The Aphids

Four North American species previously placed in Ovatus but perhaps more closely related to Myzus (Jensen & Stoetzel 1999). Characteristic features are the siphunculi with slight subapical swelling and small flange, and the reticulate dorsal cuticle.

Abstrusomyzus leucocrini (Gillette & Palmer) Apterae are pale green to brownish green or black; BL 1.6-1.8 mm. Described as abundant on leaves of Leucocrinum montanum in Colorado, USA (Gillette & Palmer 1934), but there are subsequent records from Achillea (including oviparae and apterous males collected in October) and Potentilla (A. Jensen, aphidtrek.org) so its true host associations are still uncertain.

Abstrusomyzus phloxae (Sampson) Plate 17h Apterae are pale apple-green (see aphidtrek.org and Influential Points/Gallery); BL 1.2-1.8 mm. Recorded from many plant species and genera in different families (Achillea, Agoseris, Apocynum, Capsella, Carex, Centaurea, Cerastium, Galium, Phacelia, Phlox, Plantago, Polygonum, Ranunculus, Stellaria, Trifolium, Viola). Usually it colonizes the basal or rosette leaves of low-growing plants. Widely distributed in North America; in eastern USA it is most common on Plantago, where colonies are often attended and sheltered by ants (Jensen & Stoetzel 1999). In western USA it is common on Apocynum where it feeds on undersides of leaves causing discoloration visible from above (A. Jensen, aphidtrek.org) Populations from different parts of North America in the BMNH collection show differences in ratio of R IV+V to HT II, but it is unclear whether these are related to host plant or geographical location. Apparently mostly anholocyclic, although males have been produced in culture (BMNH, leg. C.-k. Chan), and oviparae were collected from Fragaria vesca in Nova Scotia (BMNH, leg. H.T. Shultz). 2n=18*.

Abstrusomyzus reticulatus (Heie) Apterae are shining back; BL 1.6-1.9 mm. On undersides of curled leaves of Oxalis sp. (probably stricta) in North Carolina, USA. Oviparae and alate males occur in October.

Abstrusomyzus valuliae (Robinson) Apterae are blackish, BL c.1.2-1.3 mm. Alatae have 2-7 rhinaria on ANT III. On Fragaria vesca in Manitoba and Alberta (Canada).

Acanthochermes Kollar

Phylloxeridae

A genus of two palaearctic species, with spiracles on abdominal segments 2-5 as in Phylloxera and Viteus, but characterized by the curious star-like tubercles borne by immatures. The life cycle is remarkable in having only two generations per year, fundatrices and sexuales. The adult fundatrices bear small spiculose tubercles on head, thorax and abdomen.

Acanthochermes quercus Kollar Recorded from Quercus robur and Q. petraea in European Russia, Finland, Sweden, central Europe (Hungary, Germany), Portugal and the Middle East (Iran). The large fundatrices (BL up to 3 mm) induce small ring-shaped galls on the upper surface of the leaf, opening on the underside (see Chen & Qiao 2012a). Morphology and developmental stages were illustrated by Grassi (1912), and Tavares (1931) gave an account of the biology.

Acanthochermes similiquercus Jiang, Huang & Qiao Fundatrices are yellow dotted with black; BL 1.6-2.3 mm. In small yellow, yellowish green or red galls on leaves of an unidentified Quercus sp. in Sichuan, China (Jiang et al. 2009).

Acaudella Nevsky

Aphidinae: Macrosiphini

One palaearctic species on Atraphaxis (Polygonaceae) with 5-segmented antennae, no evident cauda and characteristically shaped siphunculi. A second species previously noted as belonging to Acaudella has been described in a new genus (Blackmania).

Acaudella puchovi Nevsky Apterae are dark green, BL c.1.4 mm. Alatae have 9-19 secondary rhinaria on ANT III, 0-1 on IV. In large numbers on undersides of leaves of Atraphaxis buxifolia (= caucasica) in Uzbekistan (Samarkand). It is also recorded from A. spinosa in both Kazakhstan (Kadyrbekov 2017a) and Israel (Bodenheimer & Swirski 1957). A redescription is provided by Kanturski & Wieczorek (2015b). The life cycle is unknown.

Acaudinum Börner

Aphidinae: Macrosiphini

Six palaearctic species, four of them classified in the subgenus Protacaudinum Holman, with a broadly rounded cauda living without host alternation on species of Centaurea and Serratula. Holman (1991) provided a revision and a key to species of the genus at that time, and Kadyrbekov (2022) a key to species of the nominotypical subgenus. They are adapted for ant attendance.

Acaudinum (Protacaudinum) beheni Remaudière & Davatchi Apterae are shining dark brown, BL 1.5-1.75 mm. On basal leaves of Centaurea behen in western Asia (Iran, Turkey, Armenia, Azerbaijan), attended by ants. Oviparae and apterous (alatiform) males occur in late September (Tuatay & Remaudière 1964).

Acaudinum (Protacaudinum) bulgaricum Holman Apterae are dull dark brown to black , BL 1.4-1.9 mm. On Centaurea solstitialis in Bulgaria, feeding on upper and undersides of leaves and also on stem and branches, attended by ants. An alate male was collected in early September (Holman 1991).

Acaudinum centaureae (Koch) Plate 10h Apterae are shining black (see Influential Points/Gallery), BL 1.7-2.5 mm. At bases of leaf stalks of Centaurea spp., often in ant shelters (Stroyan 1950; Holman 1991). Widely distributed in Europe. Oviparae occur in September-October. 2n=10.

Acaudinum longisetosum Holman Apterae are matt or slightly shiny jet black, BL 1.9-3.2 mm. At stem base and root collar of Centaurea spp., especially C. scabiosa, in spring, later also living on terminal parts of plants, attended by ants. There are also records from Achillea spp. (Wojciechowski et al. 2016) that perhaps require additional confirmation. In Poland, Czech Republic, Slovakia, Switzerland, Italy and Ukraine. Oviparae and apterous males occur in October.

Acaudinum rogeri Kadyrbekov Apterae are dark green, BL 2.1-2.6 mm. At stem bases and leaves of Serratula coronate in Kazakhstan (Kadyrbekov 2022). Closely related to A. longisetosum and A. roumanicum, which live on Centaurea. Alate viviparae and sexual morphs are unknown.

Acaudinum roumanicum Holman Apterae are matt or slightly shiny jet black, BL 1.6-2.7 mm. At stem bases of Centaurea stoebe subsp. stoebe (= S. rhenana)in spring, later on terminal parts of plants, attended by ants. Closely related to A. longisetosum; the discriminant given in the key is based on work of Holman (1991).Romania, Bulgaria and Slovakia. Sexual morphs are unknown.

Acuticauda Hille Ris Lambers

Aphidinae: Macrosiphini

Two nearctic and one palaearctic species on Compositae, possibly related to Brachycaudus but with secondary rhinaria in apterae, capitate dorsal hairs and a differently shaped cauda (Hille Ris Lambers 1956c). Leclant & Remaudière (1967) compared and keyed the species.

Acuticauda asterensis Gillette & Palmer (Fig.15g,h) Apterae are pale yellowish green; BL c.1.3 mm. Alatae have black wing veins. On leaves and stems of Aster and Symphyotrichum spp. in USA (Colorado, New York and Wisconsin) and Canada (Quebec); there is one record from Solidago requiring substantiation. Oviparae and alate males in autumn (Palmer 1952).

Acuticauda erigerontis Leclant & Remaudière Apterae are deep golden yellow with brown antennae and siphunculi; BL 1.2-1.5 mm. Alatae have dark wing veins. On Erigeron acer in the French Alps, rolling the leaves of the basal rosettes (Leclant & Remaudière 1967). Monoecious holocyclic, with apterous males.

Acuticauda solidaginifoliae (Williams) Plate 10c Apterae are shining reddish or greenish brown with a black dorsal shield; BL 1.8-2.0 mm. On leaves of Solidago spp., which are folded along mid-ribs to become pod-like. Widely distributed in northern and western USA. Monoecious holocyclic, with apterous males in September-October (Palmer 1952).

Acutosiphon Basu, Ghosh & Raychaudhuri

Aphidinae: Macrosiphini

One species near Vesiculaphis but with distinctive siphunculi. Hille Ris Lambers (1973) discussed its taxonomic position and probable host relationship..

Acutosiphon obliquoris R.C. Basu, A.K.Ghosh & Raychaudhuri (Fig.18a) Apterae are dark or reddish brown; BL 1.6-2.1 mm. Secondary rhinaria in alata III 25-28, IV 12-14, V 1-3. Originally described from Cotula (Compositae), but this is an unlikely host, and subsequent collections indicate possible host alternation between Ericaceae (Lyonia ovalifolia) and Cyperaceae (Carex, Cyperus); see Hille Ris Lambers (1973b) and Miyazaki (1977). In India (Assam) and Nepal, and more recently fully redescribed and illustrated from China (Fang et al. 2006).

Acyrthosiphon Mordvilko

Aphidinae: Macrosiphini

A genus of about 70 palaearctic and 9 nearctic species living without host alternation on various dicots, particularly Leguminosae/Fabaceae, Rosaceae, and Euphorbiaceae. They are typically rather large broadly spindle-shaped, short-haired aphids with long antennae, legs, siphunculi and cauda, usually green but sometimes brownish, pink, or yellow. Eastop (1971) gave keys for the identification of the species then known, and there are accounts for Fennoscandia and Denmark (Heie 1994), Britain and Ireland (Blackman 2010), Japan (Miyazaki 1971), India (Raychaudhuri, M.R.Ghosh & R.C. Basu 1978) and the Russian far east (Pashchenko 2005). Meier (1958) reviewed, and provided photographs of, the European legume feeders. Nieto Nafría et al. (2015b) provided a list of species.

Acyrthosiphon argus Miyazaki Apterae are pale green, with a dark green spinal stripe; BL 1.8-2.9 mm. Described from Daphne odora in Japan, colonising the undersides of leaves and the flowers. Infested leaves turn yellowish or dark brown, and heavy infestation can cause early defoliation and death of shrubs (original description). Previously collected on Daphne sp. in China (BNMH collection, coll. VFE, 31.v.1985). Populations in Japan (perhaps introduced from the Asian mainland) were apparently anholocyclic.

Acyrthosiphon assiniboinensis Robinson Apterae are green; BL 2.2-2.8 mm. Alatae have 15-18 secondary rhinaria on ANT III. On Potentilla fruticosa in Canada (Manitoba, Northwest Territories, Yukon).

Acyrthosiphon astragali Eastop Apterae are green, BL 2.2-2.8 mm. In large colonies on stems of Astragalus sp(p)., falling readily when disturbed (Narzikulov 1972). Recorded from Afghanistan, Pakistan, Tajikistan and Kashmir (BMNH collection). Alatae are undescribed.

Acyrthosiphon auctum (Walker) Apterae are pale yellow-green, green or pale pink, slightly wax bordered intersegmentally; BL 1.9-2.8 mm. Alatae have a dark head and thorax and reddish abdomen. On various Caryophyllaceae (Cerastium, Honkenya, Silene, Stellaria) and Brassicaceae (Cakile, Capsella), mainly in sand dune habitats, in north Europe, including Iceland (Heie 1994), and Greenland (BMNH collection, leg. J.M. Cotton). Monoecious holocyclic with oviparae and apterous males in September-October (in Scotland; BMNH collection, leg. H.L.G. Stroyan).

Acyrthosiphon auriculae Martin Apterae are pale to bright green; BL 1.8-2.85 mm. On upper surfaces of young apical leaves of a cultivated hybrid Primula (section Auricula); experimental host transfers indicated specificity to this section of the genus (Martin 1981). Only known from England, although alatae of a possible geographic variant of this species were trapped in Corsica. Monoecious holocyclic with apterous males. 2n=8.

Acyrthosiphon bidenticola Smith Apterae are bright green with mainly dark antennae, legs and siphunculi; BL 1.6-2.0 mm. On various Compositae (Aster, Bidens, Erechtities, Gnaphalium, Helianthus, Lagascea, Tithonia, Wedelia), and populations have also been found on Bocconia frutescens (Mexico, BMNH collection, leg. R. van den Bosch) and Salvia coccinea (Jamaica, Tannice Hall, pers. comm.). Central and South America (Brazil, Cuba, Jamaica, Costa Rica, Mexico, Puerto Rico, Venezuela). 2n=8*

Acyrthosiphon bistorti Ivanoskaya Apterae are yellowish green; BL c. 4.1 mm. Alatae have about 8 secondary rhinaria on ANT III. In inflorescences of Persicaria bistorta in west Siberia, and also now recorded from Polygonum and Persicaria spp. in Kazakhstan (Kadyrbekov 2012a) and Russia (Altai; Kadyrbekov 2017a).

Acyrthosiphon boreale Hille Ris Lambers Apterae are green, yellowish green or yellowish (see influentialpoints.com/Gallery); BL 2.7-3.2 mm. Apterae have 2-9 secondary rhinaria on ANT III. On Potentilla spp. in northern Europe, Greenland, Canada (Baffin Island) and north-east Siberia (Chukotka). There are also records from Switzerland, Hungary and Kazakhstan, although the distinction from other members of the malvae group is not clear enough to be certain about the extent of its distribution. It seems likely that A. malvae ssp. pentaphylloides, described from Potentilla fruticosa in east Siberia (Pashchenko 2005) and also recorded from Kazakhstan (Kadyrbekov 2016b), is a synonym. Monoecious holocyclic, with apterous males. Stekolshchikov & Khruleva (2015) provided detailed descriptions of the male and ovipara. 2n=10.

Acyrthosiphon brachysiphon Hille Ris Lambers Apterae are waxy green in life, 1.8-2.8 mm, with siphunculi hardly longer than cauda, their apices slanted outwards. This aphid has a boreoalpine distribution on Vaccinium spp. (Blackman & Eastop 2006), but there is a record of apterae and oviparae from Salix lapponum in Norway (Heikinheimo 1966, as A. aurlandicum).

Acyrthosiphon brevicorne Hille Ris Lambers Colour of apterae unrecorded, probably green, like oviparae; BL of aptera 1.8-2.3 mm. On Dryas spp. Distribution is boreoalpine; northern Canada, Greenland, Sweden and Switzerland. Monoecious holocyclic, with oviparae and apterous males in July-August in the Arctic, and in October in Switzerland (Meier 1972). Strathdee et al. (1995) studied cold-hardiness of eggs under laboratory conditions. A. dryasae described from Sakhalin is possibly this species.

Acyrthosiphon capitellum Zhang Apterae are green with dark apices to antennal segments, femora and tibiae, and dark basal parts of siphunculi, BL c. 3.3 mm. On Fragaria vesca in China (G. Zhang & Qiao 1998c, 1999d).

Acyrthosiphon caraganae (Cholodkosvsky) Apterae are green with variably developed greyish wax stripes between segments, and often a bright green spinal stripe; BL 2.6-4.3 mm. On leaves and young growth of woody Leguminosae, especially species of Caragana and Colutea, often forming large colonies. Probably originating in the Altai region, it now occurs in parks and gardens in temperate regions throughout the Northern Hemisphere. Monoecious holocyclic; males are apterous in western Europe and alate in Russia, Switzerland and eastern Canada (Eastop 1971). Populations in western Europe, Tajikistan and Sakhalin have been distinguished as subspecies; A. caraganae occidentale Hille Ris Lambers, A. caraganae tadzhikistanicum Narzikulov and A. caraganae sachalinense Pashchenko (2005). 2n=10 (also for ssp. occidentale).

