A taxonomic and nomenclatural Catalogue of the adelgids (Hemiptera: Adelgidae) is presented. Six family-group names are listed, five being synonyms of Adelgidae. Twenty-two genus-group names, of which nine are subjectively valid and in use, are presented with their type species, etymology, and grammatical gender. One hundred and six species-group names are listed, of which 70 are considered subjectively valid.

Aphidomorpha , nomenclature, Sternorrhyncha , taxonomy, woolly adelgid

Adelgidae is a small family of Hemiptera with 65 species, closely related to Aphididae. They exhibit a two-year life cycle, with some species alternating hosts between spruce (Picea) one year and species of another conifer genus (Abies, Larix, Pinus, Pseudotsuga, Tsuga) the next. Other species or populations do not alternate hosts, feeding only on Picea or one of the other conifer genera. Like other Aphidomorpha, Adelgidae exhibit cyclical parthenogenesis, although they are oviparous unlike the viviparous Aphididae. Some adelgid species are important forestry pests, most notably the hemlock woolly adelgid, Adelges tsugae (Annand) and the balsam woolly adelgid, Adelges piceae (Ratzeburg). Havill and Foottit (2007) present a thorough overview of the biology and evolution of the family.

The closest relatives of the Adelgidae are two extinct families, Elektraphididae and Mesozoicaphididae, the three families comprising the superfamily Adelgoidea (Heie and Wegierek 2009). The oldest fossils of the extinct families date to the Cretaceous, whereas the only fossil adelgid, Adelges balticus Wegierek, 2003, is Eocene in age. Heie and Wegierek (2011) present a list of the fossil Aphidomorpha, including the one adelgid and the 20 other Adelgoidea species.

Adelgid classification has long been unstable. It was first hampered by the adoption of the genus name Chermes Linnaeus, 1758, which had also been used for psyllids and scale insects (Favret et al. 2014). The name was eventually suppressed by the International Commission on Zoological Nomenclature (Evans and China 1965), but not before 137 nominal species had been described in combination with Chermes, including 43 Adelgidae (Favret et al. 2014). Towards the end of the 19^th Century and in the first half of the 20^th Century, several workers described many new species and erected new genera. Most notably, there was little agreement about adelgid taxonomy between the two most prolific adelgid workers, the Russian N.A. Cholodkovsky and the German C. Börner. With the application of molecular data and explicit systematic analyses (Havill et al. 2007, Havill and Foottit 2007), adelgid classification has become more stable. Many researchers have adopted a system with two genera (Adelges Annand and Pineus Shimer) and several subgenera, although some have preferred to treat the latter as full genera (Binazzi 1984, Zurovcová et al. 2010).

While several world catalogs and species lists of the more diverse Aphididae have been published (Wilson and Vickery 1918, Patch 1938, Hille Ris Lambers and Eastop 1976, Remaudière and Remaudière 1997), a comprehensive catalog of the Adelgidae has never been produced. Perhaps the small size of the family contributed to its neglect, but the relative inaccessibility of the taxonomic literature, written in multiple languages and in often hard-to-find sources, possibly made the task too daunting. We here present a taxonomic and nomenclatural list of the extant adelgid nominal taxa, including six family-group, 22 genus-group, and 106 species-group names. Of these, we list one valid family (Adelgidae), two and seven valid genera and subgenera (not counting nominotypical subgenera), and 65 and five valid species and subspecies (not counting nominotypical subspecies). We include one genus-group nomen dubium, four suppressed genus-group names, three species-group nomina dubia, and two unavailable species-group names; the many such names still in combination with Chermes are not listed here (Favret et al. 2014).

Despite having nomenclatural priority, Coccus laricis Bouché, 1834 has long been treated as a synonym of A. laricis Vallot, 1836 (Börner 1952, Steffan 1972, Eastop and Hille Ris Lambers 1976). Coccus laricis Bouché has not been used as a valid name after 1899, meeting the requirements of nomen oblitum per ICZN Article 23.9.1.1 (International Commission on Zoological Nomenclature 1999). Meanwhile, Adelges laricis Vallot has been used frequently. To address ICZN Article 23.9.1.2 and thus consider A. laricis Vallot a nomen protectum, we here list 25 works that used the name as valid, published by at least ten authors in the immediately preceding 50 years and encompassing a span of not less than ten years (Carter 1971, Podeur 1971, Steffan 1972, Li and Tsai 1973, Parry 1973, Eichhorn and Carter 1978, Zhang et al. 1980, Fang 1982, Binazzi 1984, Luo 1988, Rohfritsch 1988, Eichhorn 1994, Battisti et al. 1997, Fang and Yan 1997, Szklarzewicz et al. 2000, Wegierek 2002, Rożkowski 2004, Skrzypczyńska 2004, Havill and Foottit 2007, Havill et al. 2007, Zurovcová et al. 2010, Michalik et al. 2012, Sano and Ozaki 2012, Toenshoff et al. 2012, Gavrilov-Zimin et al. 2015).

