The Aphids


(in alphabetical order)








Xenosiphonaphis Takahashi

Aphidinae: Macrosiphini

Two little-known east Asian species with very broad antennal tubercles and flangeless siphunculi, very similar to Jacksonia, but first instars have no spinules on their hind tibiae.

Xenosiphonaphis conandri  Takahashi   Apterae are pale, colour is unrecorded; BL c.0.8 mm (?) (BL of alata is 1.6-2.1 mm). On undersides of leaves of Conandron ramondioides, growing on wet rocks, in Japan. Heteroecious holocyclic, with a sexual phase on Stachyurus preacox; there are alate fundatrigeniae from this plant in the BMNH collection, leg. M. Sorin. [A record and description by A.K.Ghosh & Raychaudhuri (1968) of an alata from a species of Gramineae/Poaceae in Sikkim, India should probably be referred to Jacksonia papillata.]

Xenosiphonaphis japonica Takahashi    Colour of apterae (fundatrices) is not recorded; BL c. 3.1 mm (Miyazaki 1971, as Jacksonia). Immature alatae are densely covered with cottony wax. On undersides of leaves of Stachyurus praecox in Japan in spring, migrating to an unknown secondary host.

Xerobion Nevsky

Aphidinae: Aphidini

About 20 species related to Aphis, Brachyunuguis and Protaphis with a short antennal terminal process, short dark volcano-shaped siphunculi and a short triangular cauda. They mostly live on Compositae/Asteraceae, especially Anthemidae, although several including the type species are on Chenopodiaceae. Many species were transferred from Absinthaphis by Kadyrbekov (2001c); Mier Durante & Nieto Nafría (1991) compared morphometric data for 8 species previously in that genus, and Nieto Nafría et al. (2005a) reviewed the Spanish species. Kadyrbekov (2014b) revised the genus, describing three new species from Kazakhstan, establishing several synonymies and  providing a key to apterae. Specimens from Sueda sp. in Sinai (BMNH collection, leg. E. Mescheloff) are similar to a specimen illustrated but not described from Korea as Absinthaphis koraiensis Paik (see Remaudière & Remaudière 1997).

Xerobion alakuli (Juchnevitch)    Apterae are dark green, covered with wax powder; BL 0.9-1.2 mm. On Artemisia and Seriphidium spp. in Kazakhstan and western China (Xinjiang). Kadyrbekov (1995a) provided a redescription.

Xerobion album (Remaudière & Davatchi)  (Fig.31d)   Apterae are white due to complete covering of fine wax powder, with basal segments of antennae, legs and siphunculi dark brown, and cauda pale; BL 1.4-2.0 mm. Apterae have secondary rhinaria distributed III 0-8, IV 1-5 and V 0-1, and alatae have them distributed III 11-12, IV 5-6, V 1-2. Found in colonies (with Brachycaudus helichrysi) on flower-stalks and inflorescences of Helichrysum armenium in Iran (original description, as Protaphis albus), and also recorded from Turkey.

Xerobion amurensis (Pashchenko)    Apterae are green, heavily wax-dusted, with dark siphunculi and a pale cauda; BL c.1.4-1.5 mm. On Artemisia desertorum, sometimes in dense colonies on the upper part of the plant, causing stunting of growth and clumping of leaves. In east Siberia (Maritime Territory).

Xerobion barsukense Kadyrbekov   Apterae are dark brown powdered with wax, with blackish siphunculi; BL c.1.4-1.5 mm. On flowers and shoots of Helichrysum arenarium in a desert habitat in south-west Kazakhstan (Aral region), attended by ants (Kadyrbekov 2014b).

Xerobion blascoi (García Prieto & Sanchís Segovia)    Apterae are dull black; BL 1.0-1.6 mm. Apterae have secondary rhinaria distributed III (0-)1-5, IV 0-3, and alatae have them distributed III (0-)4-9, IV (0-)1-3, V 0-2. In very dense colonies on terminal parts of  Artemisia herba-alba (incl. subsp. valentina), often causing deformation. In Los Monegros, Spain (García Prieto & Sanchís Segovia 1998, in Aphis subgen. Absinthaphis).