Acyrthosiphon (Liporrhinus) chelidonii (Kaltenbach) Apterae are pale green or greenish white, covered with greyish-white wax powder except along segmental borders; BL 1.5-2.3 mm. On Chelidonium majus, forming colonies on upper parts of stems and on young leaves. In Europe, Kazakhstan (Kadyrbekov 2017a), east Siberia, Korea and Japan. Oviparae and alate males in Netherlands in October-November (Hille Ris Lambers 1947).

Acyrthosiphon churchillense Robinson Apterae are green; BL 2.1-2.6 mm. On Oxytropis spp. and some other Leguminosae (Melilotus alba, and probably Hedysarum boreale ssp. mackenzii) in northern Canada (northern Manitoba, Yukon, Northwest Territories). Monoecious holocyclic with oviparae and apterous males in July-August (Robinson 1979 and BMNH collection, leg. AG Robinson). Close to A. kondoi; see couplet for separating these species on Oxytropis.

Acyrthosiphon corsicae Remaudière & Leclant Apterae are pale green; BL 2.2-2.6 mm. Alatae have 8-10 rhinaria on ANT III. On Euphorbia spinosa in Corsica (Remaudière & Leclant 2000).

Acyrthosiphon crepidis Holman & Szelegiewicz Apterae are green, with ventral side pruinose; BL 1.7-2.2 mm. Other morphs are unknown. On Crepis (= Youngia) tenuifolia in Mongolia.

Acyrthosiphon cyparissiae (Koch) Apterae are green with black front part of head and mainly black appendages or, especially in warmer climates, yellowish green with appendages mainly brown or brownish yellow (see influentialpoints.com/Gallery); BL 2.4-3.4 mm. On Euphorbia spp., feeding mainly on upper sides of upper leaves, often in large numbers. In Europe (but UK records apply to other species), Middle East, Central Asia and China . Monoecious holocyclic with alate males (Hille Ris Lambers 1947). In warmer regions the less-pigmented form A. cyparissiae ssp. propinquum Mordvilko is more common (Eastop 1971), although it needs to be confirmed that this is not merely an effect of the environment. Another nominal subspecies, A. cyparissiae ssp. turkestanicum Nevsky, is reported from Central Asia.

Acyrthosiphon daphnidis Ilharco Apterae are green to yellowish green with red eyes, body with greyish white powder accentuating dorsal segmentation; BL 2.4-2.8 mm. In colonies on undersides of leaves of Daphne gnidium; severe infestations can cause yellowing and dropping of leaves. In Portugal, Spain, Canary Islands (Ilharco 1996b), Corsica and northern Italy (Barbagallo et al. 2011), and also reported from Iran (Rezwani & Parvizi 1990).

Acyrthosiphon dauricum Szelegiewicz Apterae are green; BL 2.3-2.6 mm. Described from fundatrices and apterae on a composite plant in Mongolia, that was subsequently identified as Saussurea salicifolia (Szelegiewicz 1964a). However, it could not be verified that this was the true host, and subsequent records from Delphinium grandiflorum and Linaria buriatica (Holman & Szelegiewicz 1972) have not helped to clarify the situation.

Acyrthosiphon dryasae Pashchenko Apterae are pale green: BL c.2.1 mm (Pashchenko 2005). On Dryas punctata in east Siberia (Kamchatka). Apart from the slightly longer siphunculi, the description does not clearly distinguish this species from A. brevicorne.

Acyrthosiphon echinospartii Nieto Nafría & Mier Durante Apterae are pale green, with somewhat darker appendages; BL 2.6-3.1 mm. On Echinospartium lusitanicum in Spain, where it has also subsequently been recorded from Genista sp. (Mier Durante et al. 1995).

Acyrthosiphon elaeocarpi Tao Apterae are whitish yellow with yellowish brown siphunculi and blackish brown antennae, apices of tibiae, tarsi, cauda and anal plate; BL c. 3.5 mm. On Elaeocarpus serratus in Taiwan (Tao 1963). Biology and sexual morphs are unknown.

Acyrthosiphon ericetorum Hille Ris Lambers Colour of apterae in life unknown, probably pale green; BL c. 1.9-2.3 mm. On Genista spp. in the Netherlands. Monoecious holocyclic; the original description was of oviparae and apterous males. Apterous viviparae were described by Nieto Nafrìa & Mier Durante (1985). This aphid only seems to differ from A. genistae by its small size, which could be an effect of the host plant.

Acyrthosiphon euphorbiae Börner Apterae are green, tinged with yellow; BL 2.3-4.4 mm. On Euphorbia spp., in small colonies on upper sides of leaves, sometimes mixed with E. cyparissiae. In Europe, North Africa, the Middle East and Central Asia. Alatae have 11-25 rhinaria on ANT III. Sexual morphs are undescribed and the life cycle is apparently unknown. Narzikulov (1953) described a population in Tajikistan as a subspecies, A. euphorbiae ssp. tadzhikistanicum. Another subspecies described from western Europe, A. euphorbiae ssp. neerlandicum Hille Ris Lambers, was established to be a synonym by Remaudière & Leclant (2000). The range of measurements recorded for populations in Kazakhstan described as another subspecies, A. euphorbiae ssp. brevisiphon Kadyrbekov, is within that recorded for A. euphorbiae from other parts of the world (Kadyrbekov 2005e, as A. neerlandicum ssp. brevisiphon). A closely related form, A. pareuphorbiae, occurs in China.

Acyrthosiphon evodiae (Takahashi) Apterae are yellow in life, about 2.3 mm, with black antennae and legs, brown-black siphunculi and a pale cauda. Alatae have wings “somewhat clouded along the veins”. Described (in Macrosiphum) from specimens found feeding on young leaves of Euodia (= Melicope) triphylla in Taiwan (Takahashi 1929). Aphids collected on Acronychia pedunculata in Hong Kong were provisionally identified as this species (Martin & Lau 2011, as Sinomegoura evodiae). The generic placement of this species is uncertain; it may belong in Sinomegoura.The life cycle is unknown.

Acyrthosiphon extremiorientale Pashchenko Apterae are green, emerald green or orange, with antennae and siphunculi mainly dark; BL 4.2-4.9 mm. The single known alata had 40 rhinaria on ANT III. Feeding on Vicia spp. and Astragalus schelichowii in east Siberia (Pashchenko 2005). The life cycle is unknown.

Acyrthosiphon fragariaevescae Nevsky Apterae are brownish or green, with pale antennae, siphunculi and cauda; BL c.2 mm. Alatae have 13-22 rhinaria on ANT III, and ANT PT/BASE c. 4.1. On Fragaria vesca in Central Asia. Kadyrbekov (2005e) provided a redescription of the aptera. Apart from the shorter ANT PT this species seems close to A. rubi.

Acyrthosiphon fragum Zhang Apterae are green with black siphunculi; BL c. 3.9 mm. On Fragaria sp. in China (G. Zhang & Qiao 1998c, 1999d).

Acyrthosiphon galijae Kadyrbekov Apterae are green; BL 3.3-4.0 mm. On stems and leaves of Sanguisorba officinalis in Kazakhstan (Kadyrbekov 2005e). Very similar to A. sanguisorbae;, and the differences (size, hair length, etc.) could be due to host plant and other environmental factors.

Acyrthosiphon genistae Mordvilko Apterae are green to yellow green, with dark brown siphunculi; BL c. 3.4 mm. On Genista tinctoria, at ends of stems and branches. In Russia and Ukraine (Bozhko 1976). A. ericetorum from other Genista spp. in Austria could be small specimens of this aphid.

Acyrthosiphon ghanii Eastop Colour of apterae in life unknown; BL 2.0-2.4 mm. On Compositae/Asteraceae (Sonchus, Scorzonera) in Pakistan (Naumann-Etienne & Remaudière 1995). 2n=10? (Kapoor & Gautam 1994, but aphid was possibly misidentified as host was Medicago).

Acyrthosiphon (Xanthomyzus) glaucii (Narzikulov) Apterae are shining greenish white to white; BL 1.8-2.2 mm. On Glaucium spp. in Central Asia. It is also recorded from Hypecoum parviflorum (Kadyrbekov 2017a) [Host plant of type material is given as G. fimbrilligerum in original description, but as G. corniculatum on slide labels of specimens in BMNH collection with same date and locality, leg. Narzikulov.]

Acyrthosiphon gossypicola Shinji Perhaps a synonym of Acyrthosiphon gossypii, but apterae were described as reddish brown in life, and alatae have only 7-9 secondary rhinaria on basal half of ANT III, whereas those of A. gossypii have 10-21 distributed over most of its length (Eastop 1971). On Gossypium herbaceum in Korea.

Acyrthosiphon gossypii Mordvilko Plate 23c Green in life, with adults as well as immatures dusted with fine wax (see aphids of Karnataka website); BL of apterae 2.5-3.8 mm. Host plants are mainly Leguminosae/Fabaceae especially tribe Phaseoleae (Dolochos, Phaseolus, Vigna, but not known from Glycine), Malvaceae (including cotton) and Zygophyllaceae; more rarely on other plants including Brassicaceae. Trees recorded from include Hibiscus esculentus, Robinia pseudoacacia and Sophora japonica. Intraspecific variation in host plant preferences and life cycle are discussed by Müller & El Tigani (1986). Populations with particular host preferences may occur in different regions (Blackman & Eastop 2000). In southern Europe, North Africa, Middle East, south-west and central Asia, India, China, and possibly Korea and Japan. Anholocyclic over much of its range, but there is a holocycle on Alhagi camelorum in central Asia. A. umarovi (q.v.) is probably a synonym. 2n=6.

Acyrthosiphon heptapotamicum Kadurbekov Apterae are green; BL 3.2-3.9 mm. On stems and leaves of Clematis songoricum in Kazakhstan (Kadyrbekov 2005e). This aphid has long thin siphunculi like those of A. gossypii, but with a longer ANT PT than that species, and more caudal hairs.

Acyrthosiphon hissaricum Umarov Apterae are green; BL c.3.9 mm. Alatae have BL c. 3.1 mm and 20-30 secondary rhinaria on ANT III. On Cicer songaricum in Tajikistan, and also reported to occur on Medicago falcata and Hedysarum spp. in Kazakhstan (Kadyrbekov 2012a, 2014d). Similar to A. rubi, but much larger..

Acyrthosiphon ignotum Mordvilko Apterae are pale green or yellowish green; BL 2.2-3.4 mm. Apterae have 1-8 secondary rhinaria on ANT III, and alatae have 7-13. At shoot tips of Spiraea spp., often in large colonies. Also recorded from Sibiraea laevigata. In northern Europe, Iran, Kazakhstan, north India, Siberia, Mongolia (many records are as A. spiraeae). A. leleji may be a synonym. 2n=10 (Kuznetsova & Shaposhnikov 1973), but Khuda-Bukhsh & Pal (1986b) record 2n=14 for an aphid identified as this species collected on Deutzia corymbosa.

Acyrthosiphon ilka Mordvilko (= Acyrthosiphon bidentis Eastop) Apterae are green, often with darker green dorsal cross-bands; BL 1.4-2.7 mm. On Papaveraceae (Glaucium, Papaver), Compositae/Asteraceae (Bidens, Carthamus, Cichorium, Lactuca, Lampsana, Picris, Sonchus), Brassicaceae (Erysimum, Isatis), Linaceae (Linum) and plants in several other families; e.g. Kadyrbekov (2014d) records it (as A. bidentis ssp. montanum) from Kazakhstan on three genera of Campanulaceae (Adenophora, Campanula, Codonopsis). In southern Europe, Middle East, Central Asia, Pakistan (Naumann-Etienne & Remaudière 1995), south Siberia, China (as A. linisuctum) and Africa. The African populations are found on a very similar range of hosts to the Eurasian ones, but tend to have longer, more slender siphunculi, and were described as a distinct species, A. bidentis Eastop, but this seems only to be a geographical variant of A. ilka (Eastop & Blackman 2005), as also may be A. mordvilkoi Nevsky, described from Linum usitatissimum in Central Asia, and A. papaverisuctum (q.v.). A form described as a subspecies, A. ilka ssp. montanum Kadyrbekov occurs in mountainous regions of Kazakhstan above 1500m on a variety of host plants (Kadyrbekov 2005e, as A. bidentis ssp. montanum); the observed differences might be environmentally determined. Other possible members of this group are A. papaverinum Nevsky ex Pek and A. titovi Mordvilko, both recorded from Papaver sominiferum in Central Asia but neither formally described.

Acyrthosiphon kapustjanae Pashchenko Apterae are green in life; BL 2.7-2.8 mm (2 specimens). Described from Salix udensis in Kamchatka, east Siberia (Pashchenko 2005). Further collections are necessary to determine whether this is the normal host. This species is very similar to A. dauricum.

Acyrthosiphon knechteli (Börner) Apterae are green or red; BL 2.0-3.0 mm. On undersides of leaves and on young shoots of Vaccinium uliginosum in northern and central Europe and west Siberia. Monoecious holocyclic with oviparae and alate males at end of August (Heikinheimo 1990), and apparently specific to V. uliginosum.

However, in north-west Russia, Stekolshchikov (2017) and Stekolshchikov & Khureva (2020) have reported populations of an aphid that differed insignificantly in its morphology from A. knechteli, but was found feeding on eight plant species in six families. Further research is needed to clarify the taxonomic status of this form (“Acyrthosiphon sp. aff. knechteli”).

Acyrthosiphon kondoi Shinji Plate 23b Apterae are bluish green (see Influential Points/Gallery); BL 2.1-2.9 mm. Alatae have a darker brown thorax than A. pisum. On stems and leaves of Leguminosae/Fabaceae, mainly of tribes Trifoleae (Medicago, Melilotus, Trifolium) and Loteae (Dorycnium, Lotus) but also on Astragalus (Galegeae), and also recorded from Pisum, Vicia and Lens (Vicieae). An important pest of alfalfa (Blackman & Eastop 2000), now widely distributed in Asia, North and South America, South Africa, Australia and New Zealand, with distribution more recently extended to the Mediterranean region (Turkey, Kök & Özdemir 2021, Greece, Tsitsipis et al. 2007 and Corsica, Nieto-Nafría et al. 2007). A record from an unlikely host Fagonia in Algeria (Laamari et al. 2013) needs further confirmation. Monoecious holocyclic in Japan (Kawada 1992), apparently anholocyclic in most other places. 2n=10.

Acyrthosiphon (Tlja) lactucae (Passerini) Apterae are pale yellowish green or pink, with pale grey wax bloom (see Influential Points/Gallery); BL 1.7-2.9 mm. On stems and undersides of leaves of Lactuca spp. Europe, Middle East, Kazakhstan (Kadyrbekov 2014d), Pakistan (Naumann-Etienne & Remaudière 1995), introduced to North America and more recently recorded from Argentina (Mier Durante et al. 2011) and Chile (Nieto Nafría et al. 2018). Monoecious holocyclic, with alate males. 2n=16. A. scariolae (q.v.) is probably a distinct species with 2n=18, but further work is necessary to confirm this.

Acyrthosiphon (Xanthomyzus) lambersi Leclant & Remaudière Apterae are yellowish green, lightly wax-dusted on abdomen; BL 1.4-2.7 mm. On Glaucium spp. in southern Europe, North Africa and the Middle East. Anholocyclic in France according to original description.