In order to facilitate future species descriptions, we followed the model of recent aphid genus-group catalogs by including information on etymology and grammatical gender (Favret et al. 2008, 2009, Cortés Gabaudan et al. 2011, Nieto Nafría et al. 2011). In cases where two page numbers are provided for original descriptions, the first number refers to a nomenclaturally valid diagnosis, such as in a dichotomous key, the second references the formal description. Valid names are in bold font, synonyms are preceded by ‘=’. Synonyms of family-group names are presented with their rank-specific endings replaced by ‘__’. Nominal species are listed under their current generic placement with the original genus in parentheses. The taxonomic catalog is followed by an alphabetical index to help find the current placement of each name. It is our hope that this catalog will serve as an initial point of convergence in our understanding of adelgid systematics and a point of departure on which future research will be built. Future updates will be published on Aphid Species File (Aphid.SpeciesFile.org).

ADELGIDAEAnnand 1928:31

Original spelling. Adelginae

Type genus. Adelges Vallot 1836

=CHERMES__ Herrich-Schaeffer 1854:VIII

Original spelling. Chermiden

Type genus. Chermes Linnaeus 1758

Note. Suppressed (Evans and China 1965)

=CHERMAPH__ Hunter 1901:70

Original spelling. Chermaphinae

Type genus. Chermaphis Maskell 1884

=DREYFUSI__ Börner 1930:155

Original spelling. Dreyfusiini

Type genus. Dreyfusia Börner 1908a

=PINE__ Nüsslin in Börner 1909c:51

Original spelling. Pineini

Type genus. Pineus Shimer 1869

=SACCHIPHANT__ Steffan 1968:128

Original spelling. Sacchiphantini

Type genus. Sacchiphantes Curtis 1844

ADELGES Vallot 1836

Subgenus ADELGESVallot 1836:72

Type species. Adelges laricis Vallot 1836, by original monotypy

Etymology. Greek adelos ‘unclear’, ‘secret’ + Greek ge ‘earth’ + -s [concealed in the earth]

Gender. Masculine

=ANISOPHLEBA Koch 1857:320

Type species. Anisophleba hamadryas Koch 1857, by subsequent monotypy

Etymology. Greek anisos ‘unequal’ + Greek phleps ‘vein’ + -a

Gender. Feminine

=LARICETHUS Amyot 1847:485

Type species. Adelges laricis Hartig 1839, by original monotypy

Etymology. Greek larix ‘larch’ + Greek eth- ‘custom’, ‘habit’ + -us

Gender. Masculine

Note. Suppressed (China 1963)

aenigmaticusAnnand 1928:34,73 (Adelges)

geniculatus (Ratzeburg 1844:202) (Chermes)

isedakii Eichhorn in Eichhorn and Carter 1978:280 (Adelges)

japonicus (Monzen 1929:71) (Chermes)

karamatsuInouye 1945:54,86 (variety of Adelges laricis Vallot)

lapponicus (Cholodkovsky 1889b:390) (Chermes)

=praecox (Cholodkovsky 1898:28) (variety of Chermes lapponicus Cholodkovsky)

lariciatus (Patch 1909:137) (Chermes)

laricisVallot 1836:72 (Adelges) (nomen protectum with respect to Coccus laricis Bouché)

subspecies laricisVallot 1836

=atratus (Buckton 1883:39) (Chermes)

=coccineus (Ratzeburg 1843:202) (Chermes)

=consolidatus (Patch 1909:137) (Chermes)

=hamadryas (Koch 1857:320) (Anisophleba)

=lariceti (Altum 1889:279) (Chermes)

=laricis (Bouché 1834:22) (Coccus) (nomen oblitum with respect to Adelges laricis Vallot)

=laricis (Hartig 1839:644) (Chermes)

=obtectus (Ratzeburg 1844:200) (Chermes)