Xerobion camphorosmae (Tashev)    Apterae are dark grey, but almost covered in life with thick woolly wax, except for small dark spots in front of the siphunculi and a larger dark spot on the posterior abdomen; BL 1.1-1.6 mm. On Camphorosma in Bulgaria, colonising needle-like leaves and creeping stems; also known from Kazakhstan (as Protaphis comphorosmi; Juchnevitch 1974) and western China (Xinjiang). Kadyrbekov (1995a) provided a couplet separating this species from X. eriosomatinum.

Xerobion caspicae (Bozhko)    Apterae are dark green or blackish tinged with green, coated with greyish wax powder, with a rusty-coloured patch between siphunculi (Barbagallo 1996, as Absinthaphis brutii); BL 1.1-1.7 mm. Apterae have secondary rhinaria distributed III 0-6, IV 0-4, V 0-1, and alatae have them distributed III 6-12, IV 2-5, V 0-2. Original description is from Artemisia caspica (an unresolved plant name) in Ukraine. Kadyrbekov (2014b, 2017a) recorded this species from Artemisia sp. and A. terrae-albae in Kazakhstan, and established that X. brutii (Barbagallo),  described from upper parts of stems and leaves of A. variabilis in southern Italy, and X. georgii (Mier Durante & Nieto Nafría), described from A. campestris ssp. glutinosa in Spain, are synonyms. In Spain it is monoecious holocyclic, with oviparae in October (Pérez Hidalgo et al. 2007b) and alate males found in early November (Mier Durante & Nieto Nafría 1991).

Xerobion cinae (Nevsky)    Plate 7g (Fig.12r)   Apterae are dark green or greenish brown with blackish siphunculi, wax-coated; BL 1.2-1.8mm. Apterae have secondary rhinaria distributed III 0-5, IV 0-2 and V 0-1, and alatae have them distributed ANT III 5-11, IV 1-4, V 0-3. On Artemisia and Seriphidium spp., mainly on stems and leaf petioles, sometimes on undersides of leaves which become slightly wrinkled (Ivanoskaya 1960, as Aphis artemisiae), usually ant-attended. In Turkey, Iran, Russia (lower Volga, western Siberia), Central Asia, India (Kashmir)and western China (Xinjiang). Monoecious holocyclic with oviparae and alate males in mid-October to late November in Central Asia (original description, as Cryptosiphum). Kadyrbekov (2014b) redescribed apterae and alatae  and established that X. artemisiae (Narzikulov) and X. terraealbae (Ivanoskaya) were synonyms. 2n=8.

Xerobion compositae Kadyrbekov   Apterae are dark green with blackish siphunculi, without wax powder; BL c.1.5-1.6 mm. On undersides of leaves of Cousinia perovskiensis in southern Kazakhstan, attended by ants (Kadyrbekov 2014b). Oviparae identified as this species were collected from Cirsium arvense in September in central Kazakhstan.

Xerobion desertorum Kadyrbekov   Apterae are dark green with blackish siphunculi, without wax powder; BL c.1.0-1.2 mm. On flowers and shoots of  Helichrysum arenarium, attended by ants, in a desert habitat in south-west Kazakhstan (Aral region) (Kadyrbekov 2014b).

Xerobion eriosomatinum Nevsky  Plate 7i  (Fig.36e,f)   Apterae are dark brown with a covering of bluish-white wax, except for dark antesiphuncular and postsiphuncular spots: BL 1.3-1.8 mm. Apterae and alatae both have secondary rhinaria distributed III 0-5, IV 0-1. In ant-attended colonies on stems and lower and upper sides of leaves on Kochia prostrata in Spain, northern Italy (Jörg & Lampel 1988) and eastward to Pakistan (Naumann-Etienne & Remaudière 1995) and China (L. Zhang & G. Zhang 2000b, as Aphis (Zyxaphis) kochiae). Monoecious holocyclic; oviparae were described from Kazakhstan (Smailova 1974), and from Spain (Mier Durante et al. 1989).

