 1912) Cinnrrr pirzea DC, 1919 (Wilson Palearctic Pirzzrs (Mordvilko) 1919) Cirzarn preri~zosrr (Hartig) CO, 19 10 (Gillette Palearctic Picerr 19 1 7a [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 (Central (Lichtenstein) Asia) Color~doorrc/zillerre ON, 1962 (CNC) Palearctic (Central Hille Ris Lambers Asia) Colorrrdorr rrrtenzisirre BC, 1965 (CNC) Palearctic (Central (Del Guercio) Asia) Color~rrdorrrzfor?zrrc~rlrrtn NE, 1890 (USNM) Palearctic (Central minor pest (Wilson) Asia) Color~rrdorrtrnzrrcetirzn RI, 1974 (Smith and Palearctic (Central (Walker) Parron 1978) Asia) Corylobizn?z avellrrnrre BC, 1983 (Forbes and Palearctic (Europe) pest Corylus (Schrank) Chan 1984) rrvellrrrzn Cryptcryhis pone (Hardy) MB, 1965 (Robinson Palearctic (Europe) and Bradley 1965) Cryptor?7yszr.r gnleopsidis ME, 1914 (Patch Palearctic (Europe) (Kaltenbach) 1914) VOLUME 108. NUMBER 3 589

Table I. Continued.

Earlicst N.A. record (I) Geographic Origin1 Slalrls in North Alnerical North American Introduced Species (Colleclion or Reference) Rcfcrencc (2) Reference (2) Economic Host(s) Crypton?j~zusribis NY, 1854 (Fitch Palearctic (Europe) pest Ribes (Linnaeus) notes) Ctei7ocallis setosus OR, 1951 (Essig Palearctic (Europe) (Kaltenbach) 1952) Cter~ocollissellcoi~;ne CA, 1983 (Kono Palearctic (E Asia) Pest Kono, 1983 (Shinji) 1983) DecorosQ>horz NC, 1963 (Smith and Palearctic (Europe) coq~17othrisBorner Knowlton 1975) Dirrrrrphis fieque~zs ON, 1951 (CNC) Palearctic minor pest Halbert Walker (Europe) et al. 1992 Dizrrnphis rzosirr TX, 1986 (Stoetzel Palearctic major pest Gramineae, (Kurdjumov) 1987) (Central Asia) Quisenberry and incl. cereals Peairs 1998 Drepfllzos~~/~~n~i OR, 1914 (USNM) Palearctic Picm oregone17sis Granovsky Dre~~orzosiplz~mi CA, 1908 (Wilson Palearctic (Europe) Acer plrrtnrzoides (Schrank) 1909) Dysophis apiifolirr CA. 1934 (USNM) Palearctic pest Umbelliferae (Theobald) Dysuphis cratuegi DC, 1915 (USNM) Palearctic pest Umbelliferae (Kaltenbach) (1 1) Dysrrpkis devecta QC, 1966 (Quednau Palearctic (Europe) (Walker) 1966) Dysrrpliis joer7iczrlzrs OR, 1949 (USNM) Palearctic occasional pest Umbelliferae (Theobald) (12) (Europe) (including Daucus CCI~O~O) Dysrrpliis plrmtrrgir~ea NE, 1890 (Williams Palearctic (Europe) pest Mcrlus purnila (Passerini) 191 1) Dysopl7is pyri (Boyer de CO, 1978 (Smith and Palearctic Fonscolombe) Parron 1978) Dysophis radicolrr DC, 1894 (USNM) Palearctic (Europe) (Mordvilko) Dysnplzis tulipae (Boyer NH, 1903 (USNM) Palearctic pest, regulatory bulbs de Fonscolombe) (Europe) concern Elrrtobi~minbietilivn7 CA, DC, 1915 Palearctic pest Abies (Walker) (USNM) (Europe) Eriosoriirr gross~rlrri.irre ON, 19 13 (CNC) Palearctic (Europe) Ribes. Uli7i~ts (Schiile) (1 3) E1.ioso17irrpyricoln Baker OH, 1897 (Baker and Palearctic (Europe) Pyrus & Davidson Davidson 19 16) cor17r17ur7is Erios0117~U/II?~ BC, 2003 (CNC) Palearctic UIII~ZIS;Ribes (Linnaeus) ( 13) EtrcrrllQ~ter-z~stilirie DC, 1890 (USNM) Palearctic (Europe) pest Schouteden Eucarazzirr elegnns CA, 1984 (Stoetzel Palearctic (Europe) Labiatae (Ferrari) 1985) Eucerrrprphis betzrlue CO, 1909 (USNM) Palearctic (Europe) (Koch) (14) Euceruphis pu17ctiperz17is NY, 1847 (Fitch Palearctic (Europe) (Zetterstedt) (14) notes) Eulrrchnus ngilis NY, 1909 (Gillette Palearctic (Europe) (Kaltenbach) 1909) Errlr,clzr7us bre1~Q1i1osu.v WA, 1954 (Hottes Palearctic (Europe) Borner and Essig 1955) Eulochi7us rileyi IN, 1886 (USNM) Palearctic (Europe) minor pest (Williams) 590 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Continued. Earliest N.A. record (I) Geographic Origin1 Status in North America1 North American Introduced Species (Collection or Rererence) Reference (2) Rererenee (2) Economic Host(s) Euthoracaphis CA, 1929 (Essig Palearctic (e Asia) umbellariae Essig 1932) Fordaformicarir~von CO, 1909 (USNM) Palearctic (Europe) roots of Heyden Gramineae marginata Koch DC, 1885 (USNM) Palearctic (Europe) roots of Gramineae 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. cereals Geopemphigusfloccosus FL, 1945 (USNM) Neotropical minor pest roots of (Moreira) Zpomoeu bututas Glyphina betulue PA, 1945 (USNM) Palearctic (Europe) (Linnaeus) ficicola FL, 2002 (Halbert Oriental potential pest Ficus Takahashi 2004) Greenidea psidii van der CA, 1998 ( Oriental minor pest, Myrtus Goot 1998a) regulatory pest atriplicis CA, 1887 (USNM) Palearctic (Europe) Chenopodium (Linnaeus) including Ch. quinoa coriandri FL, 1997 (Halbert Palearctic (Central major pest Umbelliferae, Pas) et al. 2000) Asia) including Coriundrum, Foeniculum Hyadaphis foeniculi NY, 1879 (USNM) Palearctic (Central minor pest Lonicera ; (Passerini) Asia) Umbelliferae Hyadaphis tartaricae IL, 1979 (Voegtlin Palearctic (Central major pest Voegtlin Lonicera (Ajzenberg) 1981) Asia) 1981 tatarica Hyalopteroides humilis DC, MA, 1909 Palearctic (Walker) (Hayhurst 1909) amygdali IL, NJ (date ?) (16) Palearctic (e Asia) pest Prunus (s.g. (Blanchard) (1 6) (Smith and Parron Amydalus) 1978) IL, 1879 (Thomas Palearctic (Europe) pest Prunus (s.g. (Geoffroy) (16) 1879) Prunus) FL, 1997 (Halbert Palearctic (e Asia) carduellinus et al. 2000) (Theobald) IL, 1878 (Thomas Palearctic (Europe) minor pest Ribes (Linnaeus) 1879) Hyperomyzus pallidus ME, 19 14 (Hille Ris Palearctic (Europe) Ribes Hille Ris Lambers Lambers 1949) Hyperomyzus picridis PA, 1942 (USNM) Palearctic (Europe) potential pest Ribes (Borner) Hyperomyzus rhinanthi ME, 195 1 (CNC) Palearctic (Europe) Ribes (Schouteden) (1 7) Zdiopterus nephrelepidis IL, 1907 (Davis 1909) Neotropical pest Stoetzel 1990 greenhouse Davis Stoetzel 1990 ferns Jacksonia papillata ID, 1959 (USNM) Palearctic (Europe) Theobald berberidis NY, 1846 (Fitch Palearctic (Europe) (Kaltenbach) notes) Lipamyzodes matthiolae WA, 1976 (Fox 1977) Palearctic (Europe) (Doncaster) VOLUME 108, NUMBER 3 59 1

