Catalog of the phylloxerids of the world (Hemiptera, Phylloxeridae)

Colin Favret; Roger Blackman; Gary Miller; Benjamin Victor

doi:10.3897/zookeys.629.10709

Abstract

A taxonomic and nomenclatural catalog of the phylloxerids (Hemiptera, Phylloxeridae) is presented. Six family-group names are listed, three being synonyms. Thirty-five genus-group names, of which six are subjectively valid, are presented with their type species, etymology, and grammatical gender. Ninety-four species-group names are listed, of which 73 are considered subjectively valid. This is the last group of Aphidomorpha to be catalogued, bringing the list of valid extant species to 5,218.

Keywords

Aphidomorpha , nomenclature, Phylloxera , Sternorrhyncha , taxonomy

Introduction

Phylloxeridae is a small family of Hemiptera, closely related to Adelgidae and Aphididae. Little is known of the biology of most of the family’s 69 species, although that of the economically important grape phylloxeran, Daktulosphaira vitifoliae (Fitch), has been studied in detail. Most species of phylloxerid feed on species of Juglandaceae or Fagaceae, with a large number forming galls on North American hickories (Carya spp.). Host alternation exists within the family (Stoetzel 1985) but it is either rare or understudied. Two fossil species are known, Palaeophylloxera seilacheri Heie and Peñalver 1999 and Acanthochermes longirostris Wegierek 2003, from the Miocene and Eocene, respectively.

Phylloxeridae is one of three extant families in the infraorder Aphidomorpha (Hemiptera, Sternorrhyncha) (Heie and Wegierek 2009). Whereas the Aphididae have been catalogued several times (Wilson and Vickery 1918, Eastop and Hille Ris Lambers 1976, Remaudière and Remaudière 1997) and the Adelgidae recently (Favret et al. 2015), the Phylloxeridae have not been comprehensively treated until now. Including the fossil taxa (Heie and Wegierek 2011), the entire infraorder has now been fully catalogued: 5,218 valid extant and 314 valid extinct species (Aphid.SpeciesFile.org).

In this catalog, we present six family-group, 35 genus-group, and 94 species-group names of extant phylloxerids. The family-group names include two valid subfamilies and two valid tribes and three subjective synonyms. The genus-group names include six valid names, 21 junior subjective synonyms, three junior objective synonyms, three junior homonyms, and two unavailable names. The species-group names include four subspecies (not including nominotypical subspecies), 14 subjective synonyms, one junior primary homonym, two nomina dubia, and four unavailable names.

The name Phylloxeridae in English is usually pronounced with the accent on the third syllable. However, the name of its type genus, Phylloxera, is often pronounced with an accent on the second. Because the e of xērós is an eta, the word made from it, once written in Roman letters and given Latin endings, must be considered to have a long e. The penultimate syllable of a Latin word must be accented when it contains such a long vowel and it is a fixed principle that the accentuation of Latin words is to be kept when they are borrowed into English. Therefore, strictly-speaking, only accentuation of the third syllable of Phylloxera is historically justified.

Russell (1975) described the complex history of the name of the grape phylloxeran, including the correct spelling of its generic name, Daktulosphaira Shimer 1866. Shimer also established Dactylosphaera (1867), probably meaning the latter to be an emended spelling of the former. The philological side of the alternate spellings can be stated briefly: k and c have both been used to transliterate classical Greek kappa, u and y to render upsilon, ai and ae the diphthong alpha+iota. C, y and ae were the preferred transliterations in classical Latin. K, u and ai are mostly used in linguistic circles that seek a more direct reflection of the phonetics of ancient Greek, bypassing the intermediary of Latin. Zoological nomenclature imposes Latin terminations, hence supposes Latinization of Greek (and other non-Latin) elements. Dactylosphaera is therefore the spelling more in the spirit of the system, although per ICZN Article 32.5.1 (International Commission on Zoological Nomenclature 1999), “incorrect transliteration or Latinization … are not to be considered inadvertent errors.” In addition to these two official spellings, other authors have used every possible combination of c/k, u/y and ae/ai to refer to the grape phylloxeran, giving Dactulosphaera (e.g., Kleeburg and Hummel 2001), Dactulosphaira (e.g., Fahrentrapp et al. 2015), Dactylosphaira (e.g., Alleweldt et al. 1991), Daktulosphaera (e.g., Loxdale 2008), Daktylosphaera (e.g., Tecchio et al. 2007), and Daktylosphaira (e.g., Torregrossa et al. 1997).