Xerobion eteriae Barjadze & Gratiashvili  Appearance in life is unknown; BL of apterae 0.9-1.4 mm (of alatae 1.2-1.9 mm). Alatae have 2-10 secondary rhinaria on ANT III only. On Acantholimon spp. in Iran (Barjadze & Gratiashvili 2020).

Xerobion hortobagyi (Szelegiewicz)    Apterae are brownish, without wax; BL 1.0-1.2 mm. On terminal parts of Artemisia maritima ssp. monogyna (= A. santonicum), not visited by ants. In Hungary and Slovakia.

Xerobion inthybi Bozhko    Apterae are dark brownish green; BL c.1.5 mm. On undersides of leaves of Cichorium intybus in Ukraine. Possibly this species should be in Aphis (Protaphis).

Xerobion juchnevitchae Smailova    Apterae are dark green or  brownish, wax-dusted, with blackish siphunculi; BL 1.2-1.8 mm. Secondary rhinaria are confined to ANT III in both apterae (0-2) and the single known alata (9). On apical parts or on undersides of leaves of  Atriplex cana in central and eastern Kazakhstan, visited by ants. Kadyrbekov (2014b) provided a redescription.

Xerobion judenkoi (Szelegiewicz)    Apterae are green, lightly wax-powdered; BL 1.4-1.7 mm. On apical parts of Artemisia campestris in Poland, and also recorded from Artemisia spp. in Latvia, Lithuania and northern Kazakhstan. An aphid on the anomalous host Achillea millefolium in Lithuania identified as this species had a relatively long antennal terminal process, warranting further investigation (Rakauskas et al. 2008).

Xerobion lambersi (Tashev)    Apterae are pale yellow-green to green with pale brown siphunculi, without wax powder (cf. X. cinae); BL 1.3-1.7 mm. Secondary rhinaria distributed III (0-)1-7, IV (0-)1-3, V 0-2 in apterae and III 5-12, IV 2-5, V 1-3 in alatae. On flowers and shoots of Artemisia spp. of the “Oligosporus group” (campestris, dracunculus, marschalliana, scoparia), attended by ants. Severe stunting and some witch’s broom formation is reported on A. scoparia in Pakistan (Naumann-Etienne & Remaudière 1995, as Aphis (Absinthaphis) cinae). This species was redescribed and removed from synonymy with X. cinae by Kadyrbekov (2014b). It was originally described from Bulgaria and is also recorded from Moldova, Ukraine, Iran, Kazakhstan, Afghanistan and Pakistan.

Xerobion pannonica (Szelegiewicz)    Apterae are brown, without wax, almost shiny; BL 1.3-1.5 mm. In ant-attended colonies on basal parts of Artemisia absinthium in Hungary, and also recorded from Romania, Slovakia and Kazakhstan.

Xerobion zoijae (Nevsky)    Apterae are yellow-green with large dark spots on dorsal abdomen and dark brownish femora and siphunculi; BL 1.7-2.3mm. On stems and leaves of  Hyalea pulchella in Turkmenistan (Repetek) and southern Kazakhstan. Alatae have secondary rhinaria distributed III 5-6, IV c.1 (original description, as Brachyunguis zoijae; transferred to Xerobion by Kadyrbekov 2001c).

Yamatocallis Matsumura


Ten oriental species associated with Acer and perhaps most closely related to the North American genus Drepanaphis, but they are larger aphids, lack dorsal processes, and often have dark pigment on the forewings and distal reticulation on the siphunculi. All viviparae are alate. Accounts are available from India (Chakrabarti 1988, A.K. Ghosh & Quednau 1990), China (Qiao & Zhang 2001c, Qiao et al. 2005b), Korea (An & Park 1993) and Japan (Higuchi 1972, 1974; Sugimoto 2017a). Sugimoto (2017a) provided keys to viviparae, oviparae, males and first instar nymphs of the five Japanese species. The species from Acer buergerianum in China described as Y. obscura in Qiao et al. (2005b) requires a new name. Fukatsu (2001) reported that secondary intracellular symbiotic bacteria occur in at least two species of this genus.