Table I. Continued

Earliest N.A. rccord (I) Geographic Origin1 Status in North America1 North Amcrican Introduced Species (Collection or Reference) Reference (2) Rcference (2) Economic Host(s) pseudobrassicae DC, 1899 (USNM) Palearctic (Europe) major pest Paddock Cruciferae (Davis) (1 8) 1915 Longicnudus trirlzodus MA, 1908 (USNM) Palearctic (Europe) Rosa (Walker) IL, 1929 (USNM) Palearctic abrotani (Walker) Macrosiphoniella IL, 1879 (Thomas Palearctic absinthii (Linnaeus) 1879) Macrosiphoniella NB, 1950 (ACNB) Palearctic arlemisiae (Boyer de Fonscolombe) Macrosiphoniella MD, 1997 (USNM) Palearctic grandicauda Takahashi and Moritsu Macrosiphoniella PA, 1948 (USNM) Palearctic Leucanthemum leucantlzemi vulgare (Ferrari) Maoosiphoniella CT, 1937 (EMEC) Palearctic millefolii (De Geer) Macrosiphoniella NE, 1891 (USNM) Palearctic (e Asia) pest Asteraceae sanborni (Gillette) (esp. chrysanthe- mums) Macrosiphoniella PA, 1948 (USNM) Palearctic cultivated subterranea (Koch) chrysanthe- mums Macrosiphoniella CT, 1909 Palearctic minor pest cultivated tanace taria (CNC-Patch) chrysanthe- (Kaltenbach) mums (occa- sionally) Mao.osiphrrm lilii NY, 1878 (Comstock Palearctic? pest Cultivated lilies (Monell) (19) 1880) rosae MA, 1841 (Harris Palearctic (Europe) pest Rosaceae (esp. (Linnaeus) 1841) Rosa) Mucrosiphurn stellariae BC, 1980 (Forbes Palearctic pest Caryophyllaceae Theobald and Chan 1983) (esp. Dianthus) bambusae GA, 1914 (USNM) Palearctic (e Asia) bamboos Fullaway FL, 1977 (FSCA) Oriental pest White et al. Gramineae (Zehntner) 200 1 (esp. sugar- cane) Melanaphis sorini FL via CA, 1996 Oriental Halbert pest Halbert and Miscanthus Halbert and (Halbert and and Remaudikre Remaudiire 2000 Remaudikre Remaudikre 2000) 2000 UT, 1910 (USNM) Palearctic (Europe) major pest Gramineae, dirhodum (Walker) incl. cereals; Rosa Metopolophium festucae CA, 1970 (USNM, Palearctic (Europe) pest Gramineae (Theobald) Eastop 1971) carnosum NE, 1889 (Williams Palearctic (Europe) (Buckton) (20) 1911) Microlophium sibiricurn PA, 1963 (USNM, Palearctic (Europe) Urtica (Mordvilko) (20) Pepper 1965) 592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Continued.

Earliest N.A. rccard (1) Geagrapl~icOrigin/ Status in North Amcrical North Amcric;tn l~~lroducedSpccies (Collection or Rcfcrcncc) Reference (2) Refercncc (2) Econonlic Hosl(s) Mir~cIr~~sr~bietiizzr171 IL 1879 (Thomas 1879 Palearctic (Europe) Koch (21) [as Sclrizo~zerrro pirzicolrr]) Mor~~~hisariterzrza ta ON, 1996 (CNC) Palearctic (Europe) (Kaltenbach) Mir.scrrphis IIILISC~Borner NC, 1962 (Smith and Palearctic (Europe) Knowlton 1975) Myznplzis bzrckf017i ME, 1959 (USNM) Palearctic (Europe) Rosn Jacob rosrrr~rri7 NY, 1906 (USNM) Palearctic (Europe) pest Ro.m (Kaltenbach) Myzocrrllis crrrpi/?i BC, 1964 (Forbes Palearctic (Europe) Crr~pirzrrs (Koch) et al. 1973) berir1ir.r MyzoctrNis castnr?icolc~ CA, 1914 (USNM) Palearctic (Europe) pest Crrsm~zenand Baker Qi~erc~~s Myzocrrllis cor.yli CA, 1903 (Clarke Palearctic (Europe) pest Corylrls (Goeze) 1903) ~r~~ellnrzrr ~nti1.1.17i17i MD, 1996 (USN M) Palearctic (Europe) polyphagous (Macchiati) Myrus ascalonicus NB, 1954 unknown Blackman pest polyphagous Doncaster (MacGillivray and Eastop 2000 1954b) Myrus certrsi Passerini NY 1855 (Thomas Palearctic (Europe) pest Pru~zlrs(s.g. 1879) Cerrrslls) Myrirs cerrus (Walker) NB, 1952 (MacGil- Palearctic (Europe) minor pest Caryophyllaceae, livray 1954a) incl. Dirrnthus Myzirs he177eroccrllis FL, LA, 1958 Palearctic (e Asia) pest He177er~octrllis (Schrank) (USNM) Myzirs lig~rstri(Mosley) NY, 1915 (USNM) Palearctic (ELII.~~~)occasionally severe Lig~rsrrirn7sp.; pest Mason 1940 ELI~~~~I~ZLIS SP. Myzirs lythri (Schrank) DC, 191 6 (USNM) Palearctic (Europe) potential weed Lytl7ru177 biocontrol agent .rtrlict~r.iae Voegtlin 1995b CA, 1936 (Essig Palearctic pest polyphagous 1938) Myrirs persicoe (Sulzer) MA. 1841 (Harris Palearctic (e Asia) major pest polyphagous 1841) Myrirs ~~ctrinnsDavidson CA 191 1 (Davidson Palearctic (e Asia) pest Cle17?ati.r 1912) Nrrsor?ovicz ribisnigri IL, I879 (Thomas Palearctic (Europe) major pest; L~ctlrcasotivn (Mosley) 1879) regulatory concern Neophyllt~l)lri.snrrr~rcrrrine FL, 1963 (Woodruff Australasian pest Russell 1967 Arnitctrr.itr Takahashi 1963) (Australia) heierophylln Neoplq~ll~~phispoclocoipi CA, 1954 (Russell Australasian pest Russell 1967, Porlocrrry~iu Takahashi 1967) (Australia) Denmark 1969 Neotoxo~tera,fvr.r7zosar?o CA, 1927 (Essig Oriental pest Alli~r177 (Takahashi) 1935) Neotoxoptero oliveri CA, 1927 (Essig Oriental pest ANiin7z (Essig) 1935) 0vatil.s cratoegnr.irrs OR, 1893 (USNM) Palearctic (Europe) pest Merzrl?~~; (Walker) Crntaegils Pnracolophn 1norrisor7i MD 19 16 (Baker Palearctic (e Asia) bamboos (Baker) 1919) Prrr7trphis jugltrizdis OR, 1928 (USNM) Palearctic (e Asia) pest Jirgln~zsregin (Goeze) VOLUME 108. NUMBER 3

Table 1. Continued.

Earlicsl N.A. rccord (1) Gcogrilphic O~.igin/ Slatrls in Norlh Amcr.lca/ North Anicrican In~roduccdSpecics (Collccrion or Rcfcrcncc) Reference (2) Reference (2) Economlc Hosr(s) Pen7phigrts brtrsctrirts NY, 1872 (Fitch Palearctic (Europe) pest Populrts riigrn; (Linnaeus) 1872) Lnctrrcn sati~~a Pernplzigr~ss/~y~'othee~~f QC, 1972 (Alleyne and Palearctic (Europe) Poprrlrls nigrn Passerini Morrison 1974) Pe/7tn/or7irr tzigroner~~osrr DC, 1909 (USNM) Unknown (either pest Mrlsn Coquerel Neotropical or (banana); Oriental) ornamental Musaceae, Araceae, Zingiberaceae Per.i/~Iiy/hrscalifor~tiierisis WA, 1903 (Essig and Palearctic (e Asia) Acer (Shinji) Abernathy 1952) Essig and Abernathy 1952 PerG~Iz~Nr1s/)iro~~ictr~s VT, 1847 (Fitch notes Palearctic (Europe) Acer (Kessler) [as Apliis nceris L.]) plntarioides Periplzyllrls BC, 1924 (Essig and Palearctic (Europe) Acer testudinaceus Abernathy 1952) (Fernie) Phloeoil7yzus passerir~i NB? 1950, ME 1970 Palearctic (Signoret) (22) (Smith 1974a [as P. rlfar,borr~iSmith]) P/7orodo17 11~1112~11i NY, 1863 (Walsh Palearctic (Europe) pest P~LI~LIS; (Schrank) 1863) Hrrr?iulrls PIi~~llcrp/2isfi~gi IL, 1879 (Thomas Palearctic (Europe) pest Fag~ls (Linnaeus) 1879) Pleotrichopltor~~a CA, 1910 (USNM) Palearctic (Asia?) minor pest chrysanthelnums rhrysnt~t/~er?~i (Theobald) Pleotrichophor~ts CO, 191 5 (USNM) Palearctic (Europe) A177br,osicr, glnr~du/osus Arternisicr (Kaltenbach) Procij)/~il~~ssylostei ME, 1914 (Patch 1917) Palearctic Lor~iceru;roots (De Geer) (23) of Piceo Pse~~rlocn~~dellcrr~rbiclrr NC, TN, 1973 (Smith Palearctic (Europe) (Bomer) and Knowlton 1975) Pserrdoregti~npor~icolrr FL, 1928 (FSAC) Oriental (Takahashi) PterocaNis uh~i IL, 1910 (Davis 1910) Palearctic (Europe) (De Geer) Pterocon?nicr pilosr~ri~ UT, 1958 (Knowlton Palearctic Buckton 1958) Pter.ocot~~rna~O~JLI~~L~~II MT, 1904 (USNM) Palearctic (Europe) (Kaltenbach) Pteroror~~r~za.srrlici.s MN, 1886 (Oestlund Palearctic (Linnaeus) 1886) RI~o~JNIo~~~~~zLIslor~icerrre VA, 1912 (USNM) Palearctic (Europe) minor pest Lor~icera (Siebold) R/IOIJ~~~S~J/IO~I~~LI.SBC, 1975 (Forbes and Palearctic (e Asia) hydrarzgeoe Chan 1976) (Matsumura) R/IO]J~~OS~J~OII~~LISCA, 19 12 (USNM) Palearctic minor pest Bulbs; Solnrzr~m latysip/~ori(Davidson) tuberosro?~ 594 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table I. Continued.

