PROC. ENTOMOL. SOC. WASH. 119(4), 2017, pp. 565–574

THE MEALY PLUM APHID AND ITS CONGENERS: A SYNONYMIC REVISION OF THE PRUNUS-INFESTING APHID GENUS HYALOPTERUS (: )

COLIN FAVRET,NARESH M. MESHRAM,GARY L. MILLER,JUAN MANUEL NIETO NAFRI´A, AND ANDREY V. S TEKOLSHCHIKOV

(CF) University of Montreal, Department of Biological Sciences, Biodiversity Centre, 4101 rue Sherbrooke est, Montreal QC, H1X 2B2 Canada. (e-mail: [email protected]) (ORCID: 0000-0001-6243-3184); (NMM) Indian Council of Agricultural Research, Indian Agricultural Research Institute, Division of Entomology, Pusa Campus, New Delhi – 110012, India. (e-mail: [email protected]); (GLM) USDA-ARS, Systematic Entomology Laboratory, 10300 Baltimore Ave, Bldg. 005, BARC-West, Beltsville, MD 20705, U.S.A. (e-mail: [email protected]); (JMNN) Universidad de Leo´n, Departamento de Biodiversidad y Gestio´nAmbiental, 24071 Leo´n, Spain. (e-mail: [email protected]) (ORCID: 0000-0001-6714-2415); (AVS) Russian Academy of Sciences, Zoological Institute, Universitetskaya nab. 1, Saint Petersburg 199034, Russia. (e-mail: [email protected])

Abstract.—The three species of Hyalopterus Koch cause economic damage to various stone fruit trees of the genus Prunus L., H. pruni (Geoffroy), H. amygdali (Blanchard), and H. persikonus Miller et al. Although the third species was estab- lished recently, it has been suggested that one of the twelve older synonyms of the first two may be applicable to the third. We undertook a synonymic revision of the nominal species of Hyalopterus to clarify the taxonomy and nomenclature of the genus. The three valid species of Hyalopterus are affirmed to be H. pruni (the mealy plum aphid), H. amygdali, and H. arundiniformis Ghulamullah, stat. nov. We determined that H. mimulus Bo¨rner and H. persikonus are junior synonyms of H. arundiniformis, syn. nov., and that Brachysiphum kobachidzei Rusanova is not a synonym of H. amygdali but a valid species of Aphis, stat. nov., comb. nov. Aphis amygdalipersicae Mosley is likely a senior synonym of Brachycaudus helichrysi (Kaltenbach), syn. nov.; to maintain current usage, we establish the former as a nomen oblitum with respect to the latter. Finally, several other nominal species were un- evaluable and are therefore listed as nomina dubia. Key Words: almond, apricot, nomenclature, peach, plum DOI: 10.4289/0013-8797.119.4.565

Species of the aphid genus Hyalopterus market values in the United States in 2014 (Koch 1854) are important pests of stone of 6.4 billion, and 53, 629.1, and 332.2 fruits of the genus Prunus L., including million dollars (Agricultural Marketing especially almond, apricot, peach, and Resource Center 2017). Hyalopterus plum (including prunes), with respective species are known to alternate between 566 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Prunus and Phragmites Adans. (Blackman MATERIALS AND METHODS and Eastop 2000). The genus name We evaluated the status and validity Hyalopterus was once used to refer to of the fifteen nominal species of Hy- anumberofaphidspecieswithshort alopterus listed by Remaudie`re and cornicles in the subfamily Aphidinae, Remaudie`re (1997) and Lozier et al. including several currently placed else- (2008). Study consisted primarily in where, e.g., Aphis hyperici Monell, reviewing the historical literature, es- Coloradoa abrotani (Koch), Hayhurstia pecially original descriptions, and the atriplicis (Linnaeus), Hyadaphis albus examination of the primary types of (Monzen), Longicaudus trirhodus nominal species, when available. Par- (Walker), and Semiaphis sphondylii ticular attention was paid to the possi- (Koch) (Favret 2017). Due to their ble synonyms of H. persikonus;type economically-important status, Hyalopterus specimens were identified using the keys species have attracted a lot of atten- to the species of Hyalopterus published tion and various authors have estab- by Lozier et al. (2008) and Rakauskas lished nominal species based on et al. (2013). Literature and type mate- relatively minor differences. Valid rial are listed with each nominal species, Hyalopterus species have thus accu- below. mulated many synonyms. As of the last published aphid catalog, there RESULTS were 12 names associated with two Aphis amygdali Blanchard 1840: 206 valid species, H. amygdali (Blanchard 1840), and H. pruni (Geoffroy 1762) Primary type information.