=strobilobius (Kaltenbach 1843:203) (Chermes)

subspecies potaninilaricis Zhang in Zhang et al. 1980:383 (subspecies of Adelges laricis Vallot)

tardoides (Cholodkovsky 1911:175) (variety of Chermes strobilobius Kaltenbach)

tardus (Dreyfus 1888:3) (Chermes)

=affinis (Börner 1908b:167) (Cnaphalodes) (Börner 1908a:417 nomen nudum)

=niger (Solowiow 1924:41) (Chermes)

Subgenus ANNANDINAFavret et al. 2015:176

Type species Adelges tsugae Annand 1924b, by original designation

Etymology. (Percy Nicol) Annand [American entomologist] + -ina

Gender. Feminine

tsugaeAnnand 1924b:79 (Adelges)

Subgenus APHRASTASIABörner 1909a:1

Type species. Chermes pectinatae Cholodkovsky 1888, by original monotypy

Etymology. Greek aphrastos ‘unnoticed’ + -ia

Gender. Feminine

pectinatae (Cholodkovsky 1888:47) (Chermes)

subspecies ishiharai (Inouye 1936:75) (Chermes)

subspecies pectinatae (Cholodkovsky 1888)

Subgenus CHOLODKOVSKYABörner 1909b:1

Type species. Chermes viridanus Cholodkovsky 1896, by original monotypy

Etymology. (Nikolai Alexandrovitsch) Cholodkovsky [Russian entomologist] + -a

Gender. Feminine

oregonensisAnnand 1928:34,67 (Adelges)

viridanus (Cholodkovsky 1896:39) (Chermes)

=laricicola (Shinji 1930:151) (Chermes)

viridulus (Cholodkovsky 1911:175) (Chermes)

Subgenus DREYFUSIABörner 1908a:416

Type species. Chermes piceae Ratzeburg 1844, by subsequent designation (Börner 1908b:138)

Etymology. (Ludwig Theodor) Dreyfus [German entomologist] + -ia

Gender. Feminine

abietispiceae (Stebbing 1903:57) (Chermes) nomen dubium (Choldkovsky (1906:49, 1915:60) indicates that the species is probably real but that the description is insufficient to confirm its identity; Schneider-Orelli and Schneider (1954:414) agree that Stebbing’s description provides insufficient morphological information to compare with their description of Dreyfusia knucheli Schneider-Orelli and Schneider. Ghosh (1983:11) considered it a synonym of Dreyfusia knucheli, but qualified it with a question mark)

=himalayensis (Stebbing 1910:100) (Chermes) (unavailable, described in synonymy with Chermes abietispiceae Stebbing 1903)

funitectus (Dreyfus 1888:6) (Chermes) nomen dubium (Favret et al. 2015)

joshii (Schneider-Orelli and Schneider 1959:260) (Dreyfusia)

knucheli (Schneider-Orelli and Schneider 1954:416) (Dreyfusia)

merkeri (Eichhorn 1957:312) (Dreyfusia)

nebrodensis (Binazzi and Covassi 1991:246) (Dreyfusia)

nordmannianae (Eckstein 1890:90) (Chermes)

=nuesslini (Börner 1908c:739) (Dreyfusia)

piceae (Ratzeburg 1844:204) (Chermes)

subspecies canadensis (Merker and Eichhorn 1956:454) (form of Dreyfusia piceae (Ratzeburg))

subspecies occidentalisFoottit and Mackauer 1983:302 (subspecies of Adelges piceae (Ratzeburg))

subspecies piceae (Ratzeburg 1844)

=bouvieri (Cholodkovsky 1902:10) (variety of Chermes piceae Ratzeburg)

pindrowiYaseen and Ghani 1971:191,193 (Adelges)

prelli (Grosmann 1935:822) (Dreyfusia)

schneideri (Börner 1931:684) (Dreyfusia)

todomatsui (Inouye 1945:73,92) (Dreyfusia)

Subgenus GILLETTEELLABörner 1930:157

Type species. Chermes cooleyi Gillette 1907, inherited from replaced name

Etymology. (Clarence Preston) Gillette [American entomologist] + -ella [diminutive suffix]

Gender. Feminine

Note. Replacement name for Gillettea Börner 1909d

=GILLETTEA Börner 1909d:504

Type species. Chermes cooleyi Gillette 1907, by original monotypy

Etymology. (Clarence Preston) Gillette [American entomologist] + -a

Gender. Feminine

Note. Junior homonym of Gillettea Ashmead 1897:69 (Hymenoptera, Cynipidae)

cooleyi (Gillette 1907:3) (Chermes)

coweni (Gillette 1907:10) (variety of Chermes cooleyi Gillette)

cummingae (Steffan 1968:10,22,42) (Gilletteella) nomen nudum

glandulae (Zhang in Zhang et al. 1980:382) (Gilletteella)