Earliest N.A. record (1) Geographic Origin1 Slatus in North America1 North American Introduced Species (Colleclion or Rererence) ReTelence (2) Rererence (2) Economic HOSL(S) Rltopalosiphoninus BC, 1953 (ACNB) Palearctic (Europe) minor pest Polyphagous staphyleae (Koch) on roots, including stored bulbs and rootcrops Rhopalosiplzt~nzniuidis NY, 1851 (Fitch Palearctic (e Asia) serious pest Gramineae (Fitch) (24) 1856) Rlzopalosiphum NE, 1890 (USNM) Palearctic pest1 biological various aquatic nymphaeae (Linnaeus) Davidson 19 17 control plants Rhopalosiphunz NY, 1856 (Fitch Palearctic (e Asia) pest Gramineae ruj5abdorninule 1856) (Sasaki) (24) Scliizaphis granzinum MD, 1884 (Pergande Palearctic serious pest Gramineae (Rondani) 1902) rninuta FL, 1998 (Halbert et Oriental (van der Goot) al. 2001) Schizaphis palustris BC, 1974 (Forbes and Palearctic (Europe) (Theobald) Chan 1976) Schizaphis rotundiventris FL, 1988 (Halbert et Palearctic (e Asia) or (Signoret) al. 2000) Oriental pineti BC, 1961 (CNC) Palearctic (Fabricius) Shivaphis celli Das (25) GA, SC, 1996 Palearctic (E Asia) nuisance pest Celtis (Halbert et al. and Oriental Halbert and Choate 2000) 1999 Siplza eleguns NB, 1949 (ACNB) Palearctic Black- minor pest Gramineae Del Guercio man & Eastop 2000 glyceriae ME, 1909 (Patch Palearctic pest Gramineae (Kaltenbach) 1910) (incl. rice) avenue NY, 1851 (Davis Palearctic major pest Davis Gramineae (Fabricius) 1914) 1914 Sitobion fiagariae CA 1916 (USNM) Palearctic (Europe) pest Gramineae, (Walker) Rubus Sitobion luteum FL, 1940 (FSCA) unknown minor pest greenhouse (Buckton) (26) orchids Sniynthurodes betae DC, 1895 (USNM) Palearctic minor pest Stoetzel polyphagous Westwood (Mediterranean) et al. 1996 on roots incl. Pltaseolus, Solanurn, Gossypiunz Subacyrthosiplzon IA, 1966 (USNM) Palearctic (Europe) cryp~obiumHille Ris Lambers TalcecuNis arundicolr,zs CA, 1903 (USNM) Oriental bamboos (Clarke) Tulcecullis arundinariae CA, 1911 (Essig 1917) Oriental bamboos (Essig) Takecullis tairvartus FL, 1930 (USNM) Oriental bamboos (Takahashi) fusijormis FL, 1959 (FSCA, Palearctic (e Asia) possible pest Ulmus?; roots Matsumura (27) USNM) I Oriental of Gramineae inc. rice Tetraneura ulrni NE, 1890 (USNM) Palearctic (Linnaeus) Iysimachiae NY, 2000 (FSCA) Palearctic (Europe) Populus, Borner (28) Lysimachiu VOLUME 108, NUMBER 3 595

Table 1. Continued.

Earliest N.A. record (I) Geographic Origin1 Status in North America1 North American Introduced Species (Collection or Reference) Reference (2) Reference (2) Economic Host(s) riehmi KS, WA, 1948 Palearctic (Europe) Trifolium (Borner) (USNM) NY, 1879 (USNM) Palearctic (Europe) pest Leguminosae, (Monell) (29) esp. Trifolium and Medicago LA, 1925 (USNM) Oriental pest Mizell et al. Lagerslroemia kahawaluokalani 2002 indica (Kirkaldy) BC, 1927 Palearctic (Europe) (Kaltenbach) (Glendenning 1929) Tinocallis sultans ID, 1986 (Halbert Palearctic (e Asia) Ulmus (Nevsky) and Pike 1990) TinocaNis talcachihoensis MD, 1996 (USNM) Palearctic (e Asia) Ulmus Higuchi Tinocallis ulmiparvifoliae CA,FL, 1998 (Gill Palearctic (e Asia) nuisance pest Ulmus Matsumura 1998b, Halbert 1998) TinocaNis zelkorvae CA, 1983 (Kono Palearctic (e Asia) (Takahashi) 1983) CA, 1907 (USNM) Palearctic (e Asia) major pest polyphagous (Boyer de or Oriental Fonscolombe) FL, 1995 (Halbert Palearctic (e Asia) major pest Halbert Citrus (Kirkaldy) and Brown 1996) or Oriental and Brown 1996 rara Mordvilko CO, 1929 (USNM) Palearctic (Europe) roots of Compositae Trichosiphonaphis FL, 1974 (Smith and Palearctic (e Asia) Polygonum polygoni (van der Denmark 1982) punctatum Goot) annulatus WA, 1951 (USNM) Palearctic (Europe) (Hartig) (30) Tuberculatus querccus BC, 1999 (CNC) Palearctic (Europe) (Kaltenbach) Tuberolachnus IL, 1872 (Le Baron Palearctic (Europe) minor pest Salix (Gmelin) 1872) achilleae CA, 1966 (Hille Ris Palearctic (Europe) Achillea (Koch) Lambers 1966) Uroleucon cirsii ON, 1956 (CNC) Palearctic (Europe) Cirsium (Linnaeus) DC, 1892 (USNM) Palearctic (Europe) (Linnaeus) Uroleucon taraxaci NY, 1902 (USNM) Palearctic (Europe) Taraxacum (Kaltenbach) Vesiculaphis caricis VA, 1916 (USNM) Palearctic (e Asia) (Fullaway) vaccinii IL, 1929 (USNM) Palearctic (Europe) (Theobald)

1. Names of Canadian 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 on Blackman and 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 with various host associations. We have not distinguished records for the various forms. 4. Aphidounguis mali: Single collection from botanical garden. 5. : Previously infrequent on , the recent upsurge of this species on alfalfa may represent a separate introduction of an alfalfa-feeding population from Asia comparable to the situation in Therioaphis trifolii (A. craccivora is known as the alfalfa aphid in China). 596 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Continued.

