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

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.

Introduction

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

is one of three extant families in the infraorder (, ) (Heie and Wegierek 2009). Whereas the have been catalogued several times (Wilson and Vickery 1918, Eastop and Hille Ris Lambers 1976, Remaudière and Remaudière 1997) and the recently (Favret et al. 2015), the 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 in English is usually pronounced with the accent on the third syllable. However, the name of its type genus, , 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 is historically justified.

Russell (1975) described the complex history of the name of the grape phylloxeran, including the correct spelling of its generic name, Shimer 1866. Shimer also established (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. is therefore the PageBreakspelling 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 (e.g., Kleeburg and Hummel 2001), (e.g., Fahrentrapp et al. 2015), (e.g., Alleweldt et al. 1991), (e.g., Loxdale 2008), (e.g., Tecchio et al. 2007), and (e.g., Torregrossa et al. 1997).

In any case, Shimer (1866, 1867) established different type species for and , thus the two spellings must be considered independent, available genus-group names (Wilson 1910). Shimer 1867, one of a large number of North American hickory-feeding species, is the type species of its genus. As a consequence, has priority over all other generic names attributable to this distinct group. These include , described by Walsh later the same year (1867), Del Guercio 1908, Mordvilko 1909, Grassi 1912, Börner 1930, Börner 1908b, and Börner 1930. If we consider a key diagnostic character of the hickory-feeding species, the lack of abdominal spiracles, we can add Del Guercio 1908 and 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 and , and the fact that the identity and validity of the type species of may be questionable (Russell 1975), we have chosen to present a conservative classification, retaining the majority of species within the genus 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 (Favret et al. 2015), it is our hope that the present Catalog of will serve to stimulate interest and research on this insect group.

Also as with other recent catalogs of groups of , 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

Subfamily

Tribe

Original spelling.

Type genus.

Type species.

Etymology. Greek ákantha ‘thorn’ +

Gender. Masculine

=

Tribe

Original spelling. Phylloxeriden

Type genus.

=DACTYLOSPHAER–

Original spelling.

Type genus.

=MORITZIELL–

Original spelling.

Type genus.

=VACUN–

Original spelling. Vacuniden

Type genus.

Type species.

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

Gender. Neuter

=

Type species.

Etymology. Japanese Kinako ‘soybean flour’ + -ium

Gender. Neuter

Type species.

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

Gender. Feminine

=

Type species.

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

Gender. Feminine

Note. Some references cite

=

Type species.

Etymology. Greek ríza ‘root’ +

Gender. Feminine

=

Type species. None

Etymology. Greek ríza ‘root’ + Greek xērós ‘dry’ [“root drier” per

Gender. Feminine

Note. Unavailable, not proposed as a valid name. Often misattributed to

=

Type species.

Etymology. Latin ‘of or pertaining to the vine’

Gender. Masculine

Note. Junior objective synonym of

=

Type species.

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

Gender. Masculine

Note. Junior objective synonym of

=

=

=

=

=

=

Type species.

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

Gender. Feminine

Note. Replacement name for

=

Type species.

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

Gender. Feminine

Note. Junior homonym of

Type species.

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

Gender. Feminine

Type species.

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

Gender. Feminine

=

Type species.

Etymology. Greek ákantha ‘thorn’ +

Gender. Feminine

Note. Junior objective synonym of

=

Type species.

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

Gender. Feminine

=

Type species.

Etymology. Greek eû ‘truly’ +

Gender. Feminine

=

Type species.

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

Gender. Feminine

Note. Described as subgenus of

=

Type species.

Etymology. Greek mikrós ‘small’ +

Gender. Feminine

=

Type species.

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

Gender. Feminine

=

Type species.

Etymology. Latin

Gender. Neuter

=

Type species.

Etymology. Greek pará ‘beside’ +

Gender. Feminine

=

Type species.

Etymology. Greek pará ‘beside’ +

Gender. Feminine

=

Type species.

Etymology. Greek pará ‘beside’ +

Gender. Feminine

=

Type species.

Etymology. Greek parthénos ‘girl, virgin’ +

Gender. Feminine

=

Type species.

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

Gender. Feminine

Note. Junior homonym of

=

Type species.

Etymology.

Gender. Feminine

=

Type species.

Etymology.

Gender. Masculine

=

Type species.

Etymology.

Gender. Feminine

=

Type species. None

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

Gender. Feminine

Note. Unavailable, described in synonymy with

=

Type species.

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

Gender. Feminine

=

Type species.

Etymology. Latin

Gender. Feminine

=

Type species.

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

Gender. Feminine

=

=

subspecies

subspecies

=

=

=

=

=

=

subspecies

subspecies

=

=

=

=

subspecies

subspecies

subspecies

Subfamily

Original spelling.