Subgenus SACCHIPHANTESCurtis 1844:831

Type species. Chermes abietis Linnaeus 1758, by original monotypy

Etymology. Greek sakkos ‘coarse cloth’, ‘sail’ + Greek hyphantēs ‘weaver’

Gender. Masculine

=ELATIPTUS Amyot 1847:484

Type species. Chermes abietis Linnaeus 1758, by original monotypy

Etymology. Greek elatē ‘fir’ + Greek ipt- ‘harm’ + -us

Gender. Masculine

Note. Suppressed (China 1963) and junior objective synonym of Sacchiphantes Curtis 1844

=PHLOEOPHTHIRIDIUM van der Hoeven 1849:509

Type species. Chermes abietis Linnaeus 1758, by original monotypy

Etymology. Greek phloios ‘bark’ + Greek phtheir- ‘destroy’ + Greek -idium [diminutive suffix]

Gender. Neuter

Note. Junior objective synonym of Sacchiphantes Curtis 1844

abietis (Linnaeus 1758:454) (Chermes)

=abietislaricis (Eckstein 1890:88) (Chermes) based on Dreyfus’s (1889:82) description of a Larix-associated form of Chermes abietis

=alaeviridis (Solowiow 1924:39) (Chermes)

=gallarumabietis (De Geer 1773:99) (Aphis) nomen dubium

=laricifoliae (Fitch 1858:752) (Chermes)

diversisAnnand 1928:34,69 (Adelges)

karafutonisKono and Inouye 1938:169,170 (Adelges)

kitamiensis (Inouye 1963:341) (Sacchiphantes)

roseigallis (Li and Tsai 1973:135,141,148) (Sacchiphantes)

segregis (Steffan 1961:67) (Sacchiphantes)

torii (Eichhorn in Eichhorn and Carter 1978:284) (Sacchiphantes)

viridis (Ratzeburg 1843:202) (Chermes)

=occidentalis (Cholodkovsky 1910:281) (Chermes)

CHERMES Linnaeus 1758:453

Type species. None (see Favret et al. 2014)

Etymology. Arabic kirmiz ‘crimson’

Gender. Masculine

Note. Suppressed (Evans and China 1965), see Favret et al. (2014) for list of species-group names described under Chermes

GISTELIELLA Strand 1928:46

Type species. Chermes lapidarius Fabricius 1803, inherited from replaced name

Etymology. (Johannes) Gistel [German entomologist] + -i + ella [diminutive suffix]

Gender. Feminine

Note. Replacement name for Aphanus Gistel 1837, nomen dubium

=APHANUS Gistel 1837:111

Type species. Chermes lapidarius Fabricius 1803, by original monotypy

Etymology. Greek aphanēs ‘invisible’

Gender. Masculine

Note. Junior homonym of Aphanus Laporte 1833:35 (Hemiptera, Lygaeidae)

lapidaria (Fabricius 1803:306) (Chermes) nomen dubium (placement unknown)

PINEUSShimer 1869:383

Subgenus PINEODESBörner 1926:240

Type species. Chermes pinifoliae Fitch 1858, by original monotypy

Etymology. Pine(us) [Hemiptera: Adelgidae]+ Greek -ōdēs ‘resembling’

Gender. Masculine

pinifoliae (Fitch 1858:741) (Chermes)

=abieticolens (Thomas 1879:156) (Chermes)

=armiger (Annand 1924a:5) (Chermes)

=montanus (Gillette 1907:14) (Chermes)

Subgenus PINEUSShimer 1869:383

Type species. Coccus pinicorticis Fitch 1855, by original monotypy

Etymology. Latin pineus ‘of or pertaining to pine’

Gender. Masculine

=CHERMAPHIS Maskell 1884:292

Type species. Kermaphis pini var. laevisMaskell 1885, by subsequent designation (Eastop and Hille Ris Lambers 1976:144)

Etymology. Cherm(es) [Hemiptera] + Aphis [Hemiptera: Aphididae]

Gender. Feminine

=CNAPHALODES Amyot and Audinet Serville 1843:595

Type species. Chermes pini Goeze 1778, by original monotypy

Etymology. Greek knaphallon ‘pillow’ + Greek -ōdēs ‘resembling’