6. Aplzis euplzorbiae: Listed in Williams (191 1) from NE: four slides in USNM, apparently from this collection, are not A. euplzorbioe. 7. Aphis fubue and Aplzis runiicis: The latter name was used in the past for the aphid now known as A. jabae. The earliest occurrence of true A. tumicis is thus uncertain. Aphis fubae is currently used for a complex of taxa usually considered as subspecies. No attempt has been made here to distinguish the members of this complex. See also under 'A. ellonyrizi' in Table 2. 8. Aphis polygorzurcr: Single record from flight intercept trap. 9. Aphis turaxicicola: Single record from trap. 10. Braclzycaudus rurnexicolerzs: Considered by Holman (1965) to be introduced into Europe from North America. However, the only recorded host, Runies oceiosellc~, is a Palearctic species naturalized in North America (Scoggan 1978). 11. Dysaplzis cratnegi: A number of similar Dysuphis species use Q.ntaegus as primary host. Early reports of D. crutnegi (e.g., Thomas 1879) based on association with Crotaegus are not verifiable. 12. Dysuplzis foerzicul~rs:Single record. 13. Eriosorna gross~~laricreand E. ulriii: These names were long considered synonyms, but Danielsson (1982) showed that two species were involved. Reexamination of older North American collections named 'ulriii' has shown that most, if not all, earlier collections are E. grossuluriae. The 2003 collection of E. ulriii in CNC is currently the earliest confirmed record of this species in North America. 14. Eucernplzis betulc~eand E. purzctipennis: These species have been confused in the past and the correct identity of specific records is uncertain. Also, E. betulue, has been used to cover a complex of related non-European taxa. Most records of E. betulae and E. purzctiperzrzis from Berulc~papyrifera are actually specimens of the recently recognized species E. papyrijericola Blackman and subarctic records from dwarf birches probably E. borealis Blackman. 15. Geoicu utr~icularia:Probably represents a complex of species (Blackman and Eastop 2000). 16. Hyulopterus ariiygrlnli and H. pr~mi:There is confusion of these two species in North America, and the exact status of individual records is uncertain. 17. Hyperonzyz~ls rlzirzurztlzi: Distribution is alpine and northern Europe, Iceland, and northeastern North America. The North American population may represent part of a natural transatlantic distribution, rather than a human-mediated introduction. 18. Lipuplzis pseudobrassicae: Records from past few decades as L. er.ysirizi (Kaltenbach). 19. Macrosiplzurii lilii: First described from bulbs imported from Japan. However, there have been no subsequent Japanese records (Blackman and Eastop 2000) 20. Microloplziurn: This is in need of revision and it is difficult to say with clarity which species are present in North America. 21. Mir7rhrus nbietirzus: Voegtlin (1995a) believes that North American populations may represent a different species from Eurasian populations. The numerous records of this species from spruce are almost all M. obliquus. 22. Pliloeornyrus pusser.ini (Signoret): Smith (1974a) refers to a collection by N.R. Brown from "Canada" in 1950. This particular collection has not been located, but other collections by Brown in ACNB from this period are all from NB. 23. xylostei: Some early North American records of P. sylostei were actually based on specimens of Hyudaplzis foerziculi (=Aphis xylostei Schrank) misfiled in USNM. 24. Rhopalosiphuriz riinidis and R. r~~fi~~brloriiirzale:Both species are assumed to be of Asian origin because complete holocyclic lifecycles are documented only in Asia. Holocyclic colonies of R. muidis were found on P~UIZLIScor~zutu in Pakistan (Remaudiere and Naumann-Etienne 1991). Similarly, holocyclic colonies of R. rufiubdoniiule occur on Prurzus in Japan (Blackman and Eastop 2000). However, a holocycle for R. r~lfc~bdorninulealso may occur in North America. (Al-Raeesi et al. 1992). There is a great deal of genetic diversity in R. ninidis populations in North America (Blackman et al. 1990). This genetic diversity may point to a North American origin for the species, or it may be the result of mutations accumulating in continuous clonal lineages. Most other Rhopalosipl7urii species, including Rhopulosipl7um pocli (L.), are thought to be of North American origin (Halbert and Voegtlin 1998). 25. Slzivuphis celti: Probably bonsai introductions 26. Sitobiorz luteuri?:Earlier North American records in USNM represent port interceptions. 27. Tetrc~rzeurujusif'orri7is: Records as T nigriabdonzinulis (Sasaki). See Eastop and Blackman (2005). 28. Thecabius lysi~izuclzicre:A 1956 collection by Hille Ris Lambers in Montreal, QC was questionably assigned to this species by Hille Ris Lambers (Smith 1974b). Recent collections in New England lend credibility to his determination. 29. Theriouphis trifolii: Eastern populations feed on clover. Alfalfa feeding populations in western USA represent a separate introduction formerly known as Tlzerioc~phis muculatu (or Pierocallidiu~ii nznculaturiz). 30. Tuberculatus nnnulutus: Record bv Lintner (1889) (as Auhis uuercus) not verified. VOLUME 108, NUMBER 3 597

Table 2. Species previously reported as introduced in North America, but now considered to be erroneous or dubious records.

Species Reason for Consideration as Erroneous or Dubious - Anzphoropkora itlbi May not occur in North America. References to this species on raspberry (Kaltenbach) in North America are mostly misidentifications of Amplzorophora agatlzonica Hottes (Blackman and Eastop 2000). However, Hille Ris Lambers believed A. rubi was present in northwestern North America (Hille Ris Lambers letter to L. Russell, 13 Nov. 1972, USNM letter file). Note that the European aphid on raspberry is actually A. idaei (Borner) while A. rubi is restricted to blackberry (Blackman et al. 1977). Anurr!phis fr~rfrirae (Koch) Listed in Bodenheimer and Swirski (1957) as occurring in North America but no supporting specimens located. Anuraphis stlbterranea Cited in Jones (1929) from Colorado. No specimens are present in (Walker) USNM. Aphis frangulae Kaltenbach North American records probably refer to Aphis nast~irtii.Aphis fiangulae is a member of a complex of taxa including Aphis gossypii Glover (Smith and Parron 1978, Stroyan 1984, Blackman and Eastop 2000). The overwintering host for A. fangulae sensu strict0 is Fiangulr~ alnus (= frangula). A. nasturtii (as A. rlzamni Boyer de Fonscolombe, see note below for Macchiatella rhamni) is the only Aplzis reported on Rhamntls in Illinois by Hottes and Frisson (193 1). Aplzis acetosae Linnaeus Listed in Borner and Heinze (1957) as occurring in North America, but no specimens have been located Aphis evonyrni Fabricius There is considerable confusion in the application of the name A. evonyrni (or euonymi). It is probable that most, if not all, of the North American records of A. evonynli (or A. euonymi) refer to other members of the Aphis fabae complex. Eastop and Hille Ris Lambers (1976) state that A. evonymi of authors prior to 1950 is a synonym of Apkisfabae solanella Theobald. Eastop and Hille Ris Larnbers (1976) listed A. euonymi Fabricius 1775 as a valid species, and Remaudiere and Remauditre (1997) treat it as a subspecies of A. fabae. Aphis nzedicriginis Koch North American records of this species are misidentifications of Aphis craccivora. Aphis grossulnrine Listed in Wilson (1913) as found in Oregon. No specimens are present in Kaltenbach USNM. Aplzis ilicis Kaltenbach Reported from California and British Columbia in Smith & Parron (1978). No supporting specimens have been located. Aplzis hieracii Schrank Listed in Rafinesque (1818). No North American specimens have been located. Aplzis nepetae Kaltenbach Listed in Smith and Parron (1978) as occurring in Idaho and Minnesota. There are no North American records in the CNC, FSCA, or USNM. Aphis plantaginis Goeze Listed in Smith and Parron (1978) as occurring in New York. No North American specimens have been located Aplzis ubnariae Schrank Listed in Smith and Parron (1978) from various states. Specimens from Colorado in USNM were actually A. spiraecola. Record by Thomas (1879) is A. pisum. Aplzis viburni Scopoli Recorded in Illinois as early as 1879 (Thomas 1879). However, records are likely the result of confusion with A. fabae and A. viburniplzila Patch. Aplzis vitis Scopoli Listed in Walsh (1863) from Illinois and Missouri. A single slide of poor quality found in USNM identified as A. vitis is likely Aphis illinoisensis. Brackycaudus prunicola Listed in Hunter (1901) as occurring in North America. There are no (Kaltenbach) North American records for this species in the CNC, FSCA, or USNM. Blackman and Eastop (2000) state that this species is "not yet recorded outside Europe." 598 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Continued. Species Reason for Consideration as Erroneous or Dubious Capizophorus archangelskii Single record from New York in 1960 (Leonard 1963). Probably C Nevsky pakansus. Borner and Heinze (1957) considered C. pakansus a synonym of C. archangelskii. Cinara piceicola North American records refer to (Hartig) (Cholodkovsky) Cinara piceae (Panzer) No North American specimens found. Cinara pini (Linnaeus) North American records are doubtful. See Smith and Parron (1978). Cinara pinicola (Kaltenbach) No authentic North American specimens found. Records of this species appear to refer to C. pinea and C. pilicornis. angelicae (Del A single published record from New York in 1913 (Olsen 1921) Guercio) represents the only record of this species in North America. There are no North American specimens in CNC, FSCA, or USNM. Cryptaphis poae (Hardy) Recorded as occurring in Colorado and Manitoba (Robinson and Bradley 1965), Robinson later stated (correspondence cited in Smith and Parron 1978) that he did not believe C. poae occurs in North America. angelicae (Koch) Recorded by Wilson (1909) for California. US specimens in USNM are Aphis species and 2 alates and alatoid nymphs of a Dysaphis species that key to crataegiltulipaelangelicae/pyrilradicola complex. The status of this species in North America is uncertain. Dysaphis sorbi (Kaltenbach) First listed by Clarke (1903) for California. Although recorded in some subsequent literature as present in North America, specimens identified in USNM as D. sorbi were actually D. plantaginea. Stroyan (1985) did not list D. sorbi as present in North America. holci (Hille Ris Although recorded as occurring in various localities in North America Lambers) (Smith and Parron 1978), D. holci probably does not occur in North America (Halbert et al. 1992). lanigerum Generally considered to be native to North America. However, as early (Hausmann) as 1841, Harris (1841) believed otherwise and cited various sources and observations to support his view. Eastop (1973) suggests the origin may be eastern Asiatic and introduced to Europe via America. Eriosoma lanuginosa (Hartig) Probably not in North America; records refer to E. pyricola (Blackman and Eastop 2000). Geoica luclyuga (Zehntner) Probably not in North America; Smith (1985) believed records dubious. No specimens located. Glyphina jacuzensis Mordvilko Listed as G. alni (Schrank) in Archibald (1957). Record is doubtful; probably actually G. longiseta Richards, but specimens not present in Archibald collection (now in CNC). roboris (Linnaeus) Smith and Parron (1 978) record from California, Connecticut, and Maine. No North American specimens have been located; material seen from these states identified as L. roboris is Longiszigma caryae. Macchiatiella rhamni Many North American records, but Aphis rhamni Boyer de Fonscolombe (Boyer de Fonscolombe) has been misapplied in the past by many authors to the species now called Aphis nasturtii Kaltenbach (Eastop and Hille Ris Lambers 1976). The presence of M. rhamni in North America is doubtful. Macrosiphum funestum Recorded by Burnham (1938) for New Brunswick in 1936. No slides have (Macchiati) been located in (CNC) or ACNB. Macrosiphum gei (Koch) Recorded from various localities in North America (Smith and Parron 1978). All North American specimens in FSCA and USNM were actually . Blackman and Eastop (2000) state that most North American records refer to M. euphorbiae. submacula North American records probably all refer to M. sgpkensis Hille Ris (Walker) Lambers (Blackman and Eastop 2000) viciae Buckton Listed in Smith and Parron (1978), citing Miiller (1969). However, Miiller (1969) actually states "NAs" while he uses "NAm" for North America. VOLUME 108, NUMBER 3