Yamatocallis acericola Higuchi   Alatae are green, with forewing membrane diffusely pigmented towards wing-tip and siphunculi blackish-brown except for pale bases; BL 3.1-4.8 mm. On upper surfaces of leaves of Acer carpinifolium in Japan (Higuchi 1974). Sexual morphs collected in November were described by Sugimoto (2017a). [The species keyed under this name in Qiao et al. (2005b, p. 407) is the one described on pp. 106-7 of the same work as Y. obscura, but is in fact another species, as yet unnamed.]

Yamatocallis acerisucta Qiao & Zhang   Alatae are pale in life, with distal halves of siphunculi dark, and wing membrane mostly unpigmented; BL c. 2.8 mm.  On an unidentified Acer sp. in Shaanxi, China (Qiao & Zhang 2001).  Biology and sexual morphs are unknown.

Yamatocallis brevicauda Chakrabarti   Colour of alatae in life is not recorded, wing membrane probably unpigmented and siphunculi black-tipped; BL 3.1-3.8 mm. On Acer villosum in Uttar Pradesh, India. Sexual morphs occur in October (Chakrabarti 1988).

Yamatocallis hirayamae Matsumura   (fig. 22G)   Alatae are pale green, green, pinkish brown or yellowish-brown, with pale green or pale brown siphunculi, and forewings with a broad dark band of pigment traversing the anterior part of the membrane from the base of Cu1a to the wing-tip; BL 2.4-3.5 mm. On various Acer spp. in Japan, China, Korea and eastern Siberia (Higuchi 1972, Pashchenko 1988a). Sexual morphs occur in late October to early December; Sugimoto (2017a) provided descriptions of all morphs.

Yamatocallis nikkoensis Sugimoto   Alatae are green with wholly black siphunculi and a broad band of dark pigment on the anterio-distal part except for a contrastingly clear area bordered by the pterostigma and radial sector; BL 2.0-2.9 mm (BL of fundatrix c. 3.4 mm). Sexual morphs were collected in October-November. All morphs were described from a single tree of Acer shirasawanum (= tenuifolium) in Honshu, Japan (Sugimoto 2017a).

Yamatocallis obscura (M.R. Ghosh, A.K. Ghosh & Raychaudhuri)   Alatae are green, and have forewings with membrane unpigmented, and siphunculi dark brown distally; BL 3.4-4.1 mm. On undersides of young leaves of an unidentified plant (assumed here to be an Acer sp.) in West Bengal (M.R Ghosh et al. 1971, as Megalophyllaphis). A.K. Ghosh & Quednau (1990) provided a redescription. The life cycle is unknown. [Qiao et al. (2005b) described a species from Acer buergerianum in China under this name, but their species differs in several respects from Y. obscura (smooth siphunculi without distal reticulation, more numerous rhinaria extending for most of length of antenna, more accessory hairs on R IV+V).]

Yamatocallis sauteri (Takahashi)   Alatae are green with yellowish-brown head and thorax, forewings with a broad band of pigment extending from pterostigma and RS along distal branches of media, and black-tipped siphunculi; BL c. 4.5 mm. On an unidentified Acer sp. in Taiwan (Takahashi 1927a). The life cycle is unknown.

Yamatocallis takagii (Takahashi)   (fig. 22F)   Alatae are green, forewings with RS and distal branches of media thickly brown-bordered, and blackish siphunculi; BL 4.5-4.7 mm. On twigs and undersides of leaves of Acer pictum and A. miyabei in Japan (Sugimoto 2017a), and on A. pictum in Korea (An & Park 1993, as Y. palgongsanensis, which is a synonym). Sexual morphs were collected in Japan in November and described by Sugimoto (2017a). 2n = c. 48 (Blackman 1986).