Acyrthosiphon leleji Pashchenko Apterae are green; BL 2.3-2.5 mm. On shoot apices and leaves of Spiraea salicifolia and Spiraea sp. in east Siberia (Pashchenko 2005). The alatae and life cycle are unknown. It is not clear how this species differs from A. ignotum.

Acyrthosiphon leonurae Pashchenko Apterae are pale green; BL 2.6-2.7 mm. Occurring singly on undersides of leaves and on apical shoots of a species of Leonurus in east Siberia (Pashchenko 2005). The alatae and life cycle are unknown. Perhaps this species should be placed in Aulacorthum.

Acyrthosiphon lobkovae Pashchenko Colour of apterae in life is unknown; BL 2.9-3.2 mm. On Myrica tomentosa (= M. gale) in east Siberia (Kamchatka). An ovipara was collected in September (Pashchenko 2005). An ovipara collected at the same time and from the same host differed in having relatively shorter ANT PT and longer siphunculi, and was described as another species, A. myricae Pashchenko.

Acyrthosiphon loti (Theobald) Apterae are green, frequently with faint greyish transverse stripes (see influentialpoints.com/Gallery), or more rarely pink; BL 1.7-2.9 mm. On Lotus and Dorycnium spp., Anthyllis vulneraria, Hippocrepis comosa, Onobrychis sativa (= viciifolia), and sometimes on certain other herbaceous Leguminosae, including Medicago sativa. In Europe, eastward to west Siberia, Turkey and Iran, and introduced to South America (Argentina: Ortego & Mier Durante 2003; Chile: Nieto Nafría et al. 2018). Monoecious holocyclic, with both apterous and alate males. 2n=10.

Acyrthosiphon macrosiphum (Wilson) Yellow-green in life, with darker antennae and tibiae (see aphidtrek.org); 1.5-1.9 mm. Holocyclic, with alate males, on leaves of Amelanchier spp. in USA (California, Colorado, Utah, Oregon, Montana) and Canada (British Columbia, Saskatchewan). Sambucus canadensis is also listed as a host, but specimens have not been seen from that plant. References: Palmer 1952, Eastop 1971. 2n=10. A photo of a live fundatrix is available at http://www.flickr.com/photos/sandnine/4545416151.

Acyrthosiphon malvae (Mosley) Plate 23a (Fig.30c) Apterae are green, yellowish or greyish green, or pinkish red (see influentialpoints.com/Gallery); BL 1.5-3.2 mm. Apterae have 1-24 secondary rhinaria on ANT III, alatae have 12-31. On many plants, but particularly herbaceous Rosaceae. This is a complex taxon, in which several forms with more specific host plant associations are recognised as subspecies, although slide-mounted specimens cannot usually be identified to subspecies level unless large samples are available. Populations of pale green or red aphids on many Geraniaceae (Geranium, Pelargonium, Erodium) and Malvaceae (Alcea, Malva), as well as many plants in other families, are regarded as A. malvae s. str., and are of almost world-wide distribution, with anholocycly in warmer climates and indoor populations. A. malvae ssp. agrimoniae (Börner) is yellowish green and found in flowerheads or on undersides of leaves of Agrimonia spp. in Europe and western and central Asia. A. malvae ssp. pentaphylloides is green and described from Pentaphylloides (=Potentilla) fruticosa in east Siberia (Pashchenko 2005; probably this is a synonym of A. boreale), A. malvae ssp. poterii Prior & Stroyan is bright salmon pink, yellowish or green and found on Poterium sanguisorba (= Sanguisorba minor), and is only known from England. A. malvae ssp. potha Börner is pale yellowish or greyish green and associated with Alchemilla spp., and possibly also Acaena spp. and Geum rivale, throughout Europe. Records of this subspecies from Potentilla are possibly all referable to the very similar A. boreale (q.v.). A. malvae ssp. rogersii (Theobald) is green or yellow-green, often shiny, and may form large colonies on young leaves of Fragaria in north and west Europe (Blackman & Eastop 2000). Populations in the BMNH collection from England and Switzerland with specificity to Geranium robertianum may also qualify for at least subspecies status.The recognised subspecies are apparently all monoecious holocyclic on their respective host plants; both apterous and alate males occur in this group, and occasional hybridisation probably occurs between host-specific and more polyphagous populations, further confusing the taxonomic situation (Prior & Stroyan 1964; Müller 1972). 2n=10 (incl. ssp. poterii, rogersii).

Acyrthosiphon matilei Remaudière & Leclant Apterae are green, BL 1.7-2.4 mm. Living all year round on Euphorbia spinosa and Eu. flavicoma ssp. occidentalis. In the south of France, Italy, Spain and Croatia. One ovipara was found in southern France in November (Remaudière & Leclant 2000), but Spanish populations are habitually anholocyclic with large overwintering populations, and can live even under snow at high altitudes (Nieto Nafría & Mier Durante 2013). The description is very close to that of A. pareuphorbiae on Euphorbia in China.

Acyrthosiphon moltshanovi Mordvilko Apterae are pale yellow or greenish; BL 2.4-3.7 mm. Described from an unidentified plant in Tajikistan, and subsequently reported from various plants in Pakistan and India; the true host is uncertain but is possibly Selinum wallichianum (= tenuifolium; David & Hameed 1975).

Acyrthosiphon nigripes Hille Ris Lambers Apterae are green with black appendages; BL 2.8-4.0 mm. On Laserpitium siler in south and central Europe (Austria, Italy, Hungary). Populations on Peucedanum are regarded as a subspecies, A. nigripes ssp. peucedani (Bohzko); this was described as shiny brown by Szelegiewicz (1967b; specimens from P. officinale), but bright shiny green in the original description (from P. ruthenicum). Another possibly subspecific form occurs on Seseli; this was described and compared with A. nigripes s. str. by Pintera (1957), under the name A. superba Börner (which is a synonym of nigripes). The name A. nigripes ssp. blattnyi Pintera ex Szelegiewicz is applicable to this form.

Acyrthosiphon ononis (Koch) Apterae are green or red; BL 2.7-3.9 mm. On Ononis spp. in Europe. Monoecious holocyclic with apterous males (Meier 1958). A close relative of A. pisum, and often regarded as a subspecies.

Acyrthosiphon papaverisuctum (Zhang, Chen, Zhong & Li) Apterae are ivory white; BL 2.3-2.8 mm. On Papaver sominiferum in Gansu province, China (Zhang 1999, as Impatientinum). Perhaps a geographical variant of A. ilka.

Acyrthosiphon pareuphorbiae Zhang Colour of apterae in life is unrecorded; BL c.2.5 mm. On Euphorbia sp. in Yunnan province, China.

Acyrthosiphon parvum Börner Apterae are green to dark green, sometimes with dusky antennae, legs and siphunculi; BL 1.9-2.5 mm. On young growth, and later on undersides of leaves, of Cytisus, Cytisophyllum, Anthyllis and Genista in eastern, central and southern Europe (Poland, Hungary, Bulgaria, Czech Republic, Austria, Switzerland, Italy, Majorca).

Acyrthosiphon pentatrichopus Hille Ris Lambers Colour of apterae in life unknown; BL 3.0-3.4 mm. Alatae are undescribed. On Rosa fendleri and Rosa sp. in western USA (Colorado, Utah). Fundatrices were collected on R. fendleri in May (Hille Ris Lambers 1974b).

Acyrthosiphon phaseoli Chakrabarti, A.K.Ghosh & Raychaudhuri Apterae are dull white to slightly greyish; BL 2.7-3.0 mm. On undersides of leaves of a Phaseolus sp. in India (HP). Alatae are undescribed. A. pisivorum, described from Pisum sativum in China, is very similar, and may be a synonym (2n=14 for pisivorum; Chen & Zhang 1985a). [If Macrocaudus phaseoli Shinji, described from Phaseolus in Japan is rediscovered and proves to be an Acyrthosiphon, then this species (if not a synonym) will require a replacement name.]

Acyrthosiphon pilosum Nieto Nafría, Aldea & Castro Apterae are green; BL 2.4-3.7 mm. On Ononis spp. in Mediterranean coastal areas of France and Spain. Oviparae were collected in mid-October (Nieto Nafría et al. 2015b).

Acyrthosiphon pisum (Harris) Pea Aphid Plate 23d Apterae are green or pink, with antennae dark at apices of segments, often lightly wax-dusted (see influentialpoints.com/Gallery); BL 2.5-4.4 (-5.5) mm. On young growth and developing pods of many herbaceous and some shrubby or woody Leguminosae/Fabaceae, and an important pest of peas and alfalfa (Blackman & Eastop 2000, p. 218). Trees recorded from are Albizia julibrissin, Robinia pseudoacacia and Sesbania grandiflora. Certain other plants (Capsella bursapastoris, Reseda spp.) may be colonised under dry conditions. A. pisum is almost cosmopolitan. It is monoecious holocyclic in temperate regions, producing both apterous and alate males in differing proportions that are dependent on host race (Frantz et al. 2010). In Europe and Central Asia, there is a complex of races and subspecies with different host ranges and preferences (Müller 1985), more than one of which has been introduced to other parts of the world, and populations on different host plants (e.g. Vicia, Trifolium, Medicago) have diverged genetically to different extents (Hawthorne & Via 2001; Simon et al. 2003; Blackman & Eastop 2007; Peccoud et al 2008, 2009a,b, 2014; Peccoud & Simon 2010; Nouhaud et al. 2014; Duvaux et al. 2015; Eyres et al. 2017). Recent work has shown that several of the “host races” are reproductively isolated, and that diversification of these occurred much earlier than previously thought, predating Neolithic agriculture (Fazalova et al. 2018; Fazalova & Nevado 2020) The A. pisum complex harbours a rich community of secondary facultative symbionts, and the roles of these in relation to genetic diversity, host specialisation and resistance to parasitism have been the subject of extensive research in Europe and North America (e.g. Ferrari et al. 2011; Russell et al. 2013).

The form usually found on Ononis is here judged to be sufficiently distinct to warrant separate species status (see A. ononis). Molecular studies have singled out other host-specific forms which qualify for separate species or at least subspecies status, on Cytisus scoparius (and probably other members of the tribe Genisteae including Spartium junceum), and on Lathyrus pratensis (Peccoud et al. 2009a,b). Available names for these are respectively spartii (Koch) and lathyri (Mosley). A. pisum is currently being studied intensively as the species selected for the aphid genome project (Brisson & Stern 2006, Tagu et al. 2010); a draft of its genome sequence has been published (International Aphid Genomics Consortium 2010) Also see Y. Li et al. 2019, 2020 and Mathers et al. (2021) for recent research involving chromosome-level genome assembly and comparisons with Myzus persicae and Rhopalosiphum maidis). 2n=8.

Acyrthosiphon porrifolii (Börner) Apterae are presumably yellow-green; BL 1.5-1.8 mm. On Hieracium porrifolium in Austria. Males are alate (Börner, 1950).

Acyrthosiphon primulae (Theobald) Apterae are pale yellow or greenish yellow, shiny, with apices of antennae and legs dark (see influentialpoints.com/Gallery); BL 2.0-2.5 mm. Alatae have dark dorsal cross-bands and postsiphuncular sclerites. On undersides of leaves of Primula spp. in Europe, and introduced to New Zealand (Cottier 1953, as Aulacorthum), Tasmania (Heie 1994, as Microlophium) and California (BMNH collection, leg. R.C. Dickson). The generic position of this species is uncertain. 2n=16.

Acyrthosiphon pseudodirhodum (Patch) Apterae are pale cream to white; BL 2.2-3.5 mm. On various Rosaceae (Geum, Potentilla, Rosa, Spiraea) in north-eastern USA and Canada. MacGillivray (1968) redescribed this species. Two other undescribed, somewhat similar spp. occur in western North America, one on Chamaebataria millefolium in California (BMNH collection, leg. P. Rude) and the other on Holodiscus discolor (California, BMNH collection, leg. C. Lagace) and Cercocarpus parvifolius (Colorado; BMNH collection, leg. F.C. Hottes & D. Hille Ris Lambers). Both have R IV+V 0.9-1.0 × HT II (0.72-0.82 in pseudodirhodum), 4-6 accessory hairs on R IV (2 in pseudodirhodum), and longer hairs (46-50 μm) on abdominal tergite 3 (14-20 μm in pseudodirhodum). The Chamaebataria aphid has siphunculi 0.18-0.2 of body length , whereas the aphid on Holodiscus and Cercocarpus has very thick siphunculi 0.125-0.15 of body length and only 2.5 –3.6 times longer than their width at midlength (6.0-7.9 × longer in pseudodirhodum and 5.3-5.8 times longer in the Chamaebataria aphid). Life cycles of all three species are unknown, but the sample from Colorado on Cercocarpus included alate males.

Acyrthosiphon purshiae (Palmer) Apterae are mid-green, shining or slightly frosted, with dark appendages; BL 2.0-2.5 mm. On leaves and stems of Purshia tridentata in western USA (Palmer 1952). [This species was described in Macrosiphum, and DNA analysis by Foottit et al. (2008) suggests that it should probably be returned to that genus.]

Acyrthosiphon ranunculum L.K.Ghosh Apterae are yellowish white, the antennae ringed with black, legs mainly pale, siphunculi brown-black , cauda pale; BL 2.2-2.6 mm. On stems, petioles and undersides of leaves of Thalictrum pauciflorum (= dioicum) in Himachal Pradesh, India.

Acyrthosiphon rockspirea Jensen Apterae are mid-green; BL 1.5-2.7 mm. living at low densities on Holodiscus spp in western USA. Monoecious holocyclic, with oviparae and alate males in September; Jensen (2021) described all morphs. This is the species on Holodiscus in California, and also on Cercocarpus in Colorado, referred to as Acyrthosiphon sp. nr. pseudodirhodum in Blackman & Eastop (2006). The record from Cercocarpus is probably based on a misidentification of the host plant (Jensen 2021).

Acyrthosiphon rubi Narzikulov (= Metopolophium sonchifoliae Raychaudhuri, L.K. Ghosh & Das) Apterae are shining green; BL 1.6-2.9 mm. Alatae have 18-34 rhinaria on ANT III. On Rubus spp. and Fragaria sp., but apparently with polyphagous tendencies, because there are also rather numerous records from plants outside Rosaceae (Catamixis, Desmodium, Fagopyrum, Hieracium, Polygonum, Rumex, Sonchus), although at least some of these are probably misidentifications. In mountainous regions of Tajikistan, Nepal (BMNH collection, leg. K.C. Sharma) and northern India. Oviparae and alate males were described (on Rubus) from Himachal Pradesh, India by Raychaudhuri et al. (1980). The specimens from Rubus ellipticus in northern India (Uttar Pradesh) described by Stroyan & Nagaich (1964) as ssp. elliptici, are smaller and have a less rugose dorsal cuticle, but other specimens from this host in Nepal are typical rubi. 2n=12.

Acyrthosiphon rubifoliae (Raychaudhuri, M.R. Ghosh & R.C. Basu) Apterae are pinkish green; BL 3.3-4.2 mm. Alatae have c.22 secondary rhinaria on ANT III and faint brown dorsal markings. Described from Rubus ellipticus (as a Metopolophium), and subsequently also recorded from Potentilla nepalensis (Raychaudhuri, L.K.Ghosh & Das 1980). In northern India (Himachal Pradesh, Nagaland, Sikkim).

Acyrthosiphon rumicis Narzykulov in Narzikulov & Umarov Apterae are shining grass-green, BL c.4.7-4.8 mm. In large colonies on stems and undersides of leaves of Rumex spp. in Tajikistan.