Table 2. Continued

Species Reason for Consideration as Erroneous or Dubious Metopolophium frisicum Hille Listed in Smith and Parron (1978), citing Borner (1952). However, no Ris Lambers specimens of this species have been located in North American collections. Myzus ajugae Schouteden Smith and Parron (1978) considered the Leonard (1968) reference to "?Myzus ajugae" suspect, believing "the determination was made because of the host association." Myzus myosotidis (Borner) Smith and Parron (1978) cite Hille Ris Lambers (1966) as the basis for the occurrence of M. myosofidis in California. However, Hille Ris Lambers (1966) mentions this species only in a key. No specimens are present in USNM. Halbert et al. (1995) considered North American distribution doubtful. 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 (Moreira) of specimens in USNM showed the collection to be from "Florida, P.R." (Puerto Rico), not Florida, U.S.A. tremulae North 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). aceris (Linnaeus) Although stated by Blackman and Eastop (1994) as introduced to North America, and recorded by various other authors, all North American collections in USNM are interceptions. Essig and Abernathy (1952) believed that reports of P. aceris were referable to other species and P. aceris had not been recorded in the Western Hemisphere. Prociphilus bumeliae (Schrank) According to Smith and Parron (1978), probably misidentifications of P. americanus. Saltusaphis scirpus Theobald A single record from Ohio, 1938 (Smith 1940). No North American specimens in USNM, CNC or FSCA. Schoutedenia ralumensis A single specimen, cited in Palmer (1952) (as Setaphis viridis van der Riibsaamen Goot) in Colorado. Not established. dauci (Fabricius) Records from District of Columbia and Maryland (Leonard and Bissell 1970, Smith and Parron 1978) may have been misidentification of . Sitobion luteum (Buckton) Cited in Smith and Parron (1978) as occurring in New Mexico and New York. All North American records in USNM are port interceptions and do not represent established populations. Thecabius affinis (Kaltenbach) Thecabius affinis is an Old World species, and Th. populiconduplifolius North American. They are morphologically and biologically indistinguishable, but Th. afjinis has 2n = 38, and Th. populiconduplifolius has 2n = 28 (Blackman and Eastop 1994). It is unknown whether Th. affinis occurs in North America. dryophila (Schrank) References to this species probably represent misidentification of Thelaxes 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 Ris Lambers 1974). Titarzosiphon artemisiae (Koch) Questionably recorded for New York by Smith and Parron (1978), citing Hunter (1901) who lists "Nectarphora arternisiae? Koch." Probably a Macrosiphoniella species. Uroleucon composilae Listed in Hille Ris Lambers (1963) as present in "America." Although (Theobald) introduced into the Caribbean, this species has not yet been found in North America. 600 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Continued. Species Reason for Consideration as Erroneous or Dubious Uroleucon nzuralis (Buckton) In Nebraska, according to in Williams (191 1). However, Robinson (1985) examined specimens from the USNM and determined that they were Uroleucon sonchellui7~(Monell). He believed U. rnuralis does not occur 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 North America." Uroleucon tanaceti (Linnaeus) Cited in Smith and Parron (1978) as occurring in several localities. However, Robinson (1985) stated that he had "not seen authentic material of this species from North America."

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

Aphis equiseticola Ossiannilson The single previous North American record from Vancouver, BC suggested an introduction of this species otherwise known from Scandinavia. However, a collection (in CNC) of ths species in 2004 from northern Qukbec indicates a possible circumboreal distribution. Aphis farinosa Gmelin Widespread in both Nearctic and Palearctic. Boernerina alni Takahashi Subspecies alaskensis Hille Ris Lambers and Hottes (originally described as a subspecies of B. variabilis Richards), widespread in Alaska and arctic Canada, is the North American segregate of this species otherwise known from east Asia. Cavariella aquatica (Gillette and Bragg) Widespread in both Nearctic and Palearctic, but infrequently collected. Takahashi According to Stroyan (1969), differences in host range among Old and New World populations indicate the possibility that this name covers a complex of taxa. Clzaitoplzorus horii Takahashi Specimens from Northwest Territories match the nominate Asian subspecies, rather than the European subspecies (beuthani Borner), and probably represent natural migration across the Bering Strait. Gllaresia poluniizi Stroyan Known from mountains of western North America and Himalayas. Maculolachnus sijpkensis Hille Ris Distributed across North America and in Mongolia (Blackman Lambers and Eastop 2000). Mindarus obliquus (Cholodkovsky) Present across boreal forest zone of North America, northern Europe, and northern Russia, but apparently not recorded in eastern Siberia. Muscaphis escherisclzi (Borner) According to Stekol'shtshikov and Shaposhnikov (1993), Toxopterella rirepanosiphoides MacGillivrary and Bradley (known from across Canada, and in northern Europe as subspecies irae Shaposhnikov) is the primary host form of M. escherisclzi. Myzodiuni modestum (Hottes) Widespread in both Nearctic and Palearctic. Thripsaphis cyperi (Walker) Although first North American records of this species in Alaska (USNM) and Ontario (CNC) date from 1961, it is distributed across boreal and subarctic Canada and is likely naturally circumboreal. Uroleucon simile (Hille Ris Lambers) Widespread in Palearctic. North American records (Alaska only) likely represent a natural expansion across Bering Strait. VOLUME 108, NUMBER 3 60 1

Before 1840- 1870- 1900- 1930- 1960- 1990- 1840 1869 1899 1929 1959 1989 present 30- intervals

Fig. 1. Adventive aphids. Number of adventive aphids in North America north of Mexico recorded at 30-year intervals. Two species not included due to confusion with related species.

(1997), except for changes noted by causing damage to several plant families Eastop and Blackman (2005). The taxo- are listed as polyphagous. nomic status and biogeographic origins A summary of species previously listed of some aphid species are difficult to for North America, but for which we determine conclusively due to ongoing consider their occurrence doubtful or taxonomic problems. We have summa- erroneous, and an explanation for their rized relevant information in footnotes exclusion is provided in Table 2. Table 3 to Table 1. contains a compilation of those species, Parameters for determining a pest are some of which were previously suspected as defined in D. R. Miller et al. (2002), as adventive to North America, which Miller and Miller (2003), and Miller et al. we consider to be most probably (2005). That is, if statements are made either Holarctic or Beringian in distribu- that a species is a pest, causes harm, tion. requires control measures, is economi- cally important, or reflects some compa- rable statement or inference, then we RESULTS consider that species a pest. Pest status Based on our findings there are 1,416 entries in Table 1 without reference to aphid species in North America north of other authors are either from Blackman Mexico, which comprises about 30% of and Eastop (1994, 2000) or our personal the world fauna of approximately 4,700 experience. The principal hosts listed are species (Remaudiere and Remaudiere those of the greatest economic impact in 1997). Table 1 provides information on North America (including crops, orna- 262 aphid species (1890 of the North mental plants, trees and weeds). Species American and approximately 6% of the PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Deast AsialOriental

HPalearctic (general) npalearctic (east Asia)

WPalearctic (cenbal Asia)

Dates

Fig. 2. Adventive aphids. Number of adventive aphids in North America north of Mexico at 10-year intervals classified by biogeographic origin. Aphids of Neotropical, Afrotropical and unknown origin combined into single category ("Other"). Two species not included due to confusion with related species. world aphid fauna) that are considered arctic Region (227), followed by the as adventive to North America. Oriental (1 5), Neotropical (4), Australa- There has been an overall increase in sian (2), and Afrotropical (1). Eight number of aphid introductions per year other species are generally distributed in from 1791 until 1929 (Fig. 1, number of east Asia, in both Oriental and Palearctic introductions plotted at 30-year intervals regions. The geographic origins of five in to smooth short term anoma- species are unknown. As shown in other lies), and there has been an overall analyses (Sailer 1978, Miller et al. 2005), decrease to the present. There are, most insects adventive to North America however, a number of irregularities in originate in the Palearctic Region. We this general trend that appear when first have further examined the rate of Asian records are plotted on a finer time scale introductions by subdividing the Pale- (Fig. 2, number at 10-year intervals, and arctic Region into European, central Fig. 3, cumulative number at 10-year Asian and east Asian components. Of intervals). Two species, Aphis rumicis L. the aphids originating from the Palearctic and Hyalopterus amygdali (Blanchard), Region, 144 are found in Europe (and are excluded from Figs. 1 to 3 due to adjacent parts of north Africa and west uncertain introduction dates because of Asia), 11 in central Asia, 25 in east Asia, confusion with related species. and 48 are widespread in Eurasia. Orig- The geographic origin of adventive inal North American populations of the aphids is illustrated in Fig. 2 and Fig. 3. majority of the widespread Palearctic The majority originated from the Pale- species likely originated from Europe. VOLUME 108, NUMBER 3 603

Dates Fig. 3. Adventive aphids. Cumulative number of adventive aphids in North America north of Mexico at 10-year intervals classified by biogeographic origin. Aphids of Neotropical, Afrotropical and unknown origin combined into single category ("Other"). Two species not included due to confusion with related species.