In any case, Shimer (1866, 1867) established different type species for Daktulosphaira and Dactylosphaera, thus the two spellings must be considered independent, available genus-group names (Wilson 1910). Dactylosphaera globosa Shimer 1867, one of a large number of North American hickory-feeding species, is the type species of its genus. As a consequence, Dactylosphaera has priority over all other generic names attributable to this distinct group. These include Xerophylla, described by Walsh later the same year (1867), Euphylloxera Del Guercio 1908, Notabilia Mordvilko 1909, Paramoritziella Grassi 1912, Parapergandea Börner 1930, Pergandea Börner 1908b, and Troitzkya Börner 1930. If we consider a key diagnostic character of the hickory-feeding species, the lack of abdominal spiracles, we can add Acanthaphis Del Guercio 1908 and Moritziella Börner 1908b to the list. It will require a thorough taxonomic revision of the phylloxerid family to correctly assign the various species, many of which are hardly known, to any of these listed generic names. Given this fact, the unfortunate history and spelling problems associated with Dactylosphaera and Daktulosphaira, and the fact that the identity and validity of the type species of Dactylosphaera may be questionable (Russell 1975), we have chosen to present a conservative classification, retaining the majority of species within the genus Phylloxera Boyer de Fonscolombe 1834. At some future date when more information is available, it may in particular be necessary to formalize a distinction between the Palearctic species (abdominal spiracular plates present) and the Nearctic species, species that typically form galls on hickories (abdominal spiracular plates absent, where known). As with the Catalog of Adelgidae (Favret et al. 2015), it is our hope that the present Catalog of Phylloxeridae will serve to stimulate interest and research on this insect group.

Also as with other recent catalogs of groups of Aphidomorpha, the etymology and grammatical gender of genus-group names has been included (Favret et al. 2008, 2009, 2015, Cortés Gabaudan et al. 2011, Nieto Nafría et al. 2011). Where original descriptions are listed with two page numbers, the first refers to a nomenclaturally valid diagnosis (e.g., a dichotomous key) and the second refers to the formal description. Valid names are listed in bold and synonyms preceded by ‘=’. The rank-specific endings of family-group synonyms are replaced by ‘–’. Species-group names are presented according to their current generic placement, their original generic placements in parentheses. An alphabetical index following the catalog provides the current placement of each name. Future updates will be published on Aphid Species File (Aphid.SpeciesFile.org).

Catalogue

PHYLLOXERIDAE Herrich-Schaeffer 1854

Subfamily PHYLLOXERINAEHerrich-Schaeffer 1854

Tribe ACANTHOCHERMESINIBörner 1913: 667

Original spelling. Acanthochermesini

Type genus. Acanthochermes Kollar 1848

ACANTHOCHERMESKollar 1848: 191

Type species. Acanthochermes quercusKollar 1848, by original monotypy

Etymology. Greek ákantha ‘thorn’ + Chermes [Hemiptera]

Gender. Masculine

quercusKollar 1848: 191 (Acanthochermes)

=balbianii (Lichtenstein 1874a: 782) (Phylloxera)

similiquercusJiang et al. 2009: 44,45 (Acanthochermes)

Tribe PHYLLOXERINIHerrich-Schaeffer 1854:VII

Original spelling. Phylloxeriden

Type genus. PhylloxeraBoyer de Fonscolombe 1834

=DACTYLOSPHAER– Shimer 1867: 2

Original spelling. Dactylosphaeridae

Type genus. DactylosphaeraShimer 1867

=MORITZIELL– Börner 1908b: 607

Original spelling. Moritziellini

Type genus. Moritziella Börner 1908b

=VACUN– Herrich-Schaeffer 1854:VII

Original spelling. Vacuniden

Type genus. Vacuna von Heyden 1837

APHANOSTIGMABörner 1909b: 61

Type species. Phylloxera piri Cholodkovsky 1904, by original monotypy

Etymology. Greek aphanḗs ‘invisible’ + -o + Greek stigma ‘spot’ [pterostigma]

Gender. Neuter

=CINACIUM Kishida 1924: 473

Type species. Cinacium iaksuienseKichida 1924, by original monotypy

Etymology. Japanese Kinako ‘soybean flour’ + -ium

Gender. Neuter

iaksuiense (Kishida 1924: 473) (Cinacium)

piri (Cholodkovsky 1904: 119) (Phylloxera)