—Host, Pru- (Remaudie`re and Remaudie`re 1997). nus persica (L.) Batsch; Locality, near However, a number of authors have Paris, France; Specimen(s), presumed lost provided morphological and molecu- or non-existent. lar evidence of a third species of Comments.—Despite its specific ep- Hyalopterus (Spampinato et al. 1988; ithet, this species was first described Mosco et al. 1997; Lozier et al. 2007, from peach with no explicit indication of 2008; Poulios et al. 2007), this third how it differed from the only other species having been given the name nominal species described at that time, H. persikonus Miller et al. in Lozier H. pruni. Boisduval (1867) associated et al. 2008. Although the evidence A. amygdali with almond (Prunus dulcis for the validity of three species of (Mill.) D.A. Webb (=P. amygdalus Hyalopterus is strong, previous stud- Batsch)). Bo¨rner (1952) treated H. pruni ies did not evaluate the possible val- and H. amygdali side by side, listing idity of the many synonyms and it does many synonyms of each, and no study not appear that the type material of has ever suggested these two should these nominal species was examined. themselves be considered synonyms. We Blackman and Eastop (2011) suggested thus treat H. amygdali as one of the three that an older name may have priority valid species of Hyalopterus. Our mod- over H. persikonus.Toestablishthe ern concept of the species has evolved validity of the three species in question and become refined over time, more by and assign them their correct names, we taxonomic tradition than by specific undertook a taxonomic and nomencla- reference to Blanchard’s (1840) de- tural study of the nominal species of scription. Several recent descriptions Hyalopterus. and diagnoses are available (Nieto VOLUME 119, NUMBER 4 567

Nafrı´aetal.2005,Lozieretal.2008, 1996; Remaudie`re and Remaudie`re 1997; Rakauskas et al. 2013). Ortego 1998; Halbert et al. 2000; Conclusion.—Valid as Hyalopterus Remaudie`re and Talhouk 2001; Noordam amygdali (Blanchard). 2004; Sorin and Arakawa 2005; Pe´rez Hidalgo et al. 2009; Favret et al. 2010; Adachi 2012; Buga and Stekolshchikov Aphis amygdalipersicae Mosley 2012; Piffaretti et al. 2012; 2013; 1841: 684 Zamora Mejı´as et al. 2012; Miller et al. Primary type information.—Host, 2016). Prunus persica (L.) Batsch; Locality, Conclusion.—Nomen dubium. Senior Britain; Specimen(s), presumed lost or synonym of Brachycaudus helichrysi non-existent. (Kaltenbach 1843), syn. nov. Nomen Comments.—Mosley (1841) described oblitum with respect to B. helichrysi; B. this species briefly: “body light green; helichrysi a nomen protectum with re- eyes deep crimson; antennae long; legs spect to A. amygdalipersicae. and tubercles, which are very short, of an amber color.” He did not mention the presence of wax, a significant oversight, Aphis arundinis Fabricius 1775: 734 and the cornicle color (tubercles) does Primary type information.—Host, not match that of Hyalopterus.We Calamagrostis epigeois (L.) Roth; Local- therefore suspect his species may not be ity, Europe; Specimen(s), non-existent. Hyalopterus at all, but more likely Comments.