Of the 262 adventive aphids, 114 quent decrease in successful entry and (about 44%) are considered to be pests. establishment of species. There are Of these, 20 (>17%) are recorded as also finer scale changes in the number of either serious or major pests (i.e., causing first detections (Fig. 2), apparently due extensive or long term damage). to a combination of major geopolitical and economic events, resulting in real changes in rates of introduction, and the The total number of adventive aphids efforts of particular individuals in re- in North America north of Mexico (262) cording the aphid fauna. For example, is remarkably similar to the 255 adven- the peak in the number of first records in tive scales insects (Miller et al. 2005). the 1870's is almost entirely due to the Also, the decrease in annual number of work of Thomas (1879), and there is introduced aphids since 1929 as shown in a major trough in the 1930's apparently Fig. 1 is mirrored in the pattern seen in related to the economic depression and adventive Coccoidea (e.g., Miller et al. associated decrease in trade activity, 2005). Sailer (1978) suggested that en- despite active work on aphids by Essig, forcement of the United States Plant Gillette, and others during this period. Quarantine Act of 19 12 has resulted in As for insects in general (Sailer 1978, greater rate of interception and conse- 1983) and for scale insects in particular 604 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(Miller et al. 2005), most adventive LITERATURECITED aphids are indigenous to the Palearctic Alleyne, E. H. and F. 0. Morrison. 1974. Some Region (especially Europe). This is to be Canadian poplar aphid . Canadian Ento- expected, given the immigration history mologist 109(3): 321-328. of the continent and the associated Al-Raeesi, A. A., S. E. Halbert, and J. B. Johnson. 1992. Rlzopnlosiplzun? rufiiabclolninnlis (Sasaki) transport of crops and their weeds. and Aplzis arrnoraceae Cowen (: However, European introductions have Aphididae) confirmed on wheat in Idaho. Pan- become relatively few (Fig. 2, and seen Pacific Entomologist 68: 70-71. as a leveling off of the European Anonymous. 2005. Soybean aphid in Minnesota. component of the cumulative curve in http://www .soybeans.umn.edu~crop/insects/aphid/ Fig. 3). Asian species now account for aphid.htm [accessed 4 May 20051. Archibald, K. D. 1957. Report of the forest a major proportion of new introduc- Aphidae (Homoptera) of Nova Scotia together tions, although the total number is with species and host plant lists. Annals of the similar to that in the early part of the Entomological Society of America 50: 3842. 20th century (Fig. 2). Because the Asian Baker, A. C. 1919. An undescribed species of fauna was poorly known at the time, Dryopen injurious to Plzyllostaclzys. (Aphidi- dae-Hom.). Proceedings of the Entomological several of these early oriental introduc- Society of Washington 21: 106106. tions were first described from popula- Baker, A. C. and W. M. Davidson. 1916. Wooly tions in the U.S.A. (Aphis spiraecola pear aphis. Journal of Agriculture Research 6: Patch, Atarsaphis agrifoliae Ferris, Eu- 351-360. thoracaphis umbellariae Essig, Neotoxop- Bartholomew, P. S. 1932. Six new species of aphids, with records of other species new to California. tera olivieri (Essig), Paracolopha morri- Annals of the Entomological Society of Amer- soni Baker, Periphyllus californicus ica 25(4): 713-729. (Shinji), arundicolens (Clarke), Bernal, J., D. Gonzhlez, E. T. Natwick, J. G. Loya, R. T. arundinariae (Essig)) as was the Leon-Lopez, and W. E. Bendixen. 1993. Natural African aphid Aloephagus myersi Essig. enemies of identified in California. California Agriculture 47: 2&28. The proportion of pests among the Blackman, R. L. and V. F. Eastop. 1994. Aphids adventive species is much greater than on the World's Trees: An Identification and the proportion among the native species. Information Guide. CAB International, Wall- This to be expected, given that most of ingford, UK. 1,004 pp. these introductions were enabled by . 2000. Aphids on the World's Crops: An Identification and Information Guide. Second human activity, and hence associated edition. John Wiley & Sons Ltd., England. with economically important plants. A 466 pp. more detailed analysis of the economic Blackman, R. L., V. F. Eastop, and M. Hills. 1977. impact of adventive and native species is Morphological and cytological separation of in preparation. An?plzo~~ol~horaBuckton (Homoptera: Aphidi- dae) feeding on European raspberry and ACKNOWLEDGMENTS blackberry (Rubus spp.). Bulletin of Entomo- logical Research 67: 285-296. We thank D. Creel and J. Eibl Blackman, R. L., S. E. Halbert, and T. W. Carroll. (USDA, ARS, Systematic Entomology 1990. Association between karyotype and host Laboratory, Beltsville, MD) for their plant in corn aphid (Homoptera: Aphidi- dae) in the northwestern United States. Envi- assistance with reference verification for ronmental Entomology 19: 609-61 1. this paper. We also thank P. Mason Bodenheimer, F. S. and E. Swirski. 1957. The (AAFC, Ottawa), E. Grissell (USDA, Aphidoidea of the Middle East. Jerusalem. ARS, Systematic Entomology Laborato- 378 pp. ry, Washington, DC) and D. Nickle Borner, C. 1952. Europae centralis Aphides. Die Blattlause Mitteleuropas. Namen, Synonyme, (USDA, ARS, Systematic Entomology Wirtspflanzen, Generationszyklen. Mitteilun- Laboratory, Beltsville, MD) for their gen der Thiiringischen Botanischen Gesell- manuscript reviews. schaft 3: 1488. VOLUME 108, NUMBER 3 605

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. 1872. Fourteenth report of the noxious, Gittins, A. R., G. W. Bishop, G. F. Knowlton, and beneficial and other insects of the State of New E. J. Parker. 1976. An annotated list of the York. Transactions of the New York State aphids of Idaho (Homoptera: Aphididae). Agriculture Society 30: 355-381. University of Idaho College of Agriculture Forbes, A. R. and C. K. Chan. 1976. The aphids Agricultural Experiment Station Research (Homoptera: Aphididae) of British Columbia Bulletin 95: 47 pp. 4. Further additions and corrections. Journal Glendenning, R. 1926. Some new aphids from of the Entomological Society of British Co- British Columbia. Canadian Entomologist 58: lumbia 73: 57-63. 95-98. Forbes, A. R. 1980. The aphids (Homoptera: . 1929. A new Callipterine from Victoria, B. Aphididae) of British Columbia 8. Further C. (Hemiptera-Aphididae). The Canadian En- additions. Journal of the Entomological Soci- tomologist 61: 237-238. ety of British Columbia 77: 3842. Hajek, A. E. and D. L. Dahlsten. 1986. Discrim- . 1983. The aphids (Homoptera: Aphididae) inating patterns of variation in aphid (Homo- of British Columbia 11. Further additions. ptera: Drepanosiphidae) distribution on Betula Journal of the Entomological Society of British pendula. Environmental Entomology 15(6): Columbia 80: 51-53. 1145-1 148. . 1984. The aphids (Homoptera: Aphididae) . 1987. The exotic aphids (Homoptera: of British Columbia 12. Further additions. Drepanosiphidae) on ornamental birch in Journal of the Entomological Society of British Northern California. Pan-Pacific Entomologist Columbia 81: 72-75. 63(4): 319-323. Forbes, A. R., B. D. Frazer, and C. K. Chan. 1974. Halbert, S. E. 1998. Triology. Entomology Section The aphids (Homoptera: Aphididae) of British 37(3), www.doacs.state.fl.uslpi/enppl98-may- Columbia 3. Additions and corrections. Jour- jun.htm [accessed 14 July 20051. nal of the Entomological Society of British . 2004. The genus Greenidea (Rhynchota: Columbia 7 1 : 4349. Aphididae) in the United States. Florida Forbes, A. R., B. D. Frazer, and H. R. MacCarthy. Entomologist 87(2): 159-163. 1973. The aphids (Homoptera: Aphididae) of Halbert, S. E. and L. G. Brown. 1996. Toxoptera British Columbia. 1. A basic taxonomic list. citricida (Kirkaldy), brown citrus aphid- Journal of the Entomological Society of British Identification, biology and management strat- Columbia 70: 43-57. egies. Florida Department of Agriculture and Fox, L. 1977. Cole crops. Insects. Cooperative Consumer Services Entomological Circular Plant Pest Report 2(10): 104. No. 374: 14. Gassmann, A., and L. T. Kok. 2002. Musk Halbert, S. E. and K. S. Pike. 1990. An Asian thistle (nodding thistle). In Van Driesche, R., aphid (Homoptera: Aphididae) new to North S. Lyon, B. Blossey, M. Hoddle, and R. America. Proceedings of the Entomological Reardon. 2002. Biological control of invasive Society of Washington 92(4): 672-674. plants in the Eastern United States. USDA Halbert, S. E. and G. Remaudiere. 2000. A new Forest Service Publication FHTET-2002-04. Oriental Melanaphis species recently intro- 413 pp. duced in North America (Hemiptera, Aphidi- Gill, R. J. 1998a. New state records: an aphid. dae). Revue fran~aised'Entomologie (Nou- California Plant Pest and Disease Report 17: 9. velle Serie) 22(2-3): 109-1 17. . 1998b. New state records: Chinese elm Halbert, S. E. and P. M. Choate. 1999. An Asian aphid. California Plant Pest and Disease Re- woolly hackberry aphid, Das port 17: 10. (Homoptera: Aphididae). Florida Department Gillette, C. P. 1908. New species of Colorado of Agriculture and Consumer Services Ento- Aphididae, with notes upon their life-habits. mological Circular No. 392: 1-2. Canadian Entomologist 40: 17-20. Halbert, S. E., S. J. Castle, and T. M. Mowry. 1995. . 1909. Plant louse notes, family Aphididae. Do Myzus (Neclarosiphon) spp. other than M. Journal of Economic Entomology 2: 385-388. persicae pose a threat to the Idaho crop? . 1910. Plant louse notes, family Aphididae American Potato Journal 72: 85-97. (Continued), Plate 26. Journal of Economic Halbert, S. E. and D. J. Voegtlin. 1998. Evidence Entomology 3(5): 403407. for the North American origin of Rhopalosi- . 1917a. Some Colorado species of the genus phum and barley yellow dwarf virus, Lachnus. Annals of the Entomological Society pp. 351-356. In Nieto, J. M., Nafria, J. M., of America 10: 133-144. and A. F. G. Dixon, eds. Aphid in natural and . 1917b. Two new aphid genera and some new managed ecosystems. Universidad de Le6n, species. Canadian Entomologist 49: 193-199. Leon, Spain. 688 pp. VOLUME 108, NUMBER 3 607