Acyrthosiphon sanguisorbae Seccombe Colour of aptera in life is unrecorded; BL 1.8-2.9 mm. On Poterium sanguisorba (= Sanguisorba minor) in Afghanistan. Alatae have 20-22 secondary rhinaria on ANT III. Placed as a subspecies of A. malvae by Remaudière & Remaudière (1997), but with several features that justify its status as a full species.

Acyrthosiphon saussureae Pashchenko Colour of aptera pale green; BL 2.5-3.0 mm. On a species of Saussurea, feeding on undersides of leaves and readily falling from plant when disturbed (Pashchenko 2005). Other morphs and life cycle are unknown. In east Siberia (Kamchatka).

Acyrthosiphon scalare (Richards) Apterae are pale green or yellow; BL 1.7-1.9 mm. Other morphs are unknown. On Potentilla fruticosa in Ontario, Canada (as Chaetosiphon).

Acyrthosiphon (Tlja) scariolae Nevsky Apterae are pale yellow-green, finely wax-powdered; BL 2.2-2.8 mm. Described from Lactuca serriola (= scariola) in Central Asia, and specimens assigned to this species have subsequently been collected on Lactuca spp. (saligna, serriola, virosa) and Sonchus arvensis in Europe, Middle East and North America (Canada), but further work is needed to confirm the distinction from A. lactucae (q.v.). 2n=18.

Acyrthosiphon shinanonum Miyazaki Apterae are green with black apices to antennae, femora and tibiae; BL 2.5-3.1 mm. On Geum calthifolium (var. nipponicum) in Japan.

Acyrthosiphon soldatovi Mordvilko Apterae are green, with dark antennae and dusky yellow, black-tipped siphunculi; BL 1.9-3.2 mm. On Spiraea spp., especially salicifolia, in east Siberia. Also in north India (Simla: BMNH collection, leg. K. Narayanan), Kazakhstan (Kadyrbekov 2009a) and Tajikistan (Narzikulov & Umarov 1969); the latter population was monoecious holocyclic with alate males on S. hypericifolia, and regarded as a subspecies, A. soldatovi tadzhikistanicum.

Acyrthosiphon sophorae Narzikulov & Umarov Apterae are rather large (about 2.4 mm), pale green with waxy bloom, on Sophora spp. (alopecuroides, japonica, mollis) in Kazakhstan, Tajikistan and Pakistan. The life cycle is unknown. References: Narzikulov & Umarov 1969, Eastop 1971, Naumann-Etienne & Remaudière 1995.

Acyrthosiphon supranubius Carnero & Nieto Nafría Apterae are powder green with antennae, legs, siphunculi and cauda darker; BL 2.9-3.5 mm. On Cytisus supranubius, only known from the Canary Islands (Carnero & Nieto Nafría 1995). The life cycle is unknown. [This species was regarded as a possible synonym of A. parvum by Eastop & Blackman (2006), but is now distinguished from that species in the key to apterae on Cytisus.]

Acyrthosiphon svalbardicum Heikinheimo Apterae are very variable in colour, pale pink, yellow, green or brown with darker dorsal markings; BL 1.2-1.9 mm. On Dryas octopetala in Spitzbergen. Monoecious holocyclic with oviparae and apterous males in early August (Strathdee et al. 1993). The alata was described by Simon et al. (2008). Chaubet et al. (2013) described two new aphidiine parasitoids from this aphid, one of them specific to it.

Acyrthosiphon thracicum Tashev Apterae are green, BL 2.7-3.2 mm. On Euphorbia platyphyllos in Bulgaria, and also recorded from Czech Republic on E. polychroma (Holman 1965). Remaudière & Leclant (2000) compared it with the closely related A. euphorbiae.

Acyrthosiphon umarovi Narzikulov Apterae are green; BL 2.9-3.8 mm. On upper parts of stems of Astragalus sp. in Afghanistan. Alata is undescribed. It seems probable that this is a synonym of Acyrthosiphon gossypii.

Acyrthosiphon vandenboschi Hille Ris Lambers Colour of apterae in life is unknown; BL 3.7-4.4 mm. Alatae have 42-60 secondary rhinaria on ANT III. On Potentilla glandulosa in California, USA.

Acyrthosiphon vasiljevi Mordvilko Apterae are green; BL not recorded. On leaves of Cucurbita pepo in Central Asia. Similar to A. caraganae according to the original description, which does not provide enough information to include it in a key. Aphids identified as A. vasiljevi were collected on Sonchus asper by Rusanova (1942).

Acyrthosiphon wasintae (Hottes) Apterae are pale dull green; BL 1.4-1.7 mm. Living singly on undersides of leaves of Potentilla fruticosa in western USA (Colorado, Utah, Idaho, Wyoming, Montana, Oregon) and Canada (Alberta; BMNH collection, leg. D. Hille Ris Lambers). Monoecious holocyclic, with alate males (Palmer 1952).

Adelges Vallot

Adelgidae

Distinguished by having five pairs of abdominal spiracles, whereas members of the other genus of Adelgidae, Pineus, have only four distinct pairs. As in Pineus the primary host of holocylic species is Picea, but the secondary hosts are Abies, Larix, Pseudotsuga and Tsuga, never Pinus. The galls on Picea are cone-like, often resembling miniature pineapples. The complete life cycle of holocyclic species takes two years (fig. 1), and involves up to seven different morphs. Several species are anholocyclic with a reduced number of morphs and no host alternation, either living on Picea (in which case a gall may or may not be formed by a “pseudofundatrix”), or living on what was the original secondary host.

The name Chermes was once commonly applied to Adelges, but is now suppressed (Eastop 1963). Some authors have used Adelges in a narrow sense for the A. laricis group, and placed other species in Aphrastasia, Dreyfusia, Cholodkovskya, Gilletteella and Sacchiphantes, but this generic classification is based on rather slight differences, and we use Adelges in its broader sense to cover all these groups. Recent molecular work (Havill et al., 2007) provides some support for evolution in association with secondary host genera, with species migrating to Abies clustering separately from those going to Larix and Pseudotsuga, and the species associated with Tsuga apparently forming a third group. Havelka et al. (2019, 2021) analysed mitochondrial (COI) and nuclear (EF-1α) sequences of the species in Lithuania, showed that there was cophylogeny with their symbionts, and discussed the taxonomic implications of their findings. Favret et al. (2015b) provided a taxonomic and nomenclatural catalogue of the world fauna. Carter (1971) provided an account of the British species, and Carter (1976) keyed the galls of British Adelgidae on Sitka spruce. Binazzi (2000) keyed the winged morphs of Italian species. Other taxonomic accounts are by Börner (1908), Börner & Heinze (1957), Annand (1928: North America), Heinze (1962: central Europe), Binazzi & Covassi (1991: subgenus Dreyfusia in Italy), Inouye (1953: Japan) and A.K. Ghosh (1984a: India). Havill & Foottit (2007) and Havill et al. (2007) reviewed adelgid ecology and evolution, and Sano & Ozaki (2012) discussed the evolution of adelgid life cycles. Foottit et al. (2009a) and Žurovcová et al. (2010) evaluated the use of DNA bar-coding of the mitochondrial COI gene for clarifying adelgid taxonomy.

Adelges (Sacchiphantes) abietis (L.) is anholocyclic on Picea spp. (abies, glauca, jezoensis, koyamai, sitchensis), with only two generations per year; a pseudofundatrix that induces a pineapple-like gall (fig. 123A, and see also influentialpoints.com/Gallery), and yellow to yellow-green alate gallicolae that leave the gall in late summer but do not generally disperse far, often laying eggs on the same or neighbouring spruce trees. The galls are hard, green, and covered with trichomes; Rohfritsch (1990) studied their development in detail. Ewert (1967) found differential levels of infestation by A. abietis galls among spruce clones, Mitchell & Maksymov (1977) studied predators, Flaherty et al. (2010) and Pilichowski et al. (2014) studied factors influencing gall size and distribution and development of gallicolae, and Lasota et al. (1983) described a heavy fungal attack on the galls. Found throughout Europe, and in North Africa (Morocco), Central Asia (Kirghizia; Kadyrbekov 2013c), India and North America. Japanese records of abietis should probably be referred to A japonicus, and most records from Larix are probably referable to A. viridis. 2n=18 (Steffan 1968).

Adelges (Gilletteella) cooleyi (Gillette) typically host-alternates between Picea spp. (engelmanni, pungens, sitchensis) and Pseudotsuga spp. (macrocarpa, menziesii) (fig. 64G). The galls on shoot tips of Picea are characteristically elongate, with green, pink or red gall chambers and long needles protruding from them (fig. 123B, and see also bladmineerders.nl/gallen). Apterous exules on needles of Pseudotsuga are covered in white woolly wax (see influentialpoints.com/Gallery). Native to western North America, but now occurring throughout Europe and North America, and in Tasmania (BMNH collection, leg. V.F. Eastop). Cumming (1962) described a monomorphic, non-galling, anholocyclic form on Picea glauca in Canada. An anholocyclic form also occurs on Pseudotsuga in both Europe and California, to which Steffan (1970) applied the name coweni Gillette; see under that name for further information. Parry (1978a,b; 1980) and Parry & Spiers (1982) have made extensive studies of the population biology of A. cooleyi in Scotland, Teucher (1955) studied natural enemies, and Stephan (1987) reported differences in resistance of Pseudotsuga provenances in Germany. Ahern et al. (2008) compared genetic variation and secondary host preferences of native western and introduced eastern populations of A. cooleyi in North America, and Ahern et al. (2009) studied genetic variation in the indigenous western populations. 2n=22 (Steffan 1968).

Adelges (Gilletteella) coweni (Gillette) Anholocyclic, living all year on Pseudotsuga menziesii. Alate sexuparae develop in spring, but these are all gynoparae and produce only sexual females, so that without males the holocycle is defunct. Described from California, and introduced to southern Europe. Roversi & Binazzi (1996) provided a full account of the life cycle in central Italy, where it has become a serious pest in stands of Douglas Fir.

Adelges (Gilletteella?) glandulae Zhang Host-alternating between Picea spp. (brachytyla, likiangensis, purpurea) and Abies ?faberi in Sichuan and Yunnan provinces, China (Zhang et al. 1980, Chen & Qiao 2012a). The galls are ovate, cone-like, without associated needles; the form of the gall as well as DNA analysis (Havill et al. 2007) indicate that this species is wrongly placed in subgenus Gilletteella. The hiemosistens morph on Abies overwinters in second instar. This species is described as anholocyclic, presumably in error as it is both gall-forming and host-alternating. The fundatrix/?pseudofundatrix is undescribed.

Adelges (Dreyfusia) joshii Schneider-Orelli & Schneider (fig. 16G) Anholocyclic, with three generations per year (hiemosistens and two aestivosistens) on Abies pindrow in western Himalaya. Adults are dark brown, 0.7-1.2 mm, covered with wax. Ghani & Rao (1966) gave a detailed account of the morphology and life cycle.

Adelges (Sacchiphantes) karafutonis Kono & Inouye Galls of this species on young shoots of Picea jezoensis are small, with only 5-15 scales (Inouye 1953). Only the gallicolae are described, and the secondary host (presumably a Larix species), is unknown. For differences from the closely related A. torii see Eichhorn & Carter (1978). Recorded from Sakhalin and Japan (Hokkaido).

Adelges (Sacciphantes) kitamiensis (Inouye) Described from secondary host morphs attacking Larix kaempferi in Hokkaido, Japan, sometimes so heavily that the entire trunk of a 10-20 year-old tree is covered in its white wax wool. Hiemosistens immatures and adults are dirty blue in colour (cf. viridis). For further details and differences from viridis see Inouye (1963). The small (about 0.6-0.7 mm) sexuparae maturing in May-June leave Larix for a previously unknown primary host; no galls have been found on Picea abies, even where this tree occurs in mixed stands with L. kaempferi. However, the probable primary host generations of this species have now been identified, producing pineapple galls on three Picea spp. in Central Japan, and with a DNA barcode listed as that of “Adelges sp. A” in Foottit et al. (2009). Alate gallicolae (1.5-2.5 mm) emerging from these galls deposited egg masses in the laboratory on L. kaempferi. The first instars hatching from these eggs agreed morphologically with the syntypes and description of S. kitamiensis, but were yellow in colour (Sano et al. 2011). Immature and adult fundatrices from galls have not yet been described.

Adelges (Dreyfusia) knucheli Schneider-Orelli & Schneider (fig. 16F) Host-alternating between Picea smithiana and Abies spp. (pindrow, spectabilis) in western Himalaya (Pakistan and Kashmir), or anholocyclic on Abies pindrow where the primary host is absent, e.g. in Murree Hills of Pakistan. The pineapple or cone-like galls on P. smithiana vary considerably in size and shape. The sistens generations on Abies pindrow are dark brown with wax filaments radiating a star-like fashion, and occur on main stems of older firs as well as on branches of younger trees. Progrediens apterae and alate sexuparae develop on new-grown needles, the latter without secreting wax. Ghani & Rao (1966) provided detailed descriptions of all stages and morphs. Predators were discussed by Pschorn-Walcher (1964). [Stebbing’s (1910) account of Chermes himalayensis probably applies to this adelgid; but see Schneider-Orelli & Schneider (1954).]

Adelges laricis Vallot (=strobilobius Kaltenbach) Plate 1a, 1b The oldest-named member of a taxonomically very difficult complex of adelgids which have host alternation between Picea and Larix, or have an incomplete cycle restricted to either the primary or the secondary host. Galls of laricis sensu stricto on Picea are typical of the group (fig. 123D, and see also bladmineerders.nl/gallen); they are globular or ovate, with a cream or ivory coloration and waxy texture, becoming pink or brown just prior to opening. They may be terminal, or have some growth of the shoot beyond the gall. Gall development was described in detail by Rohfritsch (1990). Predation within galls by dipterous larvae was described by Mitchell & Maksymov (1977). Gallicolae of A. laricis (fig. 64F) migrate to Larix in June-July; they are 1-2 mm, blackish, and secrete little or no wax. The hiemosistens generation on Larix in spring is dark grey without wax, but the progrediens apterae left after departure of the sexuparae in June-September produce abundant wax and honeydew; these deposits from a heavy infestation cause the foliage to turn blue (Börner & Heinze 1957; Carter 1971). See influentialpoints.com/Gallery for further information and photographs. A. laricis sensu stricto occurs in Europe and North America and is recorded from Picea abies, koraiensis, mariana, rubens, sitchensis (primary hosts), and from Larix decidua, ×eurolepis, kaempferi, laricina (secondary hosts). 2n=20 (Steffan 1968).