Yamatocallis tokyoensis (Takahashi)   (fig. 22H)  Alatae are pale green with brown head and thorax (or sometimes in spring body is pinkish brown), with pale green or pale brown siphunculi, and with most of forewing membrane infuscated except for posterior margin; BL 1.5-2.9 mm. On leaves of Acer spp., especially A. palmatum, in Japan and Korea (An & Park 1993; Sugimoto 2017a). Sexual morphs occur in November; all morphs including the fundatrx were described by Sugimoto (2017a). Kan & Sasakawa (1986) studied its relations with a syrphid predator.

Yamatochaitophorus Higuchi


Two east Asian species on Acer, related to Trichaitophorus but with long, thick spinal as well as marginal hairs.

Yamatochaitophorus albus (Takahashi) Apterae are elongate oval, whitish with pale antennae and legs; BL 0.8-1.0 mm. Described from an unidentified Acer sp. at high altitude in Japan (Takahashi 1961b; as Trichaitophorus albus), and aphids identified as this species have subsequently been recorded from India on various Acer spp. (Chakrabarti & Mandal 1986), and from A. pictum in Siberia (see Holman 2009). The alate is described from India by Chakrabarti & Mandal (1986). Biology and sexual morphs are unrecorded.

Yamatochaitophorus yichunensis Jiang, Chen & Qiao   Apterae are elongate oval, pale yellow; BL 0.8-1.4 mm. Feeding dispersed on undersides of leaves of Acer tegmentosum in China (Heilonjiang Prov.). Alatae and life cycle are unknown (Jiang et al. 2016)

Zelkovaphis Barbagallo

Eriosomatinae: Eriosomatini

Two species that form open or leaf-roll galls on Zelkova. One of them (caucasica) is now known to migrate to Carex. Barbagallo (2002) provided a diagnosis, compared it with related genera and provided a key to emigrant alatae. Barjadze et al. (2011a) provided a key to fundatrices, and Sano et al. (2015) provided keys to the alate emigrants and their first instar progeny .

Zelkovaphis caucasica (Dzhibladze)  (= Colopha caucasica of Blackman & Eastop 1994)  Spring generations roll and blister the leaves of Zelkova acuminata and Z. carpinifolia (Dzhibladze 1960). Fundatrices are greyish green or dark green covered with white wax; BL 2.1-2.4 mm (Barjadze et al. 2011a). Emigrant alatae (BL 1.3-1.8 mm) emerge from the galls in May and fly to Carex capitata, where colonies in white woolly wax are formed on the aerial parts (Barjadze et al. 2011a). The alatae were redescribed by Sano et al. (2015). Sexuparae are produced in November. In south-west Asia (Georgia, Iran; BMNH collection).

Zelkovaphis trinacriae Barbagallo   Fundatrices are broadly oval, dark grey, covered with white wax, with very dark brown appendages; BL 2.8-3.3 mm.  The gall on Zelkova sicula consists of a cluster of thickened and modified leaves that overlap one another to form a rosette-like nest enclosing the fundatrix and its progeny, which are all alate (Barbagallo 2002). Emigrant alatae are blackish, BL 1.8-2.5 mm, and migrate mostly in May to an unknown secondary host. It is known only from the type locality in Italy (Sicily).

Zinia Shaposhnikov

Aphidinae: Macrosiphini

A genus for one species combining some characters of Dysaphis and Brachycaudus.

Zinia veronicae Shaposhnikov    Apterae are pale green, with a dark grey slightly shiny dorsal abdominal sclerotic shield, black antennae and siphunculi and dark grey head and eyes; BL c.1.7-1.8 mm. On roots of Veronica spp., without ants. In Russia (Urals), Moldova, Kazakhstan, Poland and Italy. The alata was redescribed from northern Italy by van Harten & Coceano (1981).