Gender. Masculine

=EOPINEUS Steffan 1968:11

Type species. Chermes strobi Ratzeburg 1844, by subsequent designation (Eastop and Hille Ris Lambers 1976:188)

Etymology. Greek ēōs ‘dawn’, ‘early’ + Pineus [Hemiptera: Adelgidae]

Gender. Masculine

=KERMAPHIS Maskell 1885:19

Type species. Anisophleba pini Koch 1857, by subsequent designation (Eastop and Hille Ris Lambers 1976:232)

Etymology. Kerm(es) [Hemiptera: Coccoidea] + Aphis [Hemiptera: Aphididae]

Gender. Feminine

=PITYOPSYLLA Amyot 1847:461

Type species. Chermes pini Goeze 1778, by original monotypy

Etymology. Greek pitys ‘pine’ + Psylla [Hemiptera: Psyllidae]

Gender. Feminine

Note. Suppressed (China 1963) and junior objective synonym of Cnaphalodes Amyot and Audinet Serville 1843

abietinusUnderwood and Balch 1964:523 (Pineus)

armandicolaZhang et al. 1992:360,394 (Pineus)

boerneriAnnand 1928:34,112 (Pineus) (Takahashi (1937:11), Inouye (1945:36), and Ghosh (1983:8) list this species as a synonym of Pineus laevis (Maskell), but Maskell’s (1885:16) description is insufficient to validate synonymy)

boyceiAnnand 1928:35,102 (Pineus)

cembrae (Cholodkovsky 1888:47) (Chermes)

subspecies cembrae (Cholodkovsky 1888)

=sibiricus (Cholodkovsky 1889b:388) (Chermes)

subspecies pinikoreanusZhang and Fang 1981:15 (subspecies of Pineus cembrae (Cholokovsky))

cladogenousFang and Sun 1985:3 (Pineus)

coloradensis (Gillette 1907:16) (Chermes)

cortecicolusFang and Sun 1985:2 (Pineus)

engelmanniiAnnand 1928:36,130 (Pineus)

floccus (Patch 1909:137) (Chermes)

ghaniiYaseen and Ghani 1971:191,199 (Pineus)

harukawaiInouye 1945:13,21,31,80 (Pineus)

havrylenkoiBlanchard 1944:52 (Pineus)

hosoyaiInouye 1945:22,32,81 (Pineus)

konowashiyaiInouye 1945:13,21,34,81 (Pineus)

laevis (Maskell 1885:16) (variety of Kermaphis pini (Koch))

matsumuraiInouye 1945:13,22,37,82 (Pineus)

orientalis (Dreyfus 1888:3,6) (Chermes)

patchaeBörner 1926:239 (Pineus)

pineoides (Cholodkovsky 1903:263) (variety of Chermes pini (Koch))

pini (Goeze 1778:328) (Chermes)

=coniferarum (Cholodkovsky 1889a:222) (Chermes)

=pini (Koch 1857:322) (Anisophleba)

=pinicola (Cholodkovsky 1915:57) (variety of Chermes pini (Koch))

piniyunnanensisZhang et al. 1992:361,394 (Pineus)

sichunanus Zhang in Zhang et al. 1980:381 (Pineus)

similis (Gillette 1907:15) (Chermes)

simmondsiYaseen and Ghani 1971:191,195 (Pineus)

strobi (Hartig 1839:643) (Coccus)

=corticalis (Kaltenbach 1843:197) (Chermes)

=pinicorticis (Fitch 1855:871) (Coccus)

=strobi (Baerensprung 1849:174) (Coccus)

=strobi (Ratzeburg 1844:203) (Chermes)

sylvestrisAnnand 1928:34,115 (Pineus)

wallichianaeYaseen and Ghani 1971:191,202 (Pineus)

We thank Andrew Carmichael (USDA Systematic Entomology Laboratory) for literature research, Juan Manuel Nieto Nafría (University of León, Spain) for expert opinion and a meticulous cross-referencing of taxon names and bibliographic citations, Ed Jarzembowski (Nanjing Institute of Geology and Palaeontology, China) for help with etymologies, and Shalva Barjadze (Agricultural University of Georgia) for translating certain Russian texts. We also thank external reviewers Anders Albrecht and Rimantas Rakauskas and editor Benjamin Price for their time handling our manuscript. Nomenclature work has become much easier with the availability of digital historical literature. We would like especially to acknowledge the Biodiversity Heritage Library and Google Books for making available valuable literature resources. That which was not available online was available through interlibrary loan. 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.