Li & Ts’ai (1973) provided a very detailed account of an adelgid under the name A. laricis in north-east China, with primary host Picea obovata. Numerous other names have been applied to members of the laricis group, some of which were only described from one or two morphs and are possibly not distinct species:

aenigmaticus Annand Only the apterous ?progrediens morph is described by Annand (1928). It was collected on Larix laricina in Maine, USA, and distinguished from laricis by its smaller, rounded dorsal sclerites bearing round, separate wax pores, and by the short and thick third antennal segment. Similar specimens were collected by F.W. Quednau on the same host in Quebec, Canada (F.W. Quednau, pers. comm. 1971). Life cycle is unknown.

diversis Annand Progrediens apterae collected on cones of European larch (Larix decidua) in Oregon, USA were thinly covered with short wax threads (Annand 1928). Only the first instar ?sistens, progrediens and immature ?sexupara are described. The progrediens is distinguished by having no sclerites on abdominal tergites 5-8 and only small sclerites with few wax pores on the more anterior tergites. Not recorded since the original description.

geniculatus (Ratzeburg) This is the name currently applied to populations that build up large populations on Larix decidua in central Europe (Austria, Germany), causing discoloration, deformation and eventual loss of the needles and also damaging buds (Perny 2004). A. geniculatus had previously been synonymised with A. laricis, but was again given separate species status by Steffan (1968) on account of its anholocycly and apparent inability to produce alate sexuparae. However the assumption that the only populations reaching damaging levels on larch are anholocyclic and can therefore be identified as geniculatus may be unwarranted, as successive generations of progredientes of laricis persist on larch and can build up heavy infestations (Carter 1971). Further work also seems necessary to confirm that A. geniculatus is functioning as a fully independent, permanent parthenogen, and that its separate species status is justified..

isedakii Eichhorn Host-alternating between Picea jezoensis (var. hondoensis) and Larix kaempferi in Japan (Eichhorn & Carter 1978). The gall on Picea is similar to that of laricis, but larger. The gallicola has a red body and greenish wings, is smaller (1.0-1.5 mm) than the non-migratory gallicola of A. japonicus (see below), and has no wax glands on the head. Only the first instar hiemosistens (neosistens) has been described from Larix.

japonicus Monzen (fig. 64L) Anholocyclic, pseudofundatrices induce cone-like galls on shoot tips of Picea jezoensis and P. sitchensis in Japan and Sakhalin (Inouye 1953; Paschenko 1988). Gallicolae emerging from galls in August-September are large (1.9-2.8 mm) and with or without cephalic wax glands (Eichhorn & Carter 1978); some do not disperse, so that individual trees may develop heavy levels of infestation (Kamata 1986). The cause of the relatively slow larval development in the galls was studied by Tabuchi et al. (2009). Sano et al. (2008) have now shown that in northern Japan there is also a holocyclic form of this species that migrates to Larix kaempferi; they reared and described alate gallicolae and first instar exules of the migratory form, and compared the two life cycles.

karamatsui Inouye Anholocyclic on secondary hosts (Larix kaempferi, gmelinii) in Japan, Sakhalin and Korea. Sistens, apterous progrediens and sexupara are described (Inouye 1953). Sexuparae, small and dark brown to black with abundant wax, fly to Picea in Japan, but die or produce eggs which do not hatch (Eichhorn & Carter 1978); an analogous situation to that with A. tsugae on Tsuga in USA.

lapponicus (Cholodkosky) Anholocyclic, pseudofundatrices inducing galls on both native and exotic Picea spp. Dark red gallicolae about 1.6 mm long energe from galls in June-July and lay reddish eggs on needles of Picea with little or no secretion of wax. In northern and central Europe, and more recently recorded from Kirghizia (Gabrid 1981). Differential infestation of Picea spp. in northern Finland was studied by Häggman and Rousi (1986).

lariciatus Patch Host-alternating between Picea spp. (abies, glauca, mariana, pungens) and Larix spp. (laricina, lyalli) in North America (Alberta, Manitoba, Saskatchewan, Utah). Galls are globular, often sub-terminal and to one side of shoot, green when young to red and brown when mature. Cumming (1968) gave a detailed account of morphs and life cycle in Canada.

oregonensis Annand Only known from the original description of sistens and progrediens morphs from Larix occidentalis in north-western USA (Oregon, Washington, Montana). Differences from laricis were discussed by Annand (1928). It feeds on the needles and on the twigs at the needle bases. The primary host, if any, is unknown.

potaninilaricis Zhang Host-alternating between Picea retroflexa and Larix potaninii in Sichuan, China. Described as a subspecies of laricis on the basis of differences in number and distribution of wax glands (Zhang et al. 1980).

tardoides (Cholodkovsky) Host-alternating between Picea abies and Larix sibirica in north-eastern Europe. The gallicolae migrate to the secondary host in July-August, later than laricis, and unlike those of laricis, secrete abundant wax (Börner & Heinze 1957, p.346).

tardus (Dreyfus) Anholocyclic on Picea spp. throughout continental Europe. The gallicolae emerge from late June to September, later than lapponicus, and lay eggs on exposed parts of the trees under a covering of white wax. Due to non-dispersal of the gallicolae, the galls may become very numerous and can severely affect growth of young spruce trees. Baurant (1968) studied the phenology of A. tardus in relation to Picea abies in Belgium, and suggested that its synchronisation with the host might be upset by selection of appropriate spruce varieties. 2n=20 (Steffan 1968).

Adelges (Dreyfusia) merkeri Eichhorn Host-alternating between Picea orientalis and Abies spp., especially A. alba, with a rather restricted distribution in Europe (Germany, southern Sweden, one record from Austria); origins unknown. Galls on Picea resemble those of A. nordmannianae but are usually larger (7-22 mm) and open in June-July (Eichhorn 1975). On Abies, colonies are found on the trunk and crown region of older firs, or on all parts of young and seedling trees, where damage may be severe with swelling and stunting of infested branches, especially at their bases. As well as the hiemosistens there are one or even two aestivosistens generations. Adults sistentes generally have less wax wool than either nordmannianae or piceae. Pschorn-Walcher & Zwolfer (1960) provided ecological notes and Eichhorn (1968) studied population dynamics on Abies, especially the influence of natural enemies. [ Havill et al. (2020) propose on morphological and molecular evidence that A. merkeri is a recent hybrid of A. piceae and A. nordmanniae, and therefore that it is not a valid species. However, A. merkeri (like A. prelli – see below) has a viable sexual phase and consistent biological characteristics, indicating that even if originally of hybrid origin, it has been functioning for some time as a good species. It is possible that their extensive sampling of populations in Europe and North America did not include A. merkeri.]

Adelges (Dreyfusia) nordmannianae (Eckstein) (=nüsslini Börner) (fig. 10B, fig. 16C) Host-alternating between Picea orientalis and Abies spp. (alba, bornmuelleriana, cilicica, nordmanniana, etc.). Believed to be endemic and relatively uninjurious in the Caucasus and East Pontus mountains (Russia and Turkey), where there are mixed stands of Picea orientalis and Abies nordmanniana, but causing severe damage to Abies where non-endemic in Europe, North America, New Zealand (see Zondag 1982) and Tasmania. Galls on Picea (fig. 123C) are 2-15 mm long, terminal, globular, pinkish and resembling strawberries when young, later becoming greenish with red or purple coloration at bases and tips of scales. They mostly open in June (Eichhorn 1975). There is never more than one aestivosistens generation, and even this is lacking in some populations. On Abies alba there appear to be two forms of A. nordmannianae: “typica” which infests branches, twigs and needles of young trees in sunny dry locations – the attack of the progrediens generation causing severe deformation of needles and shoots and often killing the tree (see Varty 1956); and form “schneideri” which infests the trunks of older firs 30-120 years old in forests (studied by Pschorn-Walcher & Zwolfer 1958, and Eichhorn & Pschorn-Walcher 1972). The latter form would not transfer to young trees (Schneider-Orelli, Schaeffer & Wiesman 1929). Colonies of sistentes on stems are black with a grey wax bloom and only a fringe of wax wool (cf. A. piceae). Eichhorn (1969a) studied the natural enemies of A. nordmannianae in Turkey, and also worked on morph determination (Eichhorn 1969b), life cycle (Eichhorn 1973, 1991) and embryonic development (Eichhorn 1970). Most recent ecological studies of this species have been in Austria (Stürzer-Gilbert 1982; Bauer-Schmid 1983). For a review of natural enemies and potential for biological control see Ravn et al. (2013). Analysis of mitochondrial and nuclear DNA sequences failed to reveal any consistent differences between nordmannianae, piceae and prelli (Ravn et al. 2013). However, analysis of microsatellite DNA of samples from the Caucasus, Europe and North America discriminates between nordmannianae and piceae, and also provides evidence that hybridisation between these two has occurred to give sexually viable new taxa such as merkeri and prelli (Havill et al. 2020). 2n=22 (Steffan 1968).

Adelges (Aphrastasia) pectinatae (Cholodkovsky) (fig. 16A) Host-alternating between Picea (abies, obovata) and Abies spp. (alba, balsamea, concolor, etc.), in Europe (Sweden, Finland, Latvia, Ukraine) and east to Siberia. Recorded also from east Norway on 10 species of Abies, although the gall generations on Picea were not found there (Austarå 1990). The galls (fig. 123E) are compact, cone-like, dull green turning reddish-brown before opening, about 10-17 mm long. Reddish-brown gallicolae fly to Abies in June-July. Feeding on Abies is restricted to the needles. Frolowa (1924) studied cytology of egg and sperm maturation in this species. 2n=20 (Steffan 1968).

Populations alternating between Picea spp. (glehni, jezoensis, etc.) and Abies (sachalinensis, veitchii) in Japan are regarded as a separate subspecies, A. pectinatae ishiharai Inouye (Eichhorn & Carter 1978). Its morphs and life cycle are described in detail by Inouye (1953). Fang (1986) described the pest status of what is possibly this subspecies on Picea jezoensis, P. koraiensis and Abies nephrolepis in China.

An anholocyclic form in the pectinatae group, Adelges pindrowi Yaseen & Ghani, occurs on Abies pindrow in Pakistan. Information about its life cycle is provided by Yaseen & Ghani (1971).

Adelges (Dreyfusia) piceae (Ratzeburg) Balsam Woolly Aphid (fig. 16D) Anholocyclic on Abies spp. (alba, balsamea, cilicica, fraseri, etc.), with one hiemosistens and usually one aestivosistens generation. Sexual generations and galls on Picea orientalis have not been described, although there is now evidence from microsatellite DNA studies that sexual reproduction may sometimes occur if P. orientalis is available, and that hybridisation may occur with A. nordmannianae (Havill et al. 2020). Throughout Europe, and introduced to North America, and also to Chile (Fuentes-Contreras et al. 1997). In favourable conditions a second, or even a third, aestivosistens generation may occur (Pschorn-Walcher 1964), and in North America there is sometimes also a generation of progrediens apterae and alatae. On A. alba in Europe this adelgid normally infests the trunk and larger branches of older trees (30-100 years), and is of little economic importance. Introduced populations in North America, however, attack stems and buds of the crown region of Abies balsamea and A. fraseri, causing extensive injury. Young trees of A. balsamea are killed by “gout disease” (excessive shoot swellings near the buds); heavy stem attack may give rise to severe transpiration stresses by the formation of rotholz or compression wood (Balch 1952), causing crown dieback and a degrade of the timber (Carter 1971). Hain et al. (1983) studied the host’s defence reaction against stem attack. See influentialpoints.com/Gallery for illustrations and further information.

The economic importance of A. piceae has resulted in an extensive literature; Schooley & Oldford (1981) provide an annotated bibliography, and Johnson & Lyon (1988) gave a short, well-illustrated account. Biological control measures up until 1958 were summarised by McGugan & Coppel (1962). Foottit & Mackauer (1983) used morphometric studies to distinguish and key three forms of A. piceae in North America, thought to represent separate introductions. They designated these as subspecies (p. piceae, p. canadensis and p. occidentalis), although it is questionable whether the subspecies category is appropriate for anholocyclic taxa. The form known as canadensis is similar or identical to populations of piceae causing compression wood in Abies grandis in Scotland (Busby 1964). Binazzi & Covassi (1991) differentiated an anholocyclic population on Abies rosi in Italy as a new species, A. (D.) nebrodensis, which has now also been given subspecies status, as A. (D.) piceae nebrodensis (Havill et al. 2020). 2n is unknown.

Adelges (Dreyfusia) prelli (Grosmann) (fig. 16E) Host-alternating between Picea orientalis and Abies nordmanniana or (rarely) Abies cephalonica. Like A. nordmannianae, A. prelli is indigenous to Turkey (Eichhorn 1969) and introduced to central and western Europe. The galls are bluish-green, larger than those of A. nordmannianae (6-30 mm long), and open later, in July-August (Eichhorn 1975). On Abies nordmanniana in western Europe a hiemosistens and one or two aestivosistens generations occur, almost exclusively attacking the crown region of mature trees (Eichhorn 1956). The hiemosistens on young shoots is more sclerotized and has less wax than that on stems or branches (Eichhorn 1964). North American populations of A. piceae on Abies balsamea closely resemble A. prelli, which has not however been positively identified from that host (or from Abies alba). [ Havill et al. (2020) propose on morphological and molecular evidence that A. prelli is a recent hybrid of A. piceae and A. nordmanniae, and therefore that it is not a valid species. However, A. prelli (like A. merkeri – see above) has a viable sexual phase and consistent biological features, indicating that, even if originally of hybrid origin, it has been functioning for some time as a good species. It is possible that their extensive sampling of populations in Europe and North America did not include A. prelli.]

Adelges (Sacciphantes) roseigallis (Li & Tsai) Described from “red-barked Picea” (P. koraiensis?) in north-east China (Heilungkiang province), where there is host alternation to a managed species of Larix, presumably L. gmelinii (Li & Ts’ai 1973). The galls on Picea are bright red, smaller than those of A. viridis and resembling cypress cones; they are attached to bases of old branches. Alatae gallicolae are lustrous black, not powdered with wax (cf. viridis). Li & Ts’ai (1973) described other differences from A. viridis in colour and appearance in life of various stages, and in life cycle and fecundity. In Gansu province this species is reported to be anholocyclic on Picea spp. (asperata, crassifolia; Du et al. 1998, Chen & Qiao 2012a). Information is currently insufficient to include it in the key to aphids on Picea.

Adelges (Sacciphantes) segregis (Steffan) Anholocyclic on Larix decidua in central Europe, with two generations per year, one alate and the other a “pseudohiemosistens”. Closely related to A. viridis, but the overwintering first instar larva has a shorter rostrum than the neosistens of viridis (Steffan 1961a).

Adelges (Dreyfusia) todomatsui (Inouye) (fig. 16H) Anholocyclic on Abies sachalinensis (incl. var. mayriana) and A. veitchii in Hokkaido, Japan, with a hiemosistens and one or possibly two aestivosistens generations (Eichhorn & Carter 1978). No progrediens (needle-dwelling) morph has been found. Inouye (1953) observed that two biological forms occur that are indistinguishable morphologically; a stem-dwelling form on young and mature (pole-stage) trees, and a bud-dwelling form that lives under the scales of the previous year’s shoots and causes gall-like enlargement of buds. Predators were studied by Pschorn-Walcher (1964).

Adelges (Sacciphantes) torii (Eichhorn) Host-alternating between Picea jezoensis var. hondoensis and Larix kaempferi in Honshu, Japan. Galls are small and reddish, similar to those of A. karafutonis, opening in August. Eichhorn & Carter (1978) provided a detailed account.