Type genus.

Type species.

Etymology.

Gender. Feminine

=

Type species.

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

Gender. Feminine

=

Type species.

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

Gender. Feminine

Note. Replacement name for

=

Type species.

Etymology. Greek pseudo- ‘untrue’ +

Gender. Masculine

Note. Junior homonym of

=

Index of genus-group and species-group names

Del Guercio 1908 – synonym of

Kollar 1848 – ,

Börner 1909b – ,

Lichtenstein 1874a – synonym of

Lichtenstein 1874b – see

Grassi and Foà 1908 – synonym of

Börner 1942 –

Riley 1880 –

Fitch 1855 –

Riley 1874a –

Fitch 1856 –

Walsh 1863 –

Riley 1874a –

Shimer 1869 – synonym of

Walsh 1867 –

Riley 1874a –

Riley 1874a – see

Riley 1880 –

Shimer 1869 –

Shimer 1869 –

Fitch 1856 –

Haldeman 1850 –

Miyazaki 1968 –

Kishida 1924 – synonym of

von Heyden 1837 –

Grassi in Grassi et al. 1912 –

Shimer 1869 –

Shimer 1869 – subspecies of

Kaltenbach 1867 –

Shimer 1867 – synonym of

Shimer 1866 – ,

Grassi and Foà 1907 –

Iglisch 1965 –

Duncan 1922 –

Pergande 1904 –

Shimer 1869 –

Pergande 1904 –

Lichtenstein 1876 – synonym of

Del Guercio 1908 – synonym of

Targioni Tozzetti 1875 – synonym of

Börner 1909a –

Börner 1909b – ,

Shimer 1869 –

Pergande 1904 –

Stebbins 1910 –

Pergande 1904 –

von Heyden 1837 –

von Heyden 1837 – synonym of

Shimer 1867 –

Mordvilko 1909 – synonym of

Shimer 1869 – synonym of

Börner 1908b – synonym of

Kishida 1924 –

Staroselsky 1892 – synonym of

Grassi in Grassi et al. 1912 –

Pergande 1904 –

Grassi in Grassi et al. 1912 –

Shinji 1943 –

Lindinger 1933– synonym of

Balbiani 1874 – synonym of

Grassi in Grassi et al. 1912 – synonym of

Shimer 1869 –

Börner 1931 –

Börner 1908b – synonym

Mordvilko 1909 – synonym of

Pergande 1904 –

Pergande 1904 –

Shaposhnikov 1979 – ,

Grassi in Grassi et al. 1912 – synonym of

Börner 1930 – synonym of

Grassi in Grassi et al. 1912 – synonym of

Grassi in Grassi et al. 1912 – synonym of

Donnadieu 1887 – synonym of

Pergande 1904 – subspecies of

Börner 1908b – synonym of

Westwood 1869 – synonym of

Pergande 1904 –

Börner 1910 – synonym of

Boyer de Fonscolombe 1834 – ,

Grassi in Grassi et al. 1912 – synonym of

Börner 1908a –

Grassi in Grassi et al. 1912 – synonym of

Pergande 1904 –

Pergande 1904 –

Cholodkovsky 1904 –

Pergande 1904 –

Del Guercio 1900 –

Gillette 1914 – synonym of

Oestlund 1887 –

Bonfigli 1909 – synonym of

Ferrari 1872 – synonym of

Lichtenstein 1874b – synonym of

Pergande 1904 – subspecies of

Pergande 1904 –

Ferrari 1872 –

Boyer de Fonscolombe 1834 –

Kollar 1848 –

Duncan 1922 –

von Heyden 1837 – synonym of

Planchon in Bazille et al. 1868 – synonym of

Despeissis 1896 – synonym of

Riley 1874b –

Pergande 1904 –

Stoetzel 1981 –

Dreyfus 1889 – synonym of

Lichtenstein 1884 –

Pergande 1904 –

Signoret 1867 – synonym of

Targioni Tozzetti 1875 – synonym of

Duncan 1922 –

Jiang et al. 2009 –

Pergande 1904 –

Shimer 1869 –

Pergande 1904 –

Targioni Tozzetti 1875 –synonym of

Ferris 1919 –

Duncan 1922 –

Shimer 1869 –

Pergande 1904 –

Stoetzel 1981 –

Börner 1930 – synonym of

Duncan 1922 –

Aoki 1973 –

von Heyden 1837 – synonym of

Pergande 1904 – subspecies of

Planchon in Bazille et al. 1868 – synonym of

Shimer 1867 – synonym of

Fitch 1855 –

Theobald 1914 – synonym of

Westwood 1869 – synonym of

Börner 1952 – synonym of

Del Guercio 1900 – synonym of

Walsh 1867 – synonym of