DAKTULOSPHAIRAShimer 1866: 365

Type species. Pemphigus vitifoliae Fitch 1855, by original monotypy

Etymology. Greek dáktylos ‘finger’ + Greek sphaîra ‘ball’

Gender. Feminine

=PERITYMBIA Westwood 1869: 109

Type species. Peritymbia vitisana Westwood 1869, by original monotypy

Etymology. Greek perí ‘around’ + Greek týmbos ‘tomb’ [“tomb-like gall”]

Gender. Feminine

Note. Some references cite Westwood 1867: 6, but this is a note referencing an oral presentation that was never published (Westwood 1877:xlvii).

=RHIZAPHIS Planchon in Bazille et al. 1868: 336

Type species. Rhizaphis vastatrix Planchon 1868, by original monotypy

Etymology. Greek ríza ‘root’ + Aphis [Hemiptera: Aphididae]

Gender. Feminine

=RHIZOCERA Despeissis 1896: 14

Type species. None

Etymology. Greek ríza ‘root’ + Greek xērós ‘dry’ [“root drier” per Despeissis 1896, but note, Latin cēra ‘wax’]

Gender. Feminine

Note. Unavailable, not proposed as a valid name. Often misattributed to Kirk 1897: 8.

=VITEUS Shimer 1867: 6

Type species. Pemphigus vitifoliae Fitch 1855, by original monotypy

Etymology. Latin ‘of or pertaining to the vine’

Gender. Masculine

Note. Junior objective synonym of Daktulosphaira Shimer 1866

=XERAMPELUS Del Guercio 1900: 77,80

Type species. Rhizaphis vastatrix Planchon 1868, by original monotypy

Etymology. Greek xērós ‘dry’ + Greek ámpelos ‘vine’

Gender. Masculine

Note. Junior objective synonym of Rhizaphis Planchon 1868

vitifoliae (Fitch 1855: 862) (Pemphigus)

=pemphigoides (Donnadieu 1887: 1246) (Phylloxera)

=pervastatrix (Börner 1910: 4) (subspecies of Peritymbia vitifoliae (Fitch))

=vastatrix (Planchon in Bazille et al. 1868: 336) (Rhizaphis)

=vitisana (Westwood 1869: 109) (Peritymbia)

=vitis viniferae (Theobald 1914: 337) (Phylloxera) nomen nudum

=vulpinae (Börner 1952: 213) (subspecies of Viteus vitifoliae (Fitch))

FOAIELLABörner 1909b: 61

Type species. Phylloxera danesii Grassi and Foà 1907, inherited from replaced name

Etymology. (Anna) Foà [Italian entomologist] + -i + ella [diminutive suffix]

Gender. Feminine

Note. Replacement name for Boerneria Grassi and Foà 1908. Described as subgenus of Peritymbia Westwood 1869

=BOERNERIA Grassi and Foà 1908: 685

Type species. Phylloxera danesii Grassi and Foà 1907, by original monotypy

Etymology. (Carl) Börner [German entomologist] + -ia

Gender. Feminine

Note. Junior homonym of Boerneria Willem 1902: 4 (Collembola) and Boerneria Axelson 1902: 101 (Collembola). Replaced by Foaiella Börner 1909b

danesii (Grassi and Foà 1907: 431) (Phylloxera)

OLEGIAShaposhnikov 1979: 734

Type species. Aphanostigma ulmifoliae Aoki 1973, by original designation

Etymology. Oleg (Vasilyevich Kovalev) [Russian entomologist] + -ia

Gender. Feminine

ulmifoliae (Aoki 1973: 144) (Aphanostigma)

PHYLLOXERABoyer de Fonscolombe 1834: 222

Type species. Phylloxera quercus Boyer de Fonscolombe 1834, by original monotypy

Etymology. Greek phýllon ‘leaf’ + Greek xērós ‘dry’

Gender. Feminine

=ACANTHAPHIS Del Guercio 1908: 156,157

Type species. Phylloxera corticalis Kaltenbach 1867, by original designation

Etymology. Greek ákantha ‘thorn’ + Aphis [Hemiptera: Aphididae]

Gender. Feminine

Note. Junior objective synonym of Moritziella Börner 1908b

=DACTYLOSPHAERA Shimer 1867: 290

Type species. Dactylosphaera globosa Shimer 1867, by original monotypy

Etymology. Greek dáktylos ‘finger’ + Greek sphaîra ‘ball’