Halbert, S. E., B. J. Connelly, and M.-g. Feng. . 1974. On American aphids, with descrip- 1992. Diuraphis (Holcaphis)frequens (Walker), tions of a new genus and some new species on wheat, new to Idaho, and a key to North (Homoptera, Aphididae). Tijdschrift voor En- American Diuraphis. Pan-Pacific Entomologist tomologie 11 7(4): 103-155. 68: 52-61. Holman, J. 1965. Some unrecorded Middle Euro- Halbert, S. E., K. L. Hibbard, and D. J. Voegtlin. pean aphids. Acta Faunistica Entomologica 2001. Schizaphis rninuta (van der Goot) Musei Nationalis Pragae 11: 277-284. (Homoptera: Aphididae), new to the United Hottes, F. C. and E. 0.Essig. 1955. A new species of States. Insecta Mundi 15: 56. Cinara from Oregon (Aphidae). Proceedings of Halbert, S. E., G. Remaudiere, and S. E. Webb. the Biological Society of Washington 68: 61-63. 2000. Newly established and rarely collected Hottes, F. C. and T. H. Frison. 1931. The plant lice aphids (Homoptera: Aphididae) in Florida and of Illinois. Illinois Department of Registration the southeastern United States. Florida Ento- and Education Division of the Natural History mologist 83(1): 79-91. Survey Bulletin 19: 121-447. Haldeman, S. S. 1844. Descriptions of several Huber, D. M., M. E. Hugh-Jones, M. K. Rust, S. species of Aphis inhabiting Pennsylvania. R. Sheffield, D. Simberloff, and C. R. Taylor. Proceedings of the Boston Society of Natural 2002. Invasive pest species: Impacts on History 1: 168-169. agricultural production, natural resources, Harper, A. M., J. P. Miska, G. R. Manglitz, B. J. and the environment. CAST Issue Paper 20: Irwin, and E. J. Armbrust. 1978. The 1-18. literature of arthropods associated with alfalfa: Hunter, W. D. 1901. The Aphididae of North 111. A bibliography of the Pea Aphid, Acyrtho- America. Iowa Agricultural College Experi- siphon pisurn (Harris) (Homoptera: Aphidi- ment Station, Ames, Iowa. Bulletin 60: 63-138. dae). University of Illinois at Urbana-Cham- Jones, C. M. 1929. and their relation to paign, College of Agriculture, Agricultural aphids. Colorado Experiment Station Bulletin Experimental Station, Special Publication, 341: 1-96. 89 PP. Knowlton, G. F. 1958. Some Utah insects-~1958. Harris, P. 2003. Classical biological control of Utah State University Extension Service Mim- weeds. Established biocontrol agent-Aphis eograph Series 171: 1-16. chloris Koch. Foliage-sucking aphid. res2agr. Knowlton, G. F. and C. F. Smith. 1936. Aphid ca/lethbridgelweedbiolagents/aaphchl_e.htm[ac- studies. Festschrift zum 60. Geburtstage von cessed 13 July 20051. Professor Dr. Embrik Strand 1: 487492. Harris, T. W. 1841. 2. Plant-lice. (Aphididae). A Kono, T. 1977. Distribution and identification of Report on the Insects of Massachusetts, In- Blue Alfalfa Aphid jurious to Vegetation. Cambridge, Massachu- Shinji (Homoptera: Aphididae). Cooperative setts. 459 pp. Plant Pest Report 2(11): 153-156. Hayhurst, P. 1909. Observations on two species of -- . 1983. A Japanese Zelkova aphid new to the Hyalopterus (Aphididae). Journal of the U.S.A. California Plant Pest and Disease New York Entomological Society 17(3): 107- Report 2: 107-109. 115. Le Baron, W. 1872. Insects injurious to the . Hille Ris Lambers, D. 1949. Contributions to Report of the State Entomologist (Illinois) 2: a monograph of the Aphididae of Europe. 138-139. IV. The genera Aulacorthurn Mord., 1914; Leonard, M. D. 1963. A list of aphids of New Microlophiurn Mord., 1914; Hyalop- York. Proceedings of the Rochester Academy teroides Theob., 19 16; Idioprerus Davis, 1909; of Science lO(6): 289432. Coquerel, 1859; Arnphorophora . 1968. A supplement to a list of the aphids Buckton, 1876; Wahlgreniella nov. gen; of New York. Proceedings of the Rochester Megoura Buckton, 1876; Megourella nov. Academy of Science 1 1: 257-36 1. gen,; Hyperornyzus Borner, 1933; Naso- . 1970. New state records. USDA Cooper- novia Mordv., 1914. Temminckia 8: 182- ative Economic Insect Report 20: 773. 323. Leonard, M. D. and T. L. Bissell. 1970. A list of the . 1963. Notes on Surinam aphids. Land- aphids of District of Columbia, Maryland and bouwproefstation in Suriname Bulletin 80: Virginia. University of Maryland Agricultural 1-5. Experiment Station. MP 770. College Park, . 1966. Notes on California aphids, with Maryland. 129 pp. descriptions of new genera and new species Lintner, J. A. 1889. The Aphididae, or plant-lice. (Homoptera: Aphididae). Hilgardia 37(15): Transactions New York State Agricultural 569-623. Society 1883-1886: 101-1 15. 608 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

