Catalog of the adelgids of the world (Hemiptera, Adelgidae).

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.

Introduction

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

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

Adelgid classification has long been unstable. It was first hampered by the adoption of the genus name 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 , including 43 (Favret et al. 2014). Towards the end of the 19th Century and in the first half of the 20th 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 ( Annand and 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 have been published (Wilson and Vickery 1918, Patch 1938, Hille Ris Lambers and Eastop 1976, Remaudière and Remaudière 1997), a comprehensive catalog of the 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 (), 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 are not listed here (Favret et al. 2014).

Despite having nomenclatural priority, Bouché, 1834 has long been treated as a synonym of Vallot, 1836 (Börner 1952, Steffan 1972, PageBreakEastop and Hille Ris Lambers 1976). 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, Vallot has been used frequently. To address ICZN Article 23.9.1.2 and thus consider 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).

Catalogue

Original spelling.

Type genus.

=CHERMES__

Original spelling. Chermiden

Type genus.

Note. Suppressed (

=CHERMAPH__

Original spelling.

Type genus.

=DREYFUSI__

Original spelling.

Type genus.

=PINE__ Nüsslin in

Original spelling.

Type genus.

=SACCHIPHANT__

Original spelling.

Type genus.

Subgenus

Type species.

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

Gender. Masculine

=

Type species.

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

Gender. Feminine

=

Type species.

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

Gender. Masculine

Note. Suppressed (

=

subspecies

=

=

=

=

=

=

=

=

=

subspecies

=

=

Subgenus

Type species

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

Gender. Feminine

Subgenus

Type species.

Etymology. Greek aphrastos ‘unnoticed’ + -ia

Gender. Feminine

subspecies

subspecies

Subgenus

Type species.

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

Gender. Feminine

=

Subgenus

Type species.

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

Gender. Feminine

=

=

subspecies

subspecies

subspecies

=

Subgenus

Type species.

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

Gender. Feminine

Note. Replacement name for

=

Type species.

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

Gender. Feminine

Note. Junior homonym of

Subgenus

Type species.

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

Gender. Masculine

=

Type species.

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

Gender. Masculine

Note. Suppressed (

=

Type species.

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

Gender. Neuter

Note. Junior objective synonym of

=

=

=

=

=

Type species. None (see

Etymology. Arabic kirmiz ‘crimson’

Gender. Masculine

Note. Suppressed (

Type species.

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

Gender. Feminine

Note. Replacement name for

=

Type species.

Etymology. Greek aphanēs ‘invisible’

Gender. Masculine

Note. Junior homonym of

Subgenus

Type species.

Etymology.

Gender. Masculine

=

=

=

Subgenus

Type species.

Etymology. Latin pineus ‘of or pertaining to pine’

Gender. Masculine

=

Type species.

Etymology.

Gender. Feminine

=

Type species.

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

Gender. Masculine

=

Type species.

Etymology. Greek ēōs ‘dawn’, ‘early’ +

Gender. Masculine

=

Type species.

Etymology.

Gender. Feminine

=

Type species.

Etymology. Greek pitys ‘pine’ +

Gender. Feminine

Note. Suppressed (

subspecies

=

subspecies

=

=

=

=

=

=

=

Index of genus-group and species-group names

Vallot 1836

Thomas 1879 – synonym of

Underwood and Balch 1964 –

Linnaeus 1758 –

Eckstein 1890 – synonym of

Stebbing 1903 –

Annand 1928 –

Börner 1908b – synonym of

Solowiow 1924 – synonym of

Koch 1857 – synonym of

Favret et al. 2015 – subgenus of

Gistel 1837 – synonym of

Börner 1909a – subgenus of

Zhang et al. in Chen 1992 –

Annand 1924a – synonym of

Buckton 1883 – synonym of

Annand 1928 –

Cholodkovsky 1902 – synonym of

Annand 1928 –

Merker and Eichhorn 1956 – subspecies of

Cholodkovsky 1888 –

Maskell 1884 – synonym of

Linnaeus 1758

Börner 1909b – subgenus of

Fang and Sun 1985 –

Amyot and Audinet Serville 1843 – synonym of

Ratzeburg 1843 – synonym of

Gillette 1907 –

Cholodkovsky 1889a – synonym of

Patch 1909 – synonym of

Gillette 1907 –

Fang and Sun 1985 –

Kaltenbach 1843 – synonym of

Gillette 1907 –

Steffan 1968 –

Annand 1928 –

Börner 1908a – subgenus of

Amyot 1847 – synonym of

Annand 1928 –

Steffan 1968 – synonym of

Patch 1909 –

Dreyfus 1888 –

De Geer 1773 – synonym of

Ratzeburg 1844 –

Yaseen and Ghani 1971 –

Börner 1909d – synonym of

Börner 1930 – subgenus of

Strand 1928

Zhang in Zhang et al. 1980 –

Koch 1857 – synonym of

Inouye 1945 –

Blanchard 1944 –

Stebbing 1910 – synonym of

Inouye 1945 –

Eichhorn in Eichhorn and Carter 1978 –

Inouye 1936 – subspecies of

Monzen 1929 –

Schneider-Orelli and Schneider 1959 –

Kono and Inouye 1938 –

Inouye 1945 –

Maskell 1885 – synonym of

Inouye 1963 –

Schneider-Orelli and Schneider 1954 –

Inouye 1945 –

Maskell 1885 –

Fabricius 1803 –

Cholodkovsky 1889b –

Amyot 1847 – synonym of

Altum 1889 – synonym of

Patch 1909 –

Shinji 1930 – synonym of

Fitch 1858 – synonym of

Bouché 1834 – synonym of

Hartig 1839 – synonym of

Vallot 1836 –

Inouye 1945 –

Eichhorn 1957 –

Gillette 1907 – synonym of

Binazzi and Covassi 1991 –

Solowiow 1924 – synonym of

Eckstein 1890 –

Börner 1908c – synonym of

Ratzeburg 1844 – synonym of

Cholodkovsky 1910 – synonym of

Foottit and Mackauer 1983 – subspecies of

Annand 1928 –

Dreyfus 1888 –

Börner 1926 –

Cholodkovsky 1888 –

van der Hoeven 1849 – synonym of

Ratzeburg 1944 –

Yaseen and Ghani 1971 –

Börner 1926 – subgenus of

Cholodkovsky 1903 –

Shimer 1869

Goeze 1778 –

Koch 1857 – synonym of

Cholodkovsky 1915 – synonym of

Fitch 1855 – synonym of

Fitch 1858 –

Zhang and Fang 1981 – subspecies of

Zhang et al. 1992 –

Amyot 1847 – synonym of

Zhang in Zhang et al. 1980 – subspecies of

Cholodkovsky 1898 – synonym of

Grosmann 1935 –

Li and Tsai 1973 –

Curtis 1844 – subgenus of

Börner 1931 –

Steffan 1961 –

Cholodkovsky 1889b – synonym of

Zhang in Zhang et al. 1980 –

Gillette 1907 –

Yaseen and Ghani 1971 –

Baerensprung 1849 – synonym of

Hartig 1839 –

Ratzeburg 1844 – synonym of

Kaltenbach 1843 – synonym of

Annand 1928 –

Cholodkovsky 1911 –

Dreyfus 1888 –

Inouye 1945 –

Eichhorn in Eichhorn and Carter 1978 –

Annand 1924b –

Cholodkovsky 1896 –

Ratzeburg 1843 –

Cholodkovsky 1911 –

Yaseen and Ghani 1971 –