Gender. Feminine

=EUPHYLLOXERA Del Guercio 1908: 155,156

Type species. Phylloxera foveola Pergande 1904, by original designation

Etymology. Greek eû ‘truly’ + Phylloxera

Gender. Feminine

=HYSTRICHIELLA Börner 1908b: 609

Type species. Phylloxera spinulosa Targioni Tozzetti 1875, by original designation

Etymology. Greek hýstrix ‘porcupine’ + -i + -ella [diminutive suffix]

Gender. Feminine

Note. Described as subgenus of Phylloxera Boyer de Fonscolombe 1834

=MICRACANTHAPHIS Grassi in Grassi et al. 1912: 48

Type species. Vacuna coccinea von Heyden 1837, by original designation

Etymology. Greek mikrós ‘small’ + Acanthaphis

Gender. Feminine

=MORITZIELLA Börner 1908b: 608

Type species. Phylloxera corticalis Kaltenbach 1867, by original designation

Etymology. (Julius) Moritz [German entomologist] + -i + ella [diminutive suffix]

Gender. Feminine

=NOTABILIA Mordvilko 1909: 362

Type species. Phylloxera notabilis Pergande 1904, by original designation

Etymology. Latin notabilis ‘remarkable, sizeable’, inflected in the neuter plural

Gender. Neuter

=PARAMORITZIELLA Grassi in Grassi et al. 1912: 13

Type species. Phylloxera caryaefoliae Fitch 1856, by original designation

Etymology. Greek pará ‘beside’ + Moritziella

Gender. Feminine

=PARAPERGANDEA Börner 1930: 160

Type species. Phylloxera caryaevenae Fitch 1856, by original designation

Etymology. Greek pará ‘beside’ + Pergandea

Gender. Feminine

=PARAPHYLLOXERA Grassi in Grassi et al. 1912: 11,60

Type species. Vacuna glabra von Heyden 1837, by original designation

Etymology. Greek pará ‘beside’ + Phylloxera

Gender. Feminine

=PARTHENOPHYLLOXERA Grassi in Grassi et al. 1912: 11,62

Type species. Parthenophylloxera ilicisGrassi 1912, by original designation

Etymology. Greek parthénos ‘girl, virgin’ + Phylloxera

Gender. Feminine

=PERGANDEA Börner 1908b: 610

Type species. Dactylosphaera conica Shimer 1869, by original designation

Etymology. (Theodore) Pergande [American entomologist] + -a

Gender. Feminine

Note. Junior homonym of Pergandea Ashmead 1905: 382 (Hymenoptera). Described as subgenus of Dactylosphaera Shimer 1867

=PHYLLOXERELLA Grassi in Grassi et al. 1912: 11,54

Type species. Phylloxerella confusaGrassi 1912, by original designation

Etymology. Phylloxera + -ella [diminutive suffix]

Gender. Feminine

=PHYLLOXEROIDES Grassi in Grassi et al. 1912: 11,48

Type species. Phylloxera italicaGrassi 1912, by original designation

Etymology. Phylloxera + Greek –ō(i)dēs ‘resembling’

Gender. Masculine

=PSYLLOPTERA Ferrari 1872: 85

Type species. Psylloptera quercina Ferrari 1872, by original monotypy

Etymology. Psylla [Hemiptera: Psyllidae] + Greek pterá ‘wings’

Gender. Feminine

=RHANIS von Heyden 1837: 289

Type species. None

Etymology. Greek rhanís ‘drop (of a liquid)’

Gender. Feminine

Note. Unavailable, described in synonymy with Vacuna von Heyden 1837. Junior homonym of Rhanis Dejean 1836: 440 (Coleoptera)

=TROITZKYA Börner 1930: 160

Type species. Dactylosphaera caryaesemen Walsh 1867, by original designation

Etymology. (Nikolay Nikolaevich) Troitzky [Russian entomologist] + -a

Gender. Feminine

=VACUNA von Heyden 1837: 289

Type species. Vacuna coccinea von Heyden 1837, by original monotypy

Etymology. Latin Vacuna [minor goddess of ancient Italy]

Gender. Feminine

=XEROPHYLLA Walsh 1867: 283

Type species. Pemphigus caryaecaulis Fitch 1855, by subsequent designation (Börner 1930: 159)

Etymology. Greek xērós ‘dry’ + Greek phýllon ‘leaf’