MacGillivray, M. E. 1954a. Note on My-LUS certus . 1887. Synopsis of the Aphididae of (Walker), an aphid new to North America Minnesota. Geological and Natural History (Homoptera: Aphidae). Canadian Entomolo- Survey of Minnesota Bulletin 4: 1-100. gist 86(4): 190. Olsen, C. E. 1921. Two season's collecting of . 1954b. Note on Don- Aphidae, principally on Long Island, New York, caster (Homoptera: Aphidae), an aphid new with notes on some of the species. Bulletin of to North America. Canadian Entomologist the Brooklyn Entomological Society 16: 14-19. 86(10): 454. Olsen, W. H. 1974. Dusky-veined walnut aphid Mason, P. W. 1940. A revision of the North studies. California Agriculture 28: 18-19. American aphids of the genus Myzus. U.S. Paddock, F. B. 1915. The turnip louse. Texas Department of Agriculture Miscellaneous Pub- Agricultural Experiment Station Bulletin 180: lication 371, 31 pp. 77 PP. Maw, H. E. L.,R. G. Foottit, K. G. A. Hamilton, and Palmer, M. A. 1952. Aphids of the Rocky G. G. E. Scudder. 2000. Checklist of the Hemi- Mountain region. The Thomas Say Founda- ptera of Canada and Alaska. NRC Rresearch tion 5: 1452. Press, Ottawa, Ontario, Canada. 220 pp. Patch, E. M. 1910. Four rare aphid genera from Miller, D. R., G. L. Miller, and G. W. Watson. Maine. Maine Agriculture Experiment Station 2002. Invasive species of (Hemi- Bulletin 182: 241-248. ptera: Pseudococcidae) and their threat to U.S. . 1911. Aphididae. Insect Notes for 1910. agriculture. Proceedings of the Entomological Maine Agriculture Experiment Station Bulletin Society of Washington 104(4): 825-836. 187: 20-23. Miller, D. R., G. L. Miller, G. S. Hodges, and J. A. . 1912. Food plant catalogue of the Aphidae Davidson. 2005. Introduced scale insects of the world. Part I. (A bibliographical (Hemiptera: Coccoidea) of the United States appendix to aphid pests of Maine.) Maine and their impact on U.S. Agriculture. Proceed- Agriculture Experiment Station Bulletin 202: ings of the Entomological Society of Washing- 179-2 14. ton 107(1): 123-158. . 1914. Currant and gooseberry aphids in Miller, G. L. and D. R. Miller. 2003. Invasive soft Maine. Maine Agriculture Experiment Station scales (Hemiptera: ) and their threat to Bulletin 225: 49-60. U. S. agriculture. Proceedings of the Entomolog- . 1917. Eastern aphids, new or little known. ical Society of Washington 105(4): 832-846. Part I. Journal of Economic Entomology Miller, G. L., M. B. Stoetzel, R. Lopez, and D. A. lO(4): 416420. Potter. 2002. Geoica setulosa (Passerini) . 1918. Food plant catalogue of the Aphidae (Hemiptera: Aphididae): New distribution re- of the world. Part V. Maine Agriculture cords for North America. Proceedings of the Experiment Station Bulletin 270: 47-100. Entomological Society of Washington 104(1): Pergande, T. 1902. The Southern grain louse 160-163. (Toxoptera graminurn Rondani). Some mis- Mizell, R. F., F. D. Bennett, and D. K. Reed. 2002. cellaneous results of the work of the division of Unsuccessful search for parasites of the crape- entomology. VI. U.S. Department of Agricul- myrtle aphid, Tinocallis kalza~valoukalalzi ture, Division of Entomology Bulletin 38: (Homoptera: Aphididae). Florida Entomolo- 7-19. gist 85(3): 521-523. Pepper, J. 0. 1965. A list of the Pennsylvania Morrison, W. P. and F. B. Peairs. 1998. Response Aphididae and their host plants (Homoptera). model concept and economic impact. In Transactions of the American Entomological Quisenberry, S. S. and F. B. Peairs, eds. Society 91: 181-231. Response model for an Introduced Pest- Quednau, F. W. 1966. A list of aphids from Quebec The Russian wheat aphid. Proceedings Thom- with descriptions of two new species (Homo- as Say Publications in Entomology, Entomo- ptera: Aphidoidea). Canadian Entomologist logical Society of America, Lanham, Mary- 98(4): 415430. land, 442 pp. . 2003. Atlas of the Drepanosphine aphids Miiller, F. P. 1969. Homoptera Aphidina. Unter- of the world. Part 11: Oestlund, ordnung Aphidina-Blattlause, Aphiden. Ex- 1923-Panaphidina Oestlund, 1923 (Hemi- kursionsfauna von Deutschland, Wirbellose 111 ptera: Aphididae: ). Memoirs of 2: 51-141. the American Entomological Institute 72: Oestlund, 0. W. 1886. List of the Aphididae of 1-301. Minnesota, with descriptions of some new Quisenberry, S. S. and F. B. Peairs. 1998. Res- species. Annual Report Geological and Natu- ponse model for an introduced pest-the ral History Survey of Minnesota 14: 17-56. Russian Wheat Aphid. Proceedings Thomas VOLUME 108. NUMBER 3 609

Say Publications in Entomology. Entomolog- Sciences Publications in Botany. No. 7(3). ical Society of America, Lanham, Maryland. National Museums of Canada, Ottawa. 442 pp. 1,115 pp. Race, S. R. 1970. New state records. USDA Smith, C. F. 1940. Notes on some Ohio aphids. Cooperative Economic Insect Report 20: 156. Ohio Journal of Science 40(3): 139-142. Rafinesque, C. S. 1818. Second memoir on the . 1974a. The genus Pl~loeorn~~suswith the genus Aphis, containing the descriptions of 24 description of P. dearborni n. sp. from PopuIus new American species. American Monthly trernuloides Michx. (Homoptera: Aphididae). Magazine 3: 15-18. Proceedings of the Entomological Society of Ragsdale, D. W., D. J. Voegtlin, and R. J. O'Neil. Washington 76: 66-72. 2004. Soybean aphid biology in North Amer- . 1974b. Keys to and descriptions of the ica. Annals of the Entomological Society of genera of Pemphigini in North America America 97(2): 204208. (Homoptera: Aphididae: Pemphiginae). North Remaudiere, G. and K. Naumann-Etienne. 1991. Carolina Agricultural Experiment Station Dtcouverte au Pakistan de 11h8te primaire de Tcchnical Bulletin No. 226, 61 pp. Rlzopalosiphunt maidis (Fitch) (Horn. Aphidi- . 1985. Peinpltiginae in North America. In dae). Comptes rendus de 1'Acadtmie d'Agri- S7elegiewicz, H. ed. Evolution and biosyste- culture de France 77: 61-62. matics of aphids. Proceedings of the Interna- Remaudiere, G. and M. Remaudiere. 1997. Cata- tional Aphidological Symposium at Jablonna, logue des Aphididae du monde (Homoptera 5-11 April, 1981. Polska Akademia Nauk Aphidoidea). INRA Editions, Paris. 473 pp. Instytut Zoologii, 510 pp. Riley, C. V. and J. Monell. 1879. Notes on the Smith, C. F. and H. A. Denmark. 1982. Trichosi- Aphididae of the United States, with de- phonaphis polygoni (van der Goot) (Homo- scriptions of species occurr~ng west of the ptera: Aphididae), a genus and species new to Mississippi. Part 11. Notes on , the United States. Florida Entomologist 65(3): with descriptions of new species. Bulletin 381-382. of the United States Geological and Geo- Smith, C. F. and G. F. Knowlton. 1975. Moss graphical Survey of the Territories 5(1): 1- aphids in the United States (Homoptera: 32. Aphididae). USDA Cooperative Economic Robinson, A. G. 1985. Annotated list of Uroleucon Insect Report 25(2 1): 42343 1. (Uroleucon, Uronzelun, Satulu) (Homoptera: Smith, C. F. and C. S. Parron. 1978. An annotated Aphididae) of America north of Mexico, with list of Aphididae (Homoptera) of North keys and descriptions of ncw species. Canadian America. North Carolina Agricultural Entomologist 117: 1029-1054. Experiment Station Technical Bulletin 255: Robinson, A. G. and G. A. Bradley. 1965. A 1-428. preliminary list of the aphids of Manitoba. Smith, E. F. 1890. The black peach aphis. A new Proceedings of the Entomological Society of species of the genus Aplzis. Entomologica Manitoba 21: 3945. Americana 6: 101-103. Russell, L. M. 1967. Notes on, and a key to, the Stekol'shtshikov, A. V. and G. K. Shaposhnikov. species of Neophyllaphis Takahashi (Homo- 1993. Revision of aphids of the genus Muscu- ptera: Aphididae). USDA Cooperative Eco- phis Borner (Homoptera: Aphididae). Ento- nomic Insect Report 17(11): 187-194. mologicheskoye Obozreniye 72(2): 333-344. . 1968. Aphis gossypii Glover 1877 (Insecta, (In Russian; English translation, 1994 Ento- Hemiptera-Homoptera): Request for valida- mological Review 73(1): 42-54). tion under the plenary powers. Z. N. (S.). S toetzel, M. B. 1985. Eucaiazzia eleguns (Ferrari), Bulletin of Zoological Nomenclature 25: an aphid new to the western hemisphere, with 116-1 19. archival data (Homoptera: Aphididae). Pro- Sailer, R. I. 1978. Our immigrant insect fauna. ceedings of the Entomological Society of Bulletin of the Entomological Society of Washington. 87(1): 4448. America 24: 3-1 1. . 1987. Information on and identification of . 1983. History of insect introductions, Diuiuplzis noxia (Homoptera: Aphididae) and pp. 15-38. Iiz Wilson, C. L. and C. L. Graham, other aphid species colonizing of wheat eds. Exotic Plant Pests and North American and barley in the United States. Journal of Agriculture. Academic Press, New York, Economic Entomology 80: 696-704. 522 pp. -. 1990. Some aphids of importance to Scoggan, H. J. 1978. The Flora of Canada. the southeastern United States (Homoptera: Part 3 Dicotylendonae (Saururaceae to Aphididae). Florida Entomologist 73(4): 580- Violaceae). National Museum of Natural 586. 610 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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