Adelges (Annandina) tsugae Annand Hemlock Woolly Aphid (= funitecta of Blackman & Eastop 1994; see Favret et al. 2015a) Populations with host alternation between Picea and Tsuga occur in Japan and China. The galls and generations on Picea are apparently still not properly described. In Japan the primary host is Picea polita (and possibly P. jezoensis hondoensis; McClure 1996), whereas in China the primary hosts are P. likiangensis and P. brachytyla (Foottit et al. 2009a), with possible A. tsugae galls also reported on P. morrisonicola in Taiwan (Havill et al. 2016). Secondary hosts are Tsuga diversifolia and T. sieboldii in Japan, T. chinensis, T. dumosa and T. forrestii in China, and T. formosana in Taiwan (Inouye 1953; Havill et al. 2016). A. tsugae is recorded from North America, India, Japan, China and Taiwan. The introduced populations in North America are anholocyclic on both native and introduced Tsuga spp. DNA analysis has now provided evidence of separate introductions to western and eastern North America, the more recent introduction being particularly damaging to the native eastern species T. canadensis and T. caroliniana (Havill et al. 2016; and see McCarty & Addesso 2019 for a comprehensive review). McClure (1989a) described sistens and apterous progrediens morphs and also alate sexuparae from populations on T. canadensis in Connecticut, USA; the sexuparae migrated to Picea spp. and laid eggs, but sexuales failed to develop, even on Picea polita, the recorded primary host in Japan. Joseph et al. (2011) and Tobin & Turcotte (2018) reported on the phenology of A. tsugae respectively in northern Georgia and in the Central Appalachians. Analysis of mitochondrial and microsatellite DNA has shown that the lineages that are severely damaging to the two indigenous Tsuga spp. in eastern North America probably originated from southern Honshu, Japan, where the native secondary host is T. sieboldii. Other lineages, some of which had associations with particular Tsuga spp., were found elsewhere in Japan and in mainland Asia and western North America, where trees are not significantly damaged (Havill et al. 2006, 2007, 2016). Allozyme analyses have also been used to compare populations in eastern and western USA with those in China (Sánchez & Keena 2009). Evidence is accumulating that at least Chinese and Japanese populations should be regarded as distinct species. An anthocorid bug, Tetraphleps galchanoides, has been studied in China as a potential biocontrol agent (L. Li et al. 2011), and there is an ongoing biological control programme in north-western USA to find adelgid predators to use against A. tsugae (Ross et al. 2017, Rose et al. 2019).

Adelges (Cholodkovskaya) viridanus (Cholodkovsky) Anholocyclic on Larix spp. (decidua, x eurolepis, gmelinii, kaempferi, sibirica) in Europe (Sweden, UK, France, Germany, Italy, Ukraine) and east to China, Korea and Japan, although somewhat sporadic or local in occurrence. The life cycle is variable, probably depending on latitude and/or temperature. The overwintering first instar sistens is yellowish-green to green and usually under bark on the stem. The large (1.8-2.5 mm) adult sistentes, yellowish-green and surrounded by wax secretion, occur in May-June. Their eggs develop either as overwintering hiemosistentes or as alate progredientes; first instars of the latter morph are very active and move to tender young extension shoots to feed at the needle bases. The adult alate progrediens is quite large (2.0-2.6 mm), grey-green, and mostly non-migratory, laying eggs on the needles. These mostly develop as hiemosistentes but, in France and Germany (Gaumont 1954, Steffan 1964), some of them develop into a second generation of alate progredientes, and in suitable conditions these may even give rise to a third partial alate progrediens generation. 2n=24 (Steffan 1968).

In northern Russia, populations of the closely-related A. (C.) viridulus (Cholodkovsky) occur on Larix sibirica, and these seem always to have a single annual (hiemosistens) generation, no alatae being known. Its overwintering first instars have fewer wax pores on posterior abdominal segments than A. viridanus.

Adelges (Sacchiphantes) viridis (Ratzeburg) Host-alternating between Picea spp. (abies, koraiensis, orientalis, sitchensis, etc.; see Weiss 1955), and Larix spp. (decidua, x eurolepis, gmelinii, kaempferi). Steffan (1961a) made a detailed study of this adelgid in Germany (but see also Eichhorn 1989), and Li & Ts’ai (1973) described its life-cycle in north-east China.. The pineapple galls on Picea are similar to those of A. abietis but open earlier, in July or early August (see influentialpoints.com/Gallery). Miszta (1987) has studied their volatile oil composition in comparison to normal plant tissue. Immatures within the gall, and the alate gallicolae when they emerge, are reddish-yellow to brown, powdered with white wax (cf. A. abietis and A. roseigallis). Hiemosistens immatures and adults are green or brown. Sexuparae developing on Larix the following spring are pale greenish with sparse wax, and their feeding causes yellowing and kinking of the newly-grown needles (Carter 1971). The return migration to Picea is quite early, in May-June. It occurs throughout Europe, and is also recorded from China where it migrates from Picea koraiensis to Larix gmelinii (Fang et al. 1983). 2n=18 (Steffan 1968).

Aiceona Takahashi

Aiceoninae

Aiceona is often placed in the Anoeciinae, but it is not closely related to Anoecia and requires a separate subfamily. There are about 14 recognised species, all in east and south-east Asia, mostly associated with Lauraceae. Several species are described from unidentified hosts. Unusually, oviparae as well as males are alate, and they may be found in colonies at almost any time of year. Siphunculi are well-developed as hairy cones in all female morphs, but strangely absent in males. Accounts of Aiceona are available for India (M.R. Ghosh & Raychaudhuri 1973, A.K. Ghosh 1988), north-east India (Raychaudhuri et al. 1980d), China (Qiao & Zhang 2002c) and Japan (Takahashi 1960a). The genus is clearly still in need of further revision.

Aiceona actinodaphnis Takahashi Apterae are 2-3 mm, purplish-black dusted with white wax powder, living under leaves and on new growth of various Lauraceae (Actinodaphne, Cinnamomum, Lindera, Litsea, Phoebe). Alatae have dark-veined smoky wings with a pale area between the tip of the pterostigma and R_s. Sexual morphs are unknown. Recorded from China, Okinawa and Japan (Tseng & Tao 1938). A related species with paler wings, A. siamensis, is described from unidentified Lauraceae in Thailand (Takahashi 1941a).

Aiceona himalaica Miyazaki Appearance in life unrecorded, apterae 2.4-2.8 mm; cleared and mounted apterae have a pale body contrasting with the dark hind tibiae. Apterous viviparae, alate oviparae and males are described. Collected in July in Nepal, from Engelhardtia spicata (Juglandaceae), an unusual host association for this genus (Miyazaki 1977). It needs to be established whether this species is distinct from A. pseudosugii (see A.K. Ghosh, 1988)

Aiceona japonica Takahashi Apterae are about 3 mm, blackish-brown, covered with white wax powder, on leaves and young growth of Lauraceae (Actinodaphne lancifolia, Cinnamomum camphora, ?Litsea sp.) in Japan, and also recorded from Korea (Paik 1972). The wings of the alata are uniformly dusky without any clear area. Alate oviparae and males occur along with apterous and alate viviparae in May (Takahashi 1960), although Miyazaki (1977) expressed some doubt about Takahashi’s description of the ovipara of this species. The male genitalia were described and illustrated by Wieczorek et al. (2012).

Aiceona malayana Takahashi Apterae pale greenish-yellow, 2.2-2.5 mm, attacking young leaves of ?Actinodaphne sp. near Kualar Lumpur, Malaya. Alatae have wings clear or only faintly dusky. Distinguished by its short antennal hairs, this species is only known from the original collection, when it was recorded erroneously as A. osugii (Takahashi 1950,1960a). Sexual morphs and life cycle are unknown.

Aiceona (Subaiceona) manipurensis (Singh & Raychaudhuri) Apterae yellowish, BL 2.0-2.9 mm. Developing large ant-attended colonies in February to May on Persea odoratissima in Manipur, India. A.K. Ghosh (1988) provided a redescription. Sexual morphs and life cycle unknown.

Aiceona osugii Takahashi Apterae are yellowish green, with pale antennae and legs, BL about 2.5 mm. Described as feeding on undersides of leaves of an undetermined tree in Taiwan, and subsequently recorded from Lindera communis (Tao 1999). Alatae have dusky wings and dark spinal sclerites or short cross-bands on ABD TERG 1-5. Alate oviparae and males were described from Sichuan province, China, collected on Lindera sp. at the end of October (Qiao & Zhang 2002c). Alate oviparae (with spinal sclerites merged to form a broad dark abdominal spinal stripe) have also been collected in May in Taiwan on Persea sp., and (presumably vagrants) on a plant of the Amaranthaceae (BMNH collection, leg. S.E. Halbert). A. parvicornis may be a synonym.

Aiceona pallida A.K. Ghosh & Raychaudhuri Apterae are creamy yellow to dirty grey, BL 2.1-2.7 mm. In ant-attended colonies on tender shoots and undersides of young leaves which are rolled longitudinally. Alatae have pale wings with dark pterostigma. The only identified host is Machilus glaucescens (= Persea villosa) (A.K. Ghosh 1988). In West Bengal and Meghalaya, India. Sexual morphs and life cycle unknown.

Aiceona parvicornis Miyazaki Appearance in life is unknown; mounted apterae (about 2.7 mm) have dark head, prothorax, coxae and femoro-tibial joints. Alatae have dusky wings. Collected in May from Lindera pulcherrima in Nepal (Miyazaki 1977). Sexuales and life cycle are unknown. It needs to be established whether this species is distinct from A. osugii Takahashi.

Aiceona pseudosugii David, Sekhon & Bindra Apterae are 2-3 mm, pale yellow with dark hind tibiae, occurring in small colonies on undersides of young leaves, with attendant ants. Alatae have unpigmented wings. The only identified host is Persea odoratissima (David, Sekhon & Bindra 1970). Recorded from Himachal Pradesh and West Bengal, India. Sexuales and life cycle are unknown. [Differences from A. longisetosa M.R. Ghosh & Raychaudhuri, described from an unidentified plant in West Bengal and synonymised with pseudosugii by Eastop & Hille Ris Lambers (1976), are listed by A.K. Ghosh (1988). Another possible synonym is A. robustiseta (q.v.).]

Aiceona retipennis David, Narayanan & Rajasingh Apterae are 2.5-2.8 mm, yellowish-white, on undersides of leaves and young shoots (L.K. Ghosh 1972). Alatae have very dark wings with a clear patch between the pterostigma and R_s like that in A. actinodaphnis, and spinal sclerites of short cross-bands on ABD TERG 1-5. Distinguishing features of this species are the very long fine antennal hairs and short last rostral segment (R IV+V) only 0.6-0.7 times HT II. Pal & Raychaudhuri (1977) describe the alate male, collected in February. Alate oviparae (with spinal sclerites often merged between segments to give a broad longitudinal spinal stripe) have been collected in July (BMNH collection, leg. V.D. Verma). Identified hosts in the Lauraceae are Machilus gamblei (BMNH collection, 1 aptera leg. K. Narayanan), Persea sp. (Chakrabarti, Saha & Mondal 1988), and Phoebe lanceolata; the various records from plants in other families seem doubtful or due to casual occurrences. Widely distributed in northern India. 2n=18 (Khuda-Bukhsh 1980).

Aiceona robustiseta M.R. Ghosh & Raychaudhuri Apterae are 2.0-2.5 mm, creamy-yellow with a light dusting of wax, in colonies on young growth attended by ants (M.R. Ghosh & Raychaudhuri 1973). Alatae have unpigmented wings. Alate oviparae, collected in February and June, were described by Pal & Raychaudhuri (1977). A robustiseta was described from Litsea polyantha (= monopetala) in West Bengal, India. [Specimens from Machilus chekiangensis collected in Hong Kong, from Machilus gamblei (= Persea bombycina) in Assam, India and from Eurya nitida in Thailand, have been provisionally identified as this species (Martin & Lau 2011). However , apart from the lower number of secondary rhinaria in alatae, the differences from A. pseudosugii, a pale species with dark hind tibiae described from Persea odoratissima, are minimal.]

Aiceona titabarensis (Raychaudhuri & A.K. Ghosh) Plate 5c, d Apterae are 2.1-2.7 mm, dirty brownish to grey, dusted with wax powder, often heavily infesting the undersides of leaves and young shoots of Litsea polyantha (Basu & Hille Ris Lambers 1968), with or without attendant ants. Alatae have smoky wings as in A. actinodaphnis and A. retipennis, but the clear area adjacent to the pterostigma is narrower and less clearly defined. The species was originally described (in Lachnus) from Heteropanax fragrans (Araliaceae), a host which is unusual enough to need confirmation. Alate oviparae can be found in colonies with apterous and alate viviparae at least from June to December (M.R. Ghosh & Raychaudhuri 1973, as A. litseae). Recorded from Sikkim and West Bengal, India, and from Hong Kong and Vietnam (BMNH collection).

Akkaia Takahashi

Aphidinae: Macrosiphini

An east Asian genus with Phorodon-like projections on the antennal tubercles, but the head is not spiculose, the spiracular apertures on abdominal segments 1-5 are remarkably small, there are typically paired processes on abdominal tergite 7 and a larger median one on tergite 8, the subgenital plate has a posterior projection, and the cauda has a characteristic knobbed shape. Alatae look very different from apterae and have numerous tuberculate secondary rhinaria, peculiar hypha-like projections associated with the primary rhinaria, and much thinner, clavate siphunculi. Most species are only known from Polygonaceae, but one is known to have a sexual phase on Enkianthus. M.R. Ghosh et al. (1976) reviewed the species in east India, the Japanese species were reviewed by Takahashi (1961d) and Miyazaki (1971), and the Chinese species were reviewed by Qiao et al. (2006c).

Akkaia bengalensis A.N. Basu (Fig.42u) Apterae are orange with black siphunculi; BL 2.1-2.5 mm. Alatae have secondary rhinaria distributed III 25-37, IV 9-11 (M.R. Ghosh et al., 1976). On undersides of leaves of Polygonum and Persicaria spp. in India (Meghalaya, Sikkim, West Bengal) and probably Japan, as the aphid illustrated and described by Moritsu (1983) as A. odaiensis may be this species. Close to A. taiwana, and the distinction between the two needs additional verification.

Akkaia neopolygoni M.R.Ghosh, A.K.Ghosh & Raychaudhuri (Fig.42w) Apterae are green to deep green; BL c. 2.4-2.5 mm. Alatae are undescribed. On apical parts and undersides of leaves of Polygonum and Persicaria spp. in India (Arunachal Pradesh, Sikkim, W Bengal).

Akkaia odaiensis Takahashi Apterae are yellow with all appendages pale; BL 1.8-2.0 mm. Alatae are undescribed. On Polygonum thunbergii in Japan, and Tao (1999) records its occurrence in China. Moritsu’s (1983) account refers to another species, probably A. bengalensis.

Akkaia polygoni Takahashi Plate 20b (Fig.42s,v) Apterae are yellow or orange-yellow, appendages concolorous with body; BL 1.9-2.4 mm. Alatae have secondary rhinaria distributed III 40-48, IV 14-17, V 0-3. On stems and upper sides of leaves of Polygonum and Persicaria spp. in Japan, China, Taiwan and Korea (BMNH collection, leg. K.S. Boo). Heteroecious, with sexual phase on Enkianthus spp. and Pieris japonica (Takahashi 1961d, Miyazaki 1971). 2n=24.

Akkaia pseudopolygoni Qiao, Jiang & Ren Colour of aptera in life unrecorded; BL 1.7-2.0 mm. Alatae are unknown. On an unidentified Polygonum sp. in Sichuan province, China (Qiao et al. 2006c).

Akkaia sikkimensis Agarwala & Raychaudhuri Colour of aptera unknown; BL 2.5-2.6 mm. Described from Raphanus sativus, but this will not be the true host. India (Sikkim).

Akkaia taiwana Takahashi (Fig.42r,t) Apterae are pale brownish white with jet black siph.; BL 1.8-2.1 mm. Alatae have secondary rhinaria distributed III 22-32, IV 6-13 (6-segmented antennae) or III 28-34 (5-segmented antennae). On Polygonum thunbergii and Polygonum sp. in southern Japan, Taiwan, China, Sri Lanka (BMNH collection, leg. Lund University Ceylon Expedition 1962), and alatae have been trapped in New South Wales, Australia (Carver 1976).