—The original morpho- asynonymofthepolyphagousspecies logical description of this nominal spe- Brachycaudus helichrysi (Kaltenbach cies is too brief to clearly place it in any 1843). The Mosley species has nomen- specific taxon. However, Fabricius clatural priority over that of Kaltenbach. (1775) named “Arundinis epigeois” as The less-than-certain synonymy and the host, today’s Calamagrostis epigeois the long-standing use of the name B. (L.) Roth. This plant species is not a host helichrysi argue against establishing of Hyalopterus, hence the placement of a straight-forward synonymy. We there- A. arundinis as a synonym of Diuraphis fore here consider A. amygdalipersicae calamagrostis (Ossiannilsson 1959) by a nomen dubium and furthermore Eastop and Hille Ris Lambers (1976). establish it, per ICZN Article 23.9 Kaltenbach (1843), Koch (1854), Buckton (International Commission on Zoolog- (1879), Del Guercio (1900), and Patch ical Nomenclature (ICZN) 1999), as (1914) essentially stated that A. arundinis a nomen oblitum with respect to B. is closely related to but distinct from helichrysi. Aphis amygdalipersicae has H. pruni,andTheobald(1927)went not been used as a valid name after one step further treating H. arundinis 1899, whereas B. helichrysi has been and H. pruni as synonyms. However, the used as valid at least 25 times by at Fabricius (1775) description places it out- least 10 authors in the last 50 years, side the scope of Hyalopterus.Giventhe over a span of more than 10 years confused history of the name, we do not (Smith and Cermeli 1979; Raychaudhuri attempt to place A. arundinis definitively, et al. 1980, 1981; Remaudie`re 1983; nor do we attempt to rectify its nomen- Harrington 1985; Sorin 1992; Verma clatural priority over D. calamagrostis. and Das 1992; Blackman and Eastop Conclusion.—Not Hyalopterus, nomen 1994, 2000, 2006; Aldryhim and Khalil dubium. 568 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Aphis gracilis Walker 1852: 1040 With clearing and remounting, the type specimens of Oestlund’s (1886) nomi- Primary type information.—Host, nal species may be assignable to one of Salix L.; Locality, Britain; Specimen, the species in North America, but in ei- dry alata, at the Natural History Mu- ther case A. phragmitidicola would be seum, London (Doncaster 1961). a junior synonym. Comments.—The lone alate specimen Conclusion.—Nomen dubium. of this nominal species was collected on willow, a spurious host association. Doncaster (1961) listed it as a synonym Aphis pruni Geoffroy 1762: 497 of H. pruni, although given that it is an Primary type information.—Host, alate, that its host association is unes- Prunus domestica L.; Locality, France; tablished, and that the dry specimen Specimen(s), presumed lost or non-existent. prevents adequate microscopic examina- Comments.—Although Geoffroy’s tion, we are unable to place it in any of (1762) work is rejected for the purposes the known species of Hyalopterus. of nomenclature (Hemming 1954), an Conclusion.—Nomen dubium. exception was made for the validation of (Geoffroy) (Hemming Aphis persicariae Hartig 1841: 370 1956). Geoffroy references Re´aumur Primary type information.—Host, (1737) in his description, who in turn Prunus armeniaca L., P. domestica L., cites the species as infesting the plum, and P. persica (L.) Batsch; Locality, Prunus domestica. Types are unavail- Germany; Specimen(s), presumed lost or able for this species but it is the most non-existent. well-established of the Hyalopterus Comments.—Hartig’s (1841) de- names and surely the mealy plum aphid scription is probably of a Hyalopterus, is the most studied. A fair amount of but it does not permit us to distinguish literature cites Fabricius (1775) as the which of the three species he was de- author of H. pruni (Favret 2017), but as scribing. In fact, he lists apricot, peach, Fabricius explicitly cites Geoffroy and plum as hosts, suggesting that he (1762), the Fabricius attribution is sim- was probably dealing with more than ply an error propagated in the literature. one species. Given the lack of primary We consider H. pruni one of the three type specimens, we are unable to eval- valid species of Hyalopterus, but it uate the validity of this name. should be noted that there is no way to Conclusion.—Nomen dubium. definitively validate the modern con- cept of the species with that of Geoffroy Aphis phragmitidicola Oestlund (1762) or Re´aumur (1737), who very 1886: 44 likely were referencing two or all three Primary type information.—Host, species in their publications. Our mod- Phragmites Adans.; Locality, near Min- ern taxon concept of H. pruni has de- neapolis, Minnesota, USA; Specimens, veloped over time and by taxonomic 10 on a single slide, at the University of tradition. Several recent descriptions and Minnesota (Cook 1982). diagnoses are available (Nieto Nafrı´a Comments.—Both H. pruni and H. et al. 2005, Lozier et al. 2008, Rakauskas amygdali are known from North Amer- et al. 2013). ica (Foottit et al. 2006), whereas the Conclusion.—Valid as Hyalopterus third species is not (Lozier et al. 2008). pruni (Geoffroy). VOLUME 119, NUMBER 4 569