Gender. Feminine

caryaeavellanaRiley 1880: 230 (Phylloxera)

caryaecaulis (Fitch 1855: 859) (Pemphigus)

=caryaemagna (Shimer 1869: 391) (Dactylosphaera)

caryaefallaxRiley 1874a: 1387 (Phylloxera)

caryaefoliaeFitch 1856: 446 (Phylloxera)

caryaeglobuliWalsh 1863: 309 (Phylloxera)

=hemisphericum (Shimer 1869: 387) (Dactylosphaera)

caryaegummosaRiley 1874a: 1387 (Phylloxera)

caryaepilula (Walsh 1867: 283) (Xerophylla) nomen nudum

caryaerenRiley 1874a: 1387 (Phylloxera) original spelling caryaereniformis but caryaeren in prevailing usage (ICZN Article 33.3.1)

caryaescissaRiley 1880: 230 (Phylloxera)

caryaesemen (Shimer 1869: 392) (Dactylosphaera) specific epithet first used by Walsh (1867: 283), but not placed in combination with a genus and hence unavailable until Shimer established it as a binomen

caryaesepta (Shimer 1869)

subspecies caryaesepta (Shimer 1869: 389) (Dactylosphaera)

subspecies perforansPergande 1904: 188,193 (variety of Phylloxera caryaesepta (Shimer 1869))

caryaevenae (Fitch 1856: 444) (Pemphigus)

castaneae (Haldeman 1850: 106) (Chermes)

castaneivora (Miyazaki 1968: 400) (Moritziella)

coccinea (von Heyden 1837: 289) (Vacuna)

=escorialensis Lichtenstein 1876: 130 (Phylloxera) nomen nudum

=globifera (von Heyden 1837: 289) (Rhanis) unavailable, described in synonymy with Vacuna coccinea von Heyden 1837

=rutila Dreyfus 1889: 95 (Phylloxera)

confusa Grassi in Grassi et al. 1912: 54 (Phylloxera)

conica (Shimer 1869: 390) (Dactylosphaera)

corticalisKaltenbach 1867: 44 (Phylloxera)

=iberica Staroselsky 1892: 177 (Phylloxera)

=lichtensteinii Balbiani 1874: 645 (Phylloxera)

davidsoniDuncan 1922: 271 (Phylloxera)

deplanataPergande 1904: 188,205 (Phylloxera)

depressa (Shimer 1869: 390) (Dactylosphaera)

devastatrixPergande 1904: 243,248 (Phylloxera)

foaeBörner 1909a: 26 (Phylloxera)

foveata (Shimer 1869: 393) (Dactylosphaera)

foveolaPergande 1904: 188,200 (Phylloxera)

fraxini Stebbins 1910: 46 (Phylloxera) nomen dubium, only the gall was described and it is probably not a phylloxerid

georgianaPergande 1904: 243,249 (Phylloxera)

glabra (von Heyden 1837: 291) (Vacuna)

=punctata Lichtenstein 1874b:CCI (Phylloxera) original name bipunctatum but punctata in prevailing usage (ICZN Article 33.3.1)

globosa (Shimer 1867)

subspecies coniferum (Shimer 1869: 397) (Dactylosphaera)

subspecies globosa (Shimer 1867: 2) (Dactylosphaera)

ilicis (Grassi in Grassi et al. 1912: 62) (Parthenophylloxera)

intermediaPergande 1904: 188,189 (Phylloxera)

italica (Grassi in Grassi et al. 1912: 48) (Phylloxeroides)

kunugiShinji 1943: 2 (Phylloxera)

minima (Shimer 1869: 391) (Dactylosphaera)

notabilisPergande 1904: 217,235 (Phylloxera)

perniciosaPergande 1904: 244,251 (Phylloxera)

pictaPergande 1904: 188,197 (Phylloxera)

pilosulaPergande 1904: 188,203 (Phylloxera)

quercetiPergande 1904: 263 (Phylloxera)

quercina (Ferrari 1872: 85) (Psylloptera)

=spinulosa Targioni Tozzetti 1875: 308 (Phylloxera)

quercusBoyer de Fonscolombe 1834: 223 (Phylloxera)

=florentina Targioni Tozzetti 1875: 287 (Phylloxera)