Aleurodaphis van der Goot

Hormaphidinae: Cerataphidini (?)

A little-known genus with eight nominal species in eastern Asia. Molecular data has cast some doubt about whether it belongs in Cerataphidini (Stern et al. 1997). It has aleyrodiform apterae with a complete crenulate margin of wax glands, as in Cerataphis, but distinguished from that genus by absence of frontal horns and a more evident division between prothorax and mesothorax. The biology is still very unclear. Host-alternation from Styrax was proposed, but the species involved, takenouchii Takahashi, is no longer thought to belong to this genus, and is now placed in Tuberaphis. One species of Aleurodaphis has now been found to produce leaf-roll galls on Stuartia (Theaceae) in Japan, and another was obtained from boat-shaped galls on Sinojackia xylocarpa (Styracaceae) in China. Others are described from Compositae/Asteraceae (three species), Bambusa, Impatiens and Ligularia. Two of the species on Compositae/Asteraceae in Japan are monoecious and anholocyclic (Takahashi & Sorin 1958). The Japanese species were reviewed by Sorin & Miyazaki (2004), and the Chinese species by Jiang & Qiao (2011) and Qiao et al. (2018).

Aleurodaphis antennata Chakrabarti & Maity Only the aleyrodiform aptera (BL 1.3-1.5 mm) is described, from Bambusa sp. in Uttar Pradesh, India (Chakrabarti & Maity 1982). The last rostral segment (RIV+V) is shorter than in other Aleurodaphis (1.1-1.5 × HT II). Appearance in life is unknown, presumably dark with marginal wax.

Aleurodaphis asteris Takahashi & Sorin Plate 3a Apterae are black, aleyrodiform, with a fringe of wax; BL 1.0-1.1 mm. On Aster spp. in Japan and Korea, living all year on stems at or just below ground level, but young larvae move to distal parts of stems in spring and summer. Also recorded from China, where it was also found on Carpesium abrotanoides (G. Zhang 1999). Anholocyclic, with seven apparently non-overlapping generations per year on Aster (= Gymnaster) savatieri (Takahashi & Sorin 1958). Alatae are unknown. 2n= c.32.

Aleurodaphis blumeae van der Goot (= A. sinisalicis Zhang; synonymy by Jiang & Qiao 2011) Apterae are pinkish purple to red-black, not as black as A. asteris, with glassy marginal rays of wax, and developing distinctive white spinal wax markings (Moritsu 1983); BL 0.9-1.4 mm. In dense colonies on stems, and also on undersides of young leaves of hosts, causing slight leaf-curl (Calilung 1967 and J. Martin, pers.comm.). Recorded from various Compositae/Asteraceae (Adenocaulon, Aster, Blumea, Carpesium, Chrysanthemum, Cynoglossum, Erechtites, Kalimeris), and from Mazus miquelii (BMNH collection, leg. W. H. Paik). A record from Salix (as A. sinisalicis Zhang) is possibly erroneous (Jiang & Qiao 2011). In east and south-east Asia (China, Japan, Korea, Vietnam, Taiwan, Assam, Philippines, Java, Malaysia).

Aleurodaphis impatientis Sorin & Miyazaki Apterae are blackish brown to dark purple brown with marginal white wax; BL 1.2-1.3 mm. On leaves and stems of Impatiens spp. in Japan. Anholocyclic, overwintering at stem bases (Sorin & Miyazaki 2004). 2n= c.30*.

Aleurodaphis ligulariae Sorin & Miyazaki Apterae are blackish brown to dark purple brown with marginal white wax; BL 1.9-2.1 mm. On undersides of leaves, stem apices and flowerstalks of Ligularia fischeri in Japan. Anholocyclic, overwintering on basal parts of Ligularia and in leaf litter (Sorin & Miyazaki 2004).

Aleurodaphis mikaniae Takahashi Apterae are reddish brown, with a fringe of wax; BL 1.3-1.6 mm. In dense colonies on stems of Mikania scandens in Taiwan, and there are subsequent records including one from a Parasenecio sp. in China (Shaanxi, Sichuan; Huang et al. 2012). 2n= c.30.

Aleurodaphis sinojackiae Jiang & Qiao Apterae are aleyrodiform, with marginal wax: BL 1.8-2.1 mm. On Sinojackia xylocarpa, inducing the leaves to curl and form boat-shaped galls (Jiang & Qiao 2011). Alatae were collected in June. In Zhejiang and Jiangsu provinces, China. The life cycle is unknown.

Aleurodaphis stewartiae Sorin & Miyazaki Colour in life unrecorded, presumably rather dark; BL of aptera (progeny of fundatrix?) about 1.5 mm. On Stuartia monadelpha in Japan (Honshû), rolling the leaf margin upwards to form a crescent-shaped gall (Sorin & Miyazaki 2004). Emigrant alatae emerge in late July/early August; two were collected on an Aster sp., which could possibly be the secondary host.

Aleurosiphon Takahashi

Aphidinae: Aphidini

One east Asian species closely related to Aphis but with differences in chaetotaxy of dorsal body, R IV+V and second tarsal segments, and in the position of the marginal tubercles on ABD TERG 7 in fundatrices (Takahashi 1966). Alatae have dark-bordered wing veins.

Aleurosiphon smilacifoliae (Takahashi) Apterae are dark green, thickly covered with wax powder; BL 1.4-2.1 mm. On Smilax spp., feeding along main veins on underside of leaves. Japan, Korea, China and Taiwan. Monoecious holocyclic with oviparae and alate males in October (Takahashi 1923). 2n=8.

Allaphis Mordvilko

Calaphidinae: Saltusaphidini

About 12 species with narrowly elliptical body-shape, differing from other Saltusaphidini in having setiform empodial hairs. Previously these species were treated as a subgenus (Trichocallis) of Thripsaphis, but Quednau (2010) restored its full generic status, establishing the correct type species as Allaphis caricicola Mordvilko (= Thripsaphis producta Gillette).

Allaphis californica (Hille Ris Lambers) (Fig.19j) Apterae are very pale, rather shiny yellowish white with glassy transparent legs and sides of body, but black antennae; BL 2.2-2.7 mm. Alatae are unknown. On Carex ?comosa in California, USA, and also collected on C. amplifolia in Washington (BMNH collection, leg. D. Carroll). 2n=10.

Allaphis cyperi (Walker) (Fig. 19l) Apterae are described as yellowish green, covered with bluish white wax, with dark appendages, and extensive greyish dorsal sclerotisation leaving a pale spinal stripe (but see also aphidtrek.org); BL 2.3-3.1 mm. Apterae have 0-5 secondary rhinaria on ANT III; alatae have 8-13 on III and 0-3 on IV, and dorsal abdomen with broad dark bars, usually separated between tergites. On leaves of Carex spp., aften on river banks. Widely distributed in Europe and North America, and also recorded from Kazakhstan (Kadyrbekov 2014f) and east and west Siberia (Quednau & Shaposhnikov 1988; Stekolshchikov & Khureva 2020). Oviparae and apterous males occur in September (UK, BMNH collection) to late October (Spain, Nieto Nafría & Mier Durante 1998) in Europe, but an oviparae collected in early July in Wisconsin USA was depicted by Quednau (2010). A darker form with shorter appendages and a subarctic distribution in Greenland, Iceland, northern Scandinavia and northern Russia is now regarded (Quednau 2010) as a subspecies, A. cyperi ssp. amurensis (Mordvilko) (= Thripsaphis vibei in Blackman & Eastop 2006), with Thripsaphis vibei ssp. arctica Hille Ris Lambers now regarded as a seasonal variant of this subspecies). Oviparae and apterous males of this subspecies were collected in late July in Greenland, and in early August in Iceland (Hille Ris Lambers 1960b, as Thripsaphis vibei ssp. arctica). However Stekolshchikov (2015) has reported the nominal subspecies (A. cyperi cyperi) as occurring in north-east Siberia (Chukotka), and Stekolshchikov (2017) reported that specimens from the Nenets region in north-west Russia had characters of both subspecies..

Allaphis daviaulti (Quednau) (Fig.19g) Apterae are very elongate, pale whitish yellow, with pale legs, and antennae black distally; BL 1.9-2.3 mm. Alatae are unknown. In leaf sheaths of Carex ssp. (trichocarpa, haydenii) in eastern Canada (Quebec, Ontario). The sexual morphs are undescribed; the record of an ovipara by Blackman & Eastop (2006) is unsubstantiated.

Allaphis foxtonensis (Cottier) Apterae are pale green to yellowish green; BL 2.4-2.7 mm. Alatae have 12-17 rhinaria on ANT III, and dark transverse bars on dorsal abdomen (cf. T. producta). On Carex spp. (ternaria, gaudichaudiana), especially colonising bases of young leaves in centre of plant. In New Zealand and Australia (NSW), but without related species in the southern hemisphere and almost certainly an introduced species. It appears closely related to A. producta; the differences were discussed by Hille Ris Lambers (1974). 2n=10*.

Allaphis hybrida (Hille Ris Lambers) Colour of apterae in life is unknown; BL 2.2-2.5 mm. On Carex sp. in California, USA. The possibility that this population was a hybrid between A. ossiannilssoni and A. verrucosa was discussed in the description of this species (Hille Ris Lambers 1974b), but it was regarded as a good species by Quednau (2010).

Allaphis ossiannilssoni (Hille Ris Lambers) (Fig.19i) Apterae are very elongate, pale greyish yellow to greyish brown with darker marginal sclerites, and a dark grey eighth abdominal tergite, secreting bluish wax wool especially at sides and end of abdomen; BL c.1.7-1.9 mm. Alatae have 8-13 secondary rhinaria on ANT III and 1-3 on IV. On Carex spp., widely distributed in Europe and across Asia to Japan (Higuchi 1972) and China (as Thripsaphis ossiannilssoni ssp. hebeiensis Zhang & Zhang, synonymised with the nominate species by Quednau 2010; the host in China is recorded as Cyperus hebeiensis, but this is not a recognised plant name).

Allaphis pacifica (Hille Ris Lambers) Similar to A. ossiannilssoni, and described as a subspecies of that species, but raised to specific rank by Quednau (2010). Alatae are unknown. On Carex sp. in California, USA.

Allaphis producta (Gillette) (= caricicola Mordvilko 1921, = cyperi ssp. wulingshanensis Zhang & Zhang 1993; synonymy by Quednau 2010) (Fig.19e,h) Apterae are very elongate, yellowish white to pale greyish brown, secreting bluish white wax, especially at end of abdomen; BL 1.8-2.5 mm. Alatae have 10-14 rhinaria on ANT III, and broad black cross bars that are fused into a solid patch on abdominal tergites 3-6. On leaves of Carex spp. in North America, Europe, and across Asia to east Siberia ; it is now considered that palaearctic populations formerly known as Thripsaphis caricis (Mordvilko) are this species. There are oviparae in BMNH collected in mid-July in Iceland, and in mid-August to September in UK, and oviparae and apterous males were found in Spain in mid-October (Nieto Nafría & Mier Durante 1998).

Allaphis scabra (Hille Ris Lambers) (Fig.19m) Colour of apterae in life is unknown; BL 2.4-2.8 mm. Alatae have 9-11 secondary rhinaria on ANT III. On Carex sp. in western USA.

Allaphis stromi Quednau Colour of apterae in life is unknown; BL 1.5-2.6 mm. Alatae are unknown. On Carex spp. in central USA (Quednau 2010).

Allaphis utahensis (Knowlton & Hall) (Fig.19k) Colour in life is unknown, BL of apt. 1.3(?)-2.6 mm. An alata has c.9 secondary rhinaria on ANT III and broad dark cross bands on the dorsal abdomen. On Carex nebraskensis in western USA (Idaho, Utah). Oviparae and apterous males occur in October in Utah.

Allaphis verrucosa (Gillette) (Fig.19n) Apterae are yellow or pale greenish yellow, with antennae distally black and legs mainly pale; BL 2.4-2.8 mm. Alatae have 9-20 sec. rhin. on ANT III, and dorsal abdomen with broad dark bars partially separated between tergites. On Carex spp. in Europe, west Siberia, and widely distributed in North America. Hille Ris Lambers (1974) distinguished populations in Labrador/Alaska and south-west California as verrucosa ssp. nodulosa and verrucosa ssp. subverrucosa respectively, and these subspecies are retained by Quednau (2010), although it still needs to be demonstrated that the described differences are not environmentally induced. Sexuales occur in late August-September in Sweden (Ossiannilsson 1959). 2n=10.

Allocotaphis Börner

Aphidinae: Macrosiphini

A monotypic genus for a hairy aphid with elongate siphunculi and a short cauda.

Allocotaphis quaestionis (Börner) Plate 10d Apterae are rather large, green with a waxy bloom, rolling apple leaves (Malus spp.) in spring. The second generation are all alatae; green, with rust-brown thorax, a black dorsal abdominal patch, and very numerous tuberculate secondary rhinaria on ANT III-V. They migrate for the summer to Senecio doronicum. The gynoparae returning to apple initially give rise to single-parent colonies on the undersides of the leaves (Hille Ris Lambers & Wildbolz 1958). In Austria, north-west Russia, Iran and Kazakhstan. However, it may also be in China, as a gynopara probably of this species was recorded by Zhang (1999) from Pyrus xerophila, and described as a “sp. dubia” under the name A. minensis Zhang, Chen, Zhong & Li. A record from Tunisia (Ben Halima-Karmel 2012) also needs further confirmation.

Allothoracaphis Takahashi

Hormaphidinae: Nipponaphidini

The apterae of the only known species have long marginal prosomal hairs like Thoracaphis but lack siphuncular pores and marginal hairs on the reduced ABD TERG 2-7. They have a less sclerotic dorsum than Thoracaphis, with an irregular mosaic-like ornamentation.

Allothoracaphis piyananensis Takahashi Apterae are pale yellow to yellowish-brown, flattened dorsoventrally, on the undersides of leaves along the main veins. Recorded from Quercus glauca and Q. morii in Japan and Taiwan (Takahashi 1958a). The alate morph is unknown, and the species is apparently anholocyclic.

Allotrichosiphum Takahashi

Greenideinae: Greenideini

A genus for three Fagaceae-feeding species, narrow-bodied and with long siphunculi like many Eutrichosiphum, but differing in the presence of distinctly capitate or spatulate hairs. Qiao et al. (2006a) reviewed the genus.

Allotrichosiphum assamense Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae about 1.6 mm, appearance in life unrecorded but probably green or pale brown; known only from one apterous vivipara and one immature collected on Quercus dealbata (= Q. floribunda? or Lithocarpus dealbatus?) in Assam, India (Raychaudhuri et al. 1973). (Note that this is a different species from Eutrichosiphum assamense).

Allotrichosiphum cyclobalanopsidis Qiao, Jiang & Martin Appearance in life unrecorded, BL of aptera 1.9-2.3 mm. Collected in December on Quercus (Cyclobalanopsis) spp. (neglecta, bella) in China (Hong Kong, Hainan, Yunnan). The description (Qiao et al. 2006a) includes an alata and first and fourth instar nymphs.