Aphis prunicerasi Stewart 1802: 110 University collection where Matsumura’s types are deposited. Primary type information.—Host, Comments.—The lack of specimens Prunus avium L.; Locality, Europe; and the spurious host plant prevents us Specimen(s), non-existent. from properly evaluating this nominal Comments.—The name is presented species. without description. Conclusion.—Nomen dubium. Conclusion.—Nomen nudum. Hyalopterus arundiniformis Aphis spinarum Hartig 1841: 370 Ghulamullah 1942: 226 Primary type information.—Host, Primary type information.—Host, Prunus spinosa L.; Locality, Germany; Prunus armeniaca L.; Locality, Kabul, Specimen(s), presumed lost or non-existent. Afghanistan; Specimen, alata, at the Comments.—Prunus spinosa,blackthorn, Indian Agricultural Research Institute, is a species distributed over most of Eu- Pusa Campus, New Delhi. rope. Hyalopterus pruni is the only spe- Comments.—The holotype (alata) cies of the genus recorded from this host and a paratype (aptera) were examined. (Lozier et al. 2008, Holman 2009). The original description lists Prunus Conclusion.—Junior synonym of H. armeniaca, P. domestica, and P. persica pruni (Geoffroy). as hosts, but the holotype itself is la- beled with “apricot” (P. armeniaca). Brachysiphum kobachidzei Rusanova We identified the aptera using the di- 1941: 228 chotomous keys published by Lozier Primary type information.—Host, Ty- et al. (2008) and Rakauskas et al. (2013). pha angustifolia L.; Locality, Azerbaidjan; One couplet in the Rakauskas et al. Specimens, 3 apterae, one ovipara, (2013) key requires the measurement of one nymph, at the Zoological Insti- the width of the ultimate rostral segment, tute, Russian Academy of Sciences, a character unfortunately obscured in the St. Petersburg. aptera. Rostral morphology is generally Comments.—The type specimens, conserved across aphid morphs, so for with seventh abdominal marginal tuber- that measurement only, we used the cles below the level of the spiracles, are width of the ultimate rostral segment of those of an Aphis. In having much the alata. In both keys, the specimen shorter cornicles, however, they do not keyed out as H. persikonus. We thus match those of the other Typha-feeding consider it a synonym of this latter Aphis, A. typhae Mamontova. species and not of H. amygdali,as Conclusion.—Made valid and trans- previously recorded (Remaudie`re and ferred to Aphis, A. kobachidzei (Rusa- Remaudie`re 1997). nova) stat. nov., comb. nov. Conclusion.—Valid as Hyalopterus arundiniformis Ghulamullah, stat. nov. Hyalopterus abietinus Matsumura 1917: 358, 391 Hyalopterus mimulus Bo¨rner 1950: 6 Primary type information.—Host, Primary type information.—Host, Abies Mill.; Locality, Sapporo, Hokkaido, Prunus persica (L.) Batsch; Locality, Japan; Specimen(s), lost or non-existent, Germany; Specimens, apterae and alatae, confirmed absent from the Hokkaido at Bio-Test Labor, Groß Lu¨sewitz, Sanitz. 570 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Comments.—All six apterae syntypes 2017). With the addition here of Aphis were examined and identified using the amygdalipersicae, Brachycaudus heli- Lozier et al. (2008) and Rakauskas et al. chrysi has now accumulated 48 such (2013) keys. In all cases the specimens synonyms. As our methods for species keyed out as H. persikonus and not as H. diagnosis and delimitation improve, amygdali. the multiple synonyms of important Conclusion.—Transferred from being species, many with poor original de- junior synonym of H. amygdali (Blanchard) scriptions and missing type specimens, to be junior synonym of H. arundiniformis complicate the evaluation and descrip- Ghulamullah, syn. nov. tion of possible future new species. A lot of effort went into adequately es- Hyalopterus persikonus Miller, Lozier, tablishing the existence of a third spe- and Foottit in Lozier et al. 2008: 11 cies of Hyalopterus (Spampinato et al. 1988; Mosco et al. 1997; Lozier et al. Primary type information.—Host, 2007, 2008; Poulios et al. 2007) and sub- Prunus persica (L.) Batsch; Locality, sequently resurrecting, here, one out of Kala Nera, Thessaly, Greece; Specimens, twelve possible names. If the conclusions aptera, at US National Aphid Collection, of Piffaretti et al. (2012, 2013) are correct Beltsville, Maryland. and a sibling species of B. helichrysi is Comments.—The evidence is strong formally recognized, the work of estab- that this nominal species represents, lishing which of the 48 possible names it along with H. amygdali and H. pruni, the should bear, if any, will be challenging. third species of Hyalopterus (Spampinato We have surveyed and revised the nom- et al. 1988; Mosco et al. 1997; Lozier et al. inal species of the mealy plum aphid spe- 2007, 2008; Poulios et al. 2007). However, cies complex. We conclude that there are H. arundiniformis Ghulamullah 1942 and three species of Hyalopterus,eachcausing H. mimulus Bo¨rner 1950 also fit the de- varying degrees of economic damage to scription of this third species and both stone fruit trees of the genus Prunus.These have nomenclatural priority. Despite the are (1) the mealy plum aphid itself, new nomenclatural synonymy, Miller’s H. pruni,primarilycolonizingplum et al. description of the species (in Lozier trees; (2) H. amygdali, primarily colo- et al. 2008) is the most complete and nizing almond and apricot trees; (3) H. authoritative. arundiniformis,primarilycolonizingapri- Conclusion.—Junior synonym of cot and peach trees. It should be noted H. arundiniformis Ghulamullah, new that, although they do exhibit preferences, synonymy. all three of these species can be found on each other’s hosts. Care should be taken DISCUSSION to monitor regions of Prunus domestica- of agricultural and medical im- tion where the introduction of one or portance attract a lot of research attention another Hyalopterus species could have and, as a consequence, tend to have important economic repercussions (Lozier ahigher-than-averagetaxonomicandno- et al. 2008). menclatural complexity. Aphids are no Synonymy of Hyalopterus Koch 1854 exception, especially polyphagous species such as Aphis gossypii Glover or Myzus Nomina dubia persicae (Sulzer), with 41 and 38 sub- H. abietinus Matsumura 1917 jective synonyms, respectively (Favret H. gracilis (Walker 1852) VOLUME 119, NUMBER 4 571

H. persicariae (Hartig 1841) Blackman, R. L., and V. F. Eastop. 1994. Aphids H. phragmitidicola (Oestlund 1886) on the World’s Trees. Cab International, Nomina valida Wallingford, England. 986 pp. Blackman, R. L., and V. F. Eastop. 2000. Aphids H. amygdali (Blanchard 1840) on the World’s Crops, 2nd Edition. ed. Wiley, H. arundiniformis Ghulamullah 1942, Chichester. 414 pp. stat. nov. Blackman, R. L., and V. F. Eastop. 2006. Aphids on the World’s Herbaceous Plants and Shrubs. =H. mimulus Bo¨rner 1950, syn. nov. John Wiley & Sons, Chichester, England. =H. persikonus Miller et al. in Lozier 1493 pp. et al. 2008, syn. nov. Blackman, R. L., and V. F. Eastop. 2011. Addi- tions and amendments to “Aphids on the H. pruni (Geoffroy 1762) World’s Plants.” Zootaxa 2774: 57–68.  H. spinarum (Hartig 1841) Blanchard, E. 1840. Histoire Naturelle des In- = sectes. Orthopte`res, Ne´vropte`res, He´mipte`res, Hyme´nopte`res, Le´pidopte`res et Dipte`res. P. Dume´nil, Paris. 672 pp. ACKNOWLEDGMENTS Boisduval, J. A. 1867. Essai sur l’entomologie horticole : comprenant l’histoire des insectes We express our appreciation to Thomas nuisibles a` l’horticulture avec l’indication des Thieme (BTL Bio-Test Labor, Stantz, moyens propres a` les e´loigner ou a` les de´- Germany) for the loan of the Bo¨rner type truire et l’histoire des insectes et autres ani- material; to Robin Elizabeth Thomson maux utiles aux cultures. E. Donaud, Paris. (University of Minnesota, St. Paul, USA) 648 pp. for providing a photograph of the Oestlund Bo¨rner, C. 1950. Neue europa¨ische Blattlausarten. Self-published, Naumberg (Saale). 19 pp. type material; to Masakazu Sano (Hokkaido Bo¨rner, C. 1952. Europae centralis Aphides. Die University, Sapporo, Japan) who con- Blattla¨use Mitteleuropas. Namen, Synonyme, firmed the absence of the Matsumura Wirtspflanzen, Generationszyklen. Mittei- types at the Hokkaido University col- lungen der Thu¨ringischen Botanischen Ge- lection; to Roger Blackman (Natural sesellschaft 3: 1–259. History Museum, London) who confirmed Buckton, G. B. 1879. Monograph of British Aphides, Volume 2. Ray Society, London. the absence of the Mosley type but the 176 pp. presence of the Walker type at the NHM. Buga, S., and A. V. Stekolshchikov. 2012. Aphids Roger Blackman and Rimantas Rakauskas of the tribe Macrosiphini (Insecta: Homo- (Vilnius University, Lithuania) are grate- ptera: Aphididae) in Belarus. Zoosystematica fully acknowledged for providing help- Rossica 21: 63–96. ful reviews that improved the quality and Cook, E. F. 1982. Types of the Aphididae in the University of Minnesota collection, with thoroughness of the manuscript. USDA special reference to the species described by is an equal opportunity provider and O.W. Oestlund and H.F. Wilson. Annals of employer. the Entomological Society of America 75: 71–77. Literature Cited Del Guercio, G. 1900. Prospetto dell’afidofauna italica. Nuove Relazioni Intorno ai Lavori Adachi, S. 2012. Aphids of Ehime Prefecture, della Regia Stazione di Entomologia Agraria Shikoku, Japan (Hemiptera, Aphididae). di Firenze, Serie Prima 2: 1–236. Bulletin of Ehime Prefectural Science Museum Doncaster, J. P. 1961. Francis Walker’s Aphids. 17: 29–47. William Clowes and Sons, London. 165 pp. Agricultural Marketing Resource Center. 2017. Eastop, V. F. and D. Hille Ris Lambers. 1976. AgMRC. (http://www.agmrc.org/). Survey of the World’s Aphids. Dr. W. Junk, Aldryhim, Y., and A. F. Khalil. 1996. The Aphi- The Hague. 573 pp. didae of Saudi Arabia. Fauna of Saudi Arabia Fabricius, J. C. 1775. Systema Entomologiae, 15: 161–195. sistens insectorum classes, ordines, genera, 572 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

species, adiectis synonymis, locis, descrip- Media B.V., Dordrecht, The Netherlands. tionibus, observationibus. Libraria Kortii, 1216 pp. Flensburg and Leipzig. 816 pp. International Commission on Zoological No- Favret, C. 2017. Aphid Species File. (http:// menclature (ICZN). 1999. International Code Aphid.SpeciesFile.org). of Zoological Nomenclature, Fourth Edition. Favret, C., J. J. Duggan, N. J. Sanders, and L. R. International Trust for Zoological Nomen- Phillippe. 2010. Actual and inferred checklist clature, London. 306 pp. of the aphids (Hemiptera: Aphididae) of the Kaltenbach, J. H. 1843. Monographie der Familien Great Smoky Mountains National Park, with der Pflanzenla¨use (Phytophthires). P. Fagot, attendant ant and host plant associations. Aachen. 222 pp. Proceedings of the Entomological Society of Koch, C. L. 1854. Die Pflanzenla¨use Aphiden Washington 112: 381–403. getreu nach dem Leben abgebildet und Foottit, R. G., S. E. Halbert, G. L. Miller, E. Maw, beschrieben, Volume 1. von J.L. Lotzbeck, and L. M. Russell. 2006. Adventive aphids Nu¨rnberg. 36 pp. (Hemiptera: Aphididae) of America north of Lozier, J. D., R. G. Foottit, G. L. Miller, N. J. Mexico. Proceedings of the Entomological Mills, and G. K. Roderick. 2008. Molecular Society of Washington 108: 583–610. and morphological evaluation of the aphid Geoffroy, E. L. 1762. Histoire abre´ge´e des In- genus Hyalopterus Koch (Insecta: Hemi- sectes qui se trouvent aux environs de Paris. ptera: Aphididae), with a description of a new Durand, Paris. 523 pp. species. Zootaxa 1688: 1–19. Ghulamullah, C. 1942. Aphididae and some other Lozier, J. D., G. K. Roderick, and N. J. Mills. Rhynchota from Afghanistan. Indian Journal 2007. Genetic evidence from mitochon- of Entomology 3(1941): 225–243. drial, nuclear, and endosymbiont markers Halbert, S. E., G. Remaudie`re, and S. E. Webb. for the evolution of host plant associated 2000. Newly established and rarely collected species in the aphid genus Hyalopterus aphids (Homoptera: Aphididae) in Florida (Hemiptera: Aphididae). Evolution 61: 1353– and the southeastern United States. Florida 1367. Entomologist 83: 79–91. Matsumura, S. 1917. A list of the Aphididae of Harrington, R. 1985. A comparison of the Japan, with description of new species and external morphology of “scent plaques” genera. Journal of the College of Agriculture, on the hind tibiae of oviparous aphids Tohoku Imperial University 7: 351–414. (Homoptera: Aphididae). Systematic En- Miller, G. L., A. S. Jensen, C. Favret, M. A. Metz, tomology 10: 135–144. and R. R. Parmenter. 2016. The first report Hartig, T. 1841. Versuch einer Eintheilung der of the aphids (Hemiptera: : Pflanzenla¨use (Phytophthires Burm.) nach der Aphididae) of the Valles Caldera National Flu¨gelbildung. Zeitschrift fu¨r die Entomologie, Preserve, New Mexico, USA. Proceedings herausgegeben von Ernst Friedrich Germar 3: of the Entomological Society of Washington 359–376. 118: 289–296. Hemming, F. (ed.). 1954. Opinion 228. Rejection Mosco, M. C., P. Arduino, L. Bullini, and S. for nomenclatorial purposes of Geoffroy, Barbagallo. 1997. Genetic heterogeneity, re- 1762, “Histoire abre´ge´e des insectes qui se productive isolation and host preferences in trouvent aux environs de Paris.” Opinions and mealy aphids of the Hyalopterus pruni com- Declarations Rendered by the International plex (Homoptera, Aphidoidea). Molecular Commission on Zoological Nomenclature 4: Ecology 6: 667–670. 209–220. Mosley, O. 1841. Aphides (continued from Hemming, F. (ed.). 1956. Validation under p.628). The Gardeners’ Chronicle [1841] the Plenary Powers of the specific name (42): 684. pruni Geoffroy, 1762, as published in the Nieto Nafrı´a, J. M., M. P. Mier Durante, F. Garcı´a combination Aphis pruni (Class Insecta, Prieto, and N. Pe´rez Hidalgo. 2005. Hemi- Order Hemiptera). Opinions and Declara- ptera Aphididae III. In Ramos Sa´nchez, A. tions Rendered by the International Com- (ed.), Fauna Ibe´rica. Museo Nacional de mission on Zoological Nomenclature 12: Ciencias Naturales, Madrid. 362 pp. 363–376. Noordam, D. 2004. Aphids of Java. Part VI: Holman, J. 2009. Host Plant Catalog of Aphids, sixty species, six of which are newly de- Palearctic Region. Springer Science+Business scribed (Homoptera: Aphididae: Aphidinae, VOLUME 119, NUMBER 4 573

Lachninae, Neophyllaphidinae, Pemphiginae). aphids (Homoptera: Aphididae) from South Zoologische Verhandelingen 346: 85–212. India, I. Insecta Matsumurana, New Series Oestlund, O. W. 1886. List of the Aphididae of 23: 1–20. Minnesota with descriptions of some new Raychaudhuri, D. N., L. K. Ghosh, and S. K. Das. species. Annual Report of the Minnesota 1980. Studies on the aphids (Homoptera: State Geological and Natural History Survey Aphididae) from north and north-west India, 14: 17–56. 1. Insecta Matsumurana, New Series 20: 1–42. Ortego, J. 1998. Pulgones de la Patagonia Ar- Re´aumur, R. A. F. 1737. Memoires pour Servir a` gentina con la descripcio´n de Aphis intrusa l’Histoire des Insectes. L’Imprimerie Royale, sp. n. (Homoptera: Aphididae). Revista de la Paris. 532 pp. Facultad de Agronomı´a, La Plata 102(1997): Remaudie`re, G. 1983. Contribution a` la con- 59–80. naissance des aphides (Homoptera, Aphi- Ossiannilsson, F. 1959. Contributions to the doidea) de la Gre`ce et description d’un knowledge of Swedish aphids I. Descriptions of Thelaxes nouveau. Annales de l’Institut some apparently undescribed forms. Kunglig phytopathologique Benaki, Nouvelle Se´rie Lantbruksho¨gskolans Annaler 25: 1–46. 13(1982): 99–119. Patch, E. M. 1914. Maine aphids of the rose Remaudie`re, G., and M. Remaudie`re. 1997. family. Bulletin of the Maine Agicultural Catalogue of the World’s Aphididae. INRA Experiment Station 255: 253–280. Editions, Paris. 473 pp. Passerini, G. 1860. Gli afidi con un prospetto dei Remaudie`re, G., and A. S. Talhouk. 2001. Les generi ed alcune specie nuove Italiane. Ti- aphides du Liban et de la Syrie avec la de- pografia Carmignani, Parma. 40 pp. scription d’une nouvelle espe`ce de genre Pe´rez Hidalgo, N., A. Umaran, M. P. Mier Durante, Brachyunguis Das (Hemiptera, Aphididae). and J. M. Nieto Nafrı´a. 2009. Aportaciones Parasitica 55(1999): 149–183. alaafidofaunaı´bero-balear (Hemiptera, Rusanova, V. N. 1941. New aphid species Bra- Aphididae) a partir de los fotografı´as chysiphum kobachidzei, sp. n. (fam. Aphidi- (y de su metadatos) depositadas en el dae) from Georgia. Acade´mie des Sciences “Banco taxono´mico faunı´stico digital de de l’URSS, Filiale Ge´orgienne - Section de los invertebrados ibe´ricos” (B.T.F.D.I.I.). Zoologie [1941]: 227–228. Graellsia 65: 171–181. Smith, C. F., and M. Cermeli. 1979. An annotated Piffaretti, J., J.-P. Rossi, F. Vanlerberghe-Masutti, list of Aphididae (Homoptera) of the Carib- G. Genson, A. Cœur d’acier, and E. Jousselin. bean Islands and South and Central America. 2013. Molecular identification and ecological North Carolina Agricultural Experiment characteristics of two cryptic lineages within Station Technical Bulletin 259: 1–131. a cosmopolitan aphid pest, Brachycaudus Sorin, M. 1992. A list of Aphididae of Niigata helichrysi (Hemiptera: Aphididae). Applied Prefecture, Japan. Transactions of the Essa Entomology and Zoology 48: 155–164. Entomological Society of Niigata 74: 1–7. Piffaretti, J., F. Vanlerberghe-Masutti, A. Tayeh, A.-L. Sorin, M., and A. Arakawa. 2005. A list of Clamens, A. Cœur d’acier, and E. Jousselin. Aphididae of Niigata Prefecture, Japan (3). 2012. Molecular phylogeny reveals the exis- Transactions of the Essa Entomological So- tence of two sibling species in the aphid pest ciety of Niigata 92: 1–21. Brachycaudus helichrysi (Hemiptera: Aphididae). Spampinato, R., P. Arduino, S. Barbagallo, and L. Zoologica Scripta 41: 266–280. Bullini. 1988. Analisi genetica di Hyalopte- Poulios, K. D., J. T. Margaritopoulos, and J. A. rus pruni e H. amygdali, e dimostrazione Tsitsipis. 2007. Morphological separation of dell’esistenza di una nuova specie, pp. 261– host adapted taxa within the Hyalopterus 265. In Atti XV Congresso Nazionale Ital- pruni complex (Hemiptera: Aphididae). Eu- iano Di Entomologia. L’Aquila. ropean Journal of Entomology 104: 235. Stewart, C. 1802. Elements of Natural History; Rakauskas, R., J. Havelka, and A. Zaremba. Being an Introduction to the Systema Naturae 2013. Mitochondrial COI and morphological of Linnaeus; Comprising the Characters of specificity of the mealy aphids (Hyalopterus the Whole Genera, and Most Remarkable ssp.) collected from different hosts in Europe Species; Particularly of All those That Are (Hemiptera, Aphididae). ZooKeys 319: 255–267. Natives of Britain, with the Principal Cir- Raychaudhuri, D. N., D. Ghosh, D. Raychaudhuri, cumstances of their History and Manners. T. and B. K. Agarwala. 1981. Studies on the Cadell Jr. and W. Davies, London. 491 pp. 574 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Theobald, F. V. 1927. The Plant Lice or Aphidi- Walker, F. 1852. List of the Specimens of Ho- dae of Great Britain, Volume 2. Headley mopterous Insects in the Collection of the Brothers Invicta Press, London. 411 pp. British Museum 4: 909–1118. Verma, K. D., and S. M. Das. 1992. The Aphi- Zamora Mejı´as, D., N. Pe´rez Hidalgo, W. didae of North West India (With Special Villalobos Muller, andP.Hanson.2012. Reference to Aphids of Jammu and Kashmir New data about the Costa Rican aphid State). S.B. Nangia for Ashish Publishing fauna (Hemiptera, Aphididae). Graellsia 68: House, New Delhi. 171 pp. 305–312.