=scutifera Signoret 1867: 303 (Phylloxera) nomen dubium; Signoret (1867) wrote he was unable to find significant differences between this species and Phylloxera quercus Boyer de Fonscolombe except that scutifera was “slightly larger and darker”; he also drew a scale-like structure (Plate 7, Figure 6) that is not of phylloxerid origin, suggesting his description included a mixture of species

=signoreti Targioni Tozzetti 1875: 302 (Phylloxera)

reticulataDuncan 1922: 271 (Phylloxera)

rileyiRiley 1874b: 64 (Phylloxera)

rimosalisPergande 1904: 216,217 (Phylloxera)

russellaeStoetzel 1981: 128 (Phylloxera)

similansDuncan 1922: 272 (Phylloxera)

spiniferaPergande 1904: 261 (Phylloxera)

spinosa (Shimer 1869: 397) (Dactylosphaera)

spinuloidesPergande 1904: 243 (Phylloxera)

stanfordianaFerris 1919: 103 (Phylloxera)

stellataDuncan 1922: 269 (Phylloxera)

subelliptica (Shimer 1869: 389) (Dactylosphaera)

symmetrica Pergande 1904

subspecies purpureaPergande 1904: 232 (variety of Phylloxera symmetrica Pergande 1904)

subspecies symmetricaPergande 1904: 218,230 (Phylloxera)

subspecies vasculosaPergande 1904: 233 (variety of Phylloxera symmetrica Pergande 1904)

texanaStoetzel 1981: 141 (Phylloxera)

tuberculiferaDuncan 1922: 272 (Phylloxera)

Subfamily PHYLLOXERININAEBörner 1908b: 607

Original spelling. Phylloxerinini

Type genus. Phylloxerina Börner 1908a

PHYLLOXERINABörner 1908a: 94

Type species. Phylloxera salicis Lichtenstein 1884, by original monotypy

Etymology. Phylloxera + Latin -ina ‘in relation to’

Gender. Feminine

=GUERCIOJA Mordvilko 1909: 361

Type species. Chermes populi Del Guercio 1900, by original designation

Etymology. (Giacomo Del) Guercio [Italian entomologist] + -ja

Gender. Feminine

=LAUFFERELLA Lindinger 1933: 32

Type species. Chermes populi Del Guercio 1900, inherited from replaced name

Etymology. (Jorge) Lauffer [German entomologist] + -ella [diminutive suffix]

Gender. Feminine

Note. Replacement name for Pseudochermes Bonfigli 1909. Junior objective synonym of Guercioja Mordvilko 1909

=PSEUDOCHERMES Bonfigli 1909: 398

Type species. Chermes populi Del Guercio 1900, by original monotypy

Etymology. Greek pseudo- ‘untrue’ + Chermes [Hemiptera]

Gender. Masculine

Note. Junior homonym of Pseudochermes Nitsche in Judeich and Nitsche 1895: 1248 (Hemiptera: Cryptococcidae). Replaced by Lauferella Lindinger 1933

capreaeBörner 1942: 265 (Phylloxerina)

daphnoidisIglisch 1965: 424 (Phylloxerina)

moniliferae (Börner 1931: 696) (Guercioja) new name for Chermes populi Gillette 1914; possible synonym of Phylloxerina popularia (Pergande)

=populi (Gillette 1914: 269) (Chermes) junior primary homonym of Phylloxerina populi (Del Guercio 1900)

nyssae (Pergande 1904: 269) (Phylloxera)

popularia (Pergande 1904: 266) (Phylloxera)

populi (Del Guercio 1900: 81,83) (Chermes)

prolifera (Oestlund 1887: 16) (Phylloxera)

salicis (Lichtenstein 1884: 616) (Phylloxera)

salicola (Pergande 1904: 267) (Phylloxera)

Index of genus-group and species-group names

ACANTHAPHIS Del Guercio 1908 – synonym of Phylloxera

ACANTHOCHERMESKollar 1848 – Phylloxerinae, Acanthochermesini

APHANOSTIGMABörner 1909b – Phylloxerinae, Phylloxerini

balbianii Lichtenstein 1874a – synonym of Acanthochermes quercus

bipunctata Lichtenstein 1874b – see punctata

BOERNERIA Grassi and Foà 1908 – synonym of Foaiella

capreaeBörner 1942 – Phylloxerina

caryaeavellanaRiley 1880 – Phylloxera

caryaecaulisFitch 1855 – Phylloxera

caryaefallaxRiley 1874a – Phylloxera

caryaefoliaeFitch 1856 – Phylloxera

caryaeglobuliWalsh 1863 – Phylloxera

caryaegummosaRiley 1874a – Phylloxera

caryaemagnaShimer 1869 – synonym of Phylloxera caryaecaulis

caryaepilulaWalsh 1867 – Phylloxera

caryaerenRiley 1874a – Phylloxera

caryaereniformisRiley 1874a – see caryaeren

caryaescissaRiley 1880 – Phylloxera

caryaesemenShimer 1869 – Phylloxera

caryaeseptaShimer 1869 – Phylloxera

caryaevenaeFitch 1856 – Phylloxera

castaneaeHaldeman 1850 – Phylloxera

castaneivoraMiyazaki 1968 – Phylloxera

CINACIUM Kishida 1924 – synonym of Aphanostigma

coccineavon Heyden 1837 – Phylloxera

confusa Grassi in Grassi et al. 1912 – Phylloxera

conicaShimer 1869 – Phylloxera

coniferumShimer 1869 – subspecies of Phylloxera globosa

corticalisKaltenbach 1867 – Phylloxera

DACTYLOSPHAERA Shimer 1867 – synonym of Phylloxera

DAKTULOSPHAIRAShimer 1866 – Phylloxerinae, Phylloxerini

danesiiGrassi and Foà 1907 – Foaiella

daphnoidisIglisch 1965 – Phylloxerina

davidsoniDuncan 1922 – Phylloxera

deplanataPergande 1904 – Phylloxera

depressaShimer 1869 – Phylloxera

devastatrixPergande 1904 – Phylloxera

escorialensis Lichtenstein 1876 – synonym of Phylloxera coccinea

EUPHYLLOXERA Del Guercio 1908 – synonym of Phylloxera

florentina Targioni Tozzetti 1875 – synonym of Phylloxera quercus

foaeBörner 1909a – Phylloxera

FOAIELLABörner 1909b – Phylloxerinae, Phylloxerini

foveataShimer 1869 – Phylloxera

foveolaPergande 1904 – Phylloxera

fraxini Stebbins 1910 – Phylloxera

georgianaPergande 1904 – Phylloxera

glabravon Heyden 1837 – Phylloxera

globifera von Heyden 1837 – synonym of Phylloxera coccinea

globosaShimer 1867 – Phylloxera

GUERCIOJA Mordvilko 1909 – synonym of Phylloxerina

hemisphericum Shimer 1869 – synonym of Phylloxera caryaeglobuli

HYSTRICHIELLA Börner 1908b – synonym of Phylloxera

iaksuienseKishida 1924 – Aphanostigma

iberica Staroselsky 1892 – synonym of Phylloxera corticalis

ilicis Grassi in Grassi et al. 1912 – Phylloxera

intermediaPergande 1904 – Phylloxera

italica Grassi in Grassi et al. 1912 – Phylloxera

kunugiShinji 1943 – Phylloxera

LAUFFERELLA Lindinger 1933– synonym of Phylloxerina

lichtensteinii Balbiani 1874 – synonym of Phylloxera corticalis

MICRACANTHAPHIS Grassi in Grassi et al. 1912 – synonym of Phylloxera

minimaShimer 1869 – Phylloxera

moniliferaeBörner 1931 – Phylloxerina

MORITZIELLA Börner 1908b – synonym Phylloxera

NOTABILIA Mordvilko 1909 – synonym of Phylloxera

notabilisPergande 1904 – Phylloxera

nyssaePergande 1904 – Phylloxerina

OLEGIAShaposhnikov 1979 – Phylloxerinae, Phylloxerini

PARAMORITZIELLA Grassi in Grassi et al. 1912 – synonym of Phylloxera

PARAPERGANDEA Börner 1930 – synonym of Phylloxera

PARAPHYLLOXERA Grassi in Grassi et al. 1912 – synonym of Phylloxera

PARTHENOPHYLLOXERA Grassi in Grassi et al. 1912 – synonym of Phylloxera

pemphigoides Donnadieu 1887 – synonym of Daktulosphaira vitifoliae

perforansPergande 1904 – subspecies of Phylloxera caryaesepta

PERGANDEA Börner 1908b – synonym of Phylloxera

PERITYMBIA Westwood 1869 – synonym of Daktulosphaira

perniciosaPergande 1904 – Phylloxera

pervastatrixBörner 1910 – synonym of Daktulosphaira vitifoliae

PHYLLOXERABoyer de Fonscolombe 1834 – Phylloxerinae, Phylloxerini

PHYLLOXERELLA Grassi in Grassi et al. 1912 – synonym of Phylloxera

PHYLLOXERINABörner 1908a – Phylloxerininae

PHYLLOXEROIDES Grassi in Grassi et al. 1912 – synonym of Phylloxera

pictaPergande 1904 – Phylloxera

pilosulaPergande 1904 – Phylloxera

piriCholodkovsky 1904 – Aphanostigma

populariaPergande 1904 – Phylloxerina

populiDel Guercio 1900 – Phylloxerina

populi Gillette 1914 – synonym of Phylloxerina moniliferae

proliferaOestlund 1887 – Phylloxerina

PSEUDOCHERMES Bonfigli 1909 – synonym of Phylloxerina

PSYLLOPTERA Ferrari 1872 – synonym of Phylloxera

punctata Lichtenstein 1874b – synonym of Phylloxera glabra

purpureaPergande 1904 – subspecies of Phylloxera symmetrica

quercetiPergande 1904 – Phylloxera

quercinaFerrari 1872 – Phylloxera

quercusBoyer de Fonscolombe 1834 – Phylloxera

quercusKollar 1848 – Acanthochermes

reticulataDuncan 1922 – Phylloxera

RHANIS von Heyden 1837 – synonym of Phylloxera

RHIZAPHIS Planchon in Bazille et al. 1868 – synonym of Daktulosphaira

RHIZOCERA Despeissis 1896 – synonym of Daktulosphaira

rileyiRiley 1874b – Phylloxera

rimosalisPergande 1904 – Phylloxera

russellaeStoetzel 1981 – Phylloxera

rutila Dreyfus 1889 – synonym of Phylloxera coccinea

salicisLichtenstein 1884 – Phylloxerina

salicolaPergande 1904 – Phylloxerina

scutifera Signoret 1867 – synonym of Phylloxera quercus

signoreti Targioni Tozzetti 1875 – synonym of Phylloxera quercus

similansDuncan 1922 – Phylloxera

similiquercusJiang et al. 2009 – Acanthochermes

spiniferaPergande 1904 – Phylloxera

spinosaShimer 1869 – Phylloxera

spinuloidesPergande 1904 – Phylloxera

spinulosa Targioni Tozzetti 1875 –synonym of Phylloxera quercina

stanfordianaFerris 1919 – Phylloxera

stellataDuncan 1922 – Phylloxera

subellipticaShimer 1869 – Phylloxera

symmetricaPergande 1904 – Phylloxera

texanaStoetzel 1981 – Phylloxera

TROITZKYA Börner 1930 – synonym of Phylloxera

tuberculiferaDuncan 1922 – Phylloxera

ulmifoliaeAoki 1973 – Olegia

VACUNA von Heyden 1837 – synonym of Phylloxera

vasculosaPergande 1904 – subspecies of Phylloxera symmetrica

vastatrix Planchon in Bazille et al. 1868 – synonym of Daktulosphaira vitifoliae

VITEUS Shimer 1867 – synonym of Daktulosphaira

vitifoliaeFitch 1855 – Daktulosphaira

vitis viniferae Theobald 1914 – synonym of Daktulosphaira vitifoliae

vitisana Westwood 1869 – synonym of Daktulosphaira vitifoliae

vulpinae Börner 1952 – synonym of Daktulosphaira vitifoliae

XERAMPELUS Del Guercio 1900 – synonym of Daktulosphaira

XEROPHYLLA Walsh 1867 – synonym of Phylloxera

Acknowledgments

We thank Andrew Carmichael (USDA Systematic Entomology Laboratory) for literature research. Andrey V. Stekolshchikov (Russian Academy of Sciences) helped locate the Staroslesky (1892) reference, the most difficult to find. Masakazu Sano (Hokkaido Agricultural Research Center) helped research the etymology of Cinacium. We thank two external reviewers; Juan Manuel Nieto Nafría (Universidad de León, Spain) in particular provided meticulous editing and advice on nomenclatural issues. ICZN commissioner Patrice Bouchard (Canadian National Collection of Insects, Arachnids and Nematodes) also provided important nomenclatural advice. As with the adelgid catalog, we express a special appreciation for the organizations making available, in digital form, the vast and valuable historical literature. The Biodiversity Heritage Library in particular is an invaluable resource. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the USDA; USDA is an equal opportunity provider and employer.

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