Allotrichosiphum kashicola (Kurisaki) Apterae (fundatrices) 2.3-2.8 mm, colour in life unrecorded, probably green or pale brown. Alatae dark greenish-brown, clear-winged, with siphunculi nearly as long as body. On young foliage of Quercus (Cyclobalanopsis) spp. (acuta, glauca, myrsinifolia) in Honshu and Kyushu, Japan. The life cycle is greatly abbreviated, with sexual morphs (greenish alate oviparae and yellow alate males) appearing in May. The rather flat eggs are laid on the undersides of leaves. At Osaka there are only two generations per year (fundatrices and sexuales), although at Tokyo additional parthenogenetic generations have been observed (Takahashi & Sorin 1957, 1959). Apterous viviparae other than fundatrices have not been detected.

Aloephagus Essig

Eriosomatinae: Fordini

A genus with only one known species, probably of African origin. Absence of siphunculi, short, 5‑segmented antennae, elongate rostrum and characteristic habitat and appearance in life make confusion with other aphids unlikely. Hille Ris Lambers (1954a) redescribed Aloephagus and discussed its taxonomic position.

Aloephagus myersi Essig Plate 1e, fig. 81F Apterae are pale or variably pigmnted and secrete a flocculent wax; BL 1.8‑2.5 mm. On Aloe spp., living under the leaf bases, attended by ants, and also reported from the related genera Haworthia and Astroloba. Photos of live aphids and damage to Aloe can be found at:- http://www.forestryimages.org/browse/TaxThumb.cfm?fam=30&genus=Aloephagus.

Widespread in Africa, especially south of the Sahara, where it is probably native, but originally described from California. It is also found in glasshouses in Europe, has been found on A. arborescens in Japan (Sano & Matsumoto 2005), and is reported to be a major pest of cultivated aloes in Australia (Forster 1998). It is presumed to have a sexual phase on Pistacia in Africa, as emigrant alatae suspected to be this species, BL about 1.8 mm, have been collected from galls on a Pistacia sp. (presumably P. aethiopica) in Kenya (D. Hille Ris Lambers, pers. comm.; specimens in BMNH collection). Anholocyclic on Aloe in Europe and North America (Blackman & Eastop 2000, p.220). The galls and the emigrant alatae are as yet undescribed. 2n=22.

Alphitoaphis Hottes

Aphidinae: Macrosiphini

One species on Lonicera in North America related to Hyadaphis but with thinner siphunculi and hind wings of alata having only one oblique vein. Aphids collected on Eupatorium sp., at ground level with ants, in Washington DC (BMNH collection, leg. A. Jensen), and on Symphoricarpos albus in Idaho (A. Jensen colln – aphidtrek.org) may also belong to this genus.

Alphitoaphis lonicericola (Williams) Apterae yellowish brown (Hottes 1926b) or reddish purple (original description), dusted with wax powder; BL 1.6-2.1 mm. On young growth of Lonicera spp., causing leaf-curl. In USA (Minnesota, Nebraska, Illinois, Kansas). Alatae have very numerous secondary rhinaria, distributed III 35-47, IV 24-30, V 14-17 (Hottes, 1926b). Life cycle is unknown.

Amegosiphon Narzikulov

Aphidinae: Macrosiphini

A genus for one species on Berberis in Central Asia, characterised by unusual features of the siphunculi and cauda.

Amegosiphon platicaudum Narzukulov (Fig.17a,d) Apterae are rather elongate-bodied with long appendages, pale green to yellowish with an irregular red spinal stripe; BL 1.6-2.7 mm. On undersides of leaves of Berberis spp. in Iran, Kazakhstan and Tajikistan. Oviparae and alatiform (brachypterous) males occur in October-November (Remaudière & Davatchi 1959, as Elbourzaphis behboudii). Mehrparvar (2016) provided a redescription.

Ammiaphis Börner

Aphidinae: Macrosiphini

One palaearctic species on Umbelliferae/Apiaceae with well developed marginal tubercles on abdominal tergites 1-(3-) 5 and apterae with secondary rhinaria.

Ammiaphis sii (Koch) Plate 10e Apterae are pale green or yellow-green, lightly wax-dusted, with black siphunculi; BL 1.9-2.1 mm. On Falcaria vulgaris, in summer colonies on stems and in leaf-sheaths, causing swelling of sheaths and twisting and curling of leaf blades, and in spring and autumn at bases of stems and on young shoots. Central and eastern Europe, south-west and central Asia (Azerbaijan; Rusanova 1948, as Aphis falcarii; Kazakhstan). Monoecious holocyclic according to Börner (1952).

Amphicercidus Oestlund

Aphidinae: Macrosiphini

Ten species in Asia and North America mostly associated with Lonicera, characterised by having secondary rhinaria in apterae and a very short broadly rounded cauda.

Amphicercidus alticola (Narzikulov & Mukhamediev) Apterae are pale green with dark appendages; BL 2.3-2.6 mm. Alatae have 24-32 secondary rhinaria on ANT III. On Lonicera nummularifolia in Central Asia (Tajikistan). A population with apterae having more rhinaria on ANT III was described as a subspecies, A. alticola ssp. tschatcalica (Mukhamediev & Akhmedov 1982).

Amphicercidus flocculosus (Gillette & Palmer) Plate 10i Apterae have a blackish brown thorax and abdomen, dull orange to clay-coloured head, sides of body and bases of siphunculi, and are wax-powdered; BL 2.1-2.8 mm. Alatae have secondary rhinaria distributed III c.40, IV 8-16. On leaves and twigs of Lonicera and Symphoricarpos in western USA and Mexico. Monoecious holocyclic, with oviparae and alate males in October-November (Palmer 1952).

Amphicercidus forsythiae Zhang, Zhong & Zhang Apterae are greyish green with white powder; BL 3.1-3.2 mm. Alatae have broad dark dorsal cross-bands tending to merge across ABD TERG 4-6, and secondary rhinaria distributed III 113-116, IV 0-2. On Forsythia suspensa in Yunnan, China. Apart from the small number of secondary rhinaria on ANT III of apterae – a character which is likely to depend on the degree of alatiformity – this could be A. japonicus.

Amphicercidus japonicus (Hori) Apterae are dark reddish to greenish or yellowish brown, with abundant white wax (see influentialpoints.com/Gallery); BL 2.5-3.4 mm. Alatae have secondary rhinaria distributed III 80-152, IV 0-4. On young shoots and twigs of Lonicera spp. in east Asia (northern India, China, Korea, east Siberia, Japan). Monoecious holocyclic, with oviparae and alate males in October in Japan (Miyazaki 1971), but perhaps anholocyclic in India (Hille Ris Lambers & A.N. Basu 1966, as A. indicus). 2n=8 (Chen & Zhang 1985b).

Amphicercidus laniger (Takahashi) Apterae are brownish green, densely covered in white wax powder; BL 1.9-2.5 mm. Alatae have 2-5 secondary rhinaria in the middle region of ANT III. On young stems of Stauntonia hebandra (= S. obovata) in Taiwan. The alata was described by Takahashi (1929).

Amphicercidus lonicerae Maity & Chakrabarti Apterae are wax-covered; BL 3.2-3.6 mm. Alatae have secondary rhinaria distributed III 55-75, IV 4-6. In downwardly-rolled leaves of Lonicera quinquelocularis in north India. 2n=8? (Khuda-Bukhsh 1980, as Amphicercidus sp.; 2n=18 was recorded by Khuda-Bukhsh & Pal 1983b, but this was probably an error).

Amphicercidus lonicericola (Mukhamediev) Apterae are pale green, wax-covered; BL 1.8-2.2 mm. On Lonicera altmannii in Central Asia.

Amphicercidus pulverulens (Gillette) Apterae are dull greenish brown with abundant wax; BL 2.5-3.1 mm. Alatae have secondary rhinaria distributed III c.40, IV 0. On bark of Symphoricarpus spp., at or below ground level. In western USA and Canada. Monoecious holocyclic, with oviparae and apterous males in October-November (Palmer 1952).

Amphicercidus sinilonicericola Zhang Colour of apterae in life is unrecorded, presumably wax-covered; BL c. 3.0 mm. On Lonicera japonica in China. Possibly a synonym of A. japonicus.

Amphicercidus tuberculatus David, Narayanan & Rajasingh Apterae are green, wax-covered; BL 2.1-2.6 mm. Alatae have secondary rhinaria distributed III 50-63, IV 0-3 (David et al. 1972). On Lonicera quinquelocularis in north India. 2n=6 (3 samples; Chauhan & Kurl 1990) or 12 (Pal & Khuda-Bukhsh 1984).

Amphorophora Buckton

Aphidinae: Macrosiphini

About 27 species, mostly nearctic, with a few east and west palaearctic. About half live on Rubus. A few have become fern feeders. They are all medium-sized to large aphids, usually greenish and rather pale, with long appendages bearing short to medium-length hairs. The siphunculi are long, pale or dusky and slightly to moderately swollen on distal half. Most species on Rubus occur in loose colonies on shoots or solitarily on leaves. Mostly monoecious holocyclic as far as known, with many alatae produced in the third parthenogenetic generation. Heie (1995) and Blackman (2010) reviewed the species of north-west Europe. The nearctic species need revision (see aphidtrek.org). The great variation in chromosome number is discussed by Blackman (1980).

Amphorophora agathonica Hottes Apterae are pale green with siphunculi pale at base, becoming darker towards apices (see influentialpoints.com/Gallery); BL 2.4-4.7 mm. On young stems and undersides of leaves of Rubus idaeus var. strigosus, and occasionally in small numbers on other Rubus spp. Widely distributed in North America north of about latitude 38˚N. See Kennedy & Schaefers (1974) for population studies. Monoecious holocyclic with alate males. 2n=14.

Amphorophora ampullata Buckton (Fig.43e) Apterae are green, with dark-tipped siphunculi (see influentialpoints.com/Gallery); BL 3.0-5.0 mm. On undersides of fronds of ferns in numerous genera (Aspidium, Asplenium, Athyrium, Cystopteris, Dryopteris, Matteuccia, Onoclea, Polypodium, Polystichum, Thelypteris). In Europe, Asia and North America. Monoecious holocyclic with alate males. Originally described from Cystopteris montana, the name is probably being applied to a complex of species with different fern associations. In USA, populations mainly associated with Onoclea sensibilis and Matteuccia spp. (Onocleae) are regarded as a subspecies, A. ampullata ssp. laingi Mason. Populations colonising various ferns in north-east India and Nepal also have distinctive features and have also been given subspecies status, as A. ampullata ssp. bengalensis Hille Ris Lambers & A.N. Basu, which form has now been recognised as occurring in China (Su et al. 2014). Populations on Athyrium felix-femina in Netherlands and UK have a longer R IV+V and other morphological differences as well as a different karyotype (see below), and thus represent an undescribed species. Clearly there is a need for biological, morphometric and molecular studies to clarify relationships within this group. 2n=12 for samples from Dryopteris and Polystichum in UK and Japan (RLB) for ssp. bengalensis from north-east India (Kurl & Chauhan 1986a), and for ssp. laingi from Canada; 2n=10 for a sample from Athyrium filix-femina in UK.

Amphorophora amurensis (Mordvilko) Apterae are green with antennae, legs and siphunculi blackish, and a pale cauda; BL 3.0-3.5 mm. On Rubus spp. in east Asia. Takahashi (1961e) provided a description of this species as A. rubiphaga. 2n=14.

Amphorophora (Galiaphis) annae (Ossiannilsson) (Fig.29f) Apterae are whitish yellow, pale yellowish green or pale green, with darker apices to antennal segments, legs and siphunculi; BL 1.8-2.3 mm. On stems and undersides of leaves of Galium boreale in northern Europe (Sweden, Finland, eastern Germany). Oviparae occur in August-September, apterous males in September (Ossiannilsson 1954).

Amphorophora catharinae (Nevsky) Apterae are green, eyes black, antennal segments white with black apices, siphunculi pale with dusky apices; BL 2-3 mm. Alatae have 10-12 secondary rhinaria on ANT III. At shoot tips and on undersides of leaves of Rosa spp. in Uzbekistan, and also in Iran, Afghanistan, Kazakhstan (Kadyrbekov & Aoitzhanova 2005), Tajikistan and Kashmir (BMNH collection). Oviparae and alate males were described from R. ×damascena in November (Narzikulov & Umarov 1969).

Amphorophora coloutensis Smith & Knowlton Apterae are pale whitish with a greenish brown abdominal spinal stripe; BL 2.9-3.4 mm. On Geranium spp. (wild) in western USA (Colorado and Utah). Monoecious holocyclic, with oviparae and alate males in late July/August (original description).

Amphorophora (Galiaphis) cryptotaeniae (Takahashi) Apterae are pale, siphunculi dark-tipped; BL 1.0-1.3 mm. On Cryptotaenia japonica in Japan. Its generic position is uncertain.

Amphorophora filipendulae Miyazaki Apterae are green or greenish yellow, with mainly dark siphunculi; BL 2.5-3.9 mm. On undersides of leaves and young shoots of Filipendula camtschatica in Japan (Miyazaki 1971) and Kamtschatica (Pashchenko 1988a). Monoecious holocyclic.

Amphorophora forbesi Richards (= Aulacorthum capilanoense Robinson) Apterae are pale green or whitish (see aphidtrek.org); BL 2.7-3.8 mm. On undersides of leaves of Rubus spectabilis in western North America (British Columbia, Washington, Oregon). This species has an unusual combination of characters (large size, spiculose ventral side to head, steep-sided slightly scabrous antennal tubercles, no rhinaria on ANT III, slightly swollen siphunculi and a rather hairy cauda), so its generic position is not at all clear. 2n=12.

Amphorophora gei (Börner) Apterae are pale green or yellowish green, with dusky or dark siphunculi and cauda; BL 2.2-3.9 mm. On undersides of leaves of Geum spp., especially G. rivale in moist, shady places (Heie 1995). Also recorded from Alchemilla vulgaris and Filipendula ulmaria. Widespread in Europe, and introduced to western USA (Idaho: BMNH collection, leg. J. H. Martin). Monoecious holocyclic on Geum; Hille Ris Lambers (1947) described the fundatrix, but sexuales have apparently never been found. 2n=12.

Amphorophora geranii Gillette & Palmer Apteraea are “dull greenish yellow to pinkish, mottled with sooty or grass-green”; BL 2.4-3.7 mm. Alatae were described in Palmer (1952). On leaves of Geranium richardsoni and G. bicknellii in western USA (Colorado, Idaho, Utah, Wyoming).

Amphorophora idaei (Börner) Apterae are pale green, rather shiny, with pale siphunculi (see influentialpoints.com/Gallery); BL 2.6-4.1 mm. On undersides of leaves of Rubus idaeus s.str., and cultivated varieties derived from it. Europe, and also recorded from Central Asia (Kazakhstan; Kadyrbekov 2005c) and west Siberia (Stekolshchikov & Khureva 2020). Monoecious holocyclic, with alate males. Blackman et al. (1977) distinguished this species from A. rubi, under which name it was often previously included. McMenemy et al. (2009) reviewed its biology, and Mitchell et al. (2010) investigated biocontrol using a combination of plant resistance and the parasitoid Aphidius ervi. 2n=18.

Amphorophora (Galiaphis) japonica (Takahashi) Apterae are green; BL c.1.5 mm. Host recorded as “Anemone (?)”. Japan. Host and generic position are uncertain.

Amphorophora kesocqua Hottes Apterae are bluish green, siphunculi becoming darker distally; BL c.3 mm. On Rubus sp. in Colorado, USA. Possibly these are fundatrices of A. tigwatensa.

Amphorophora pacifica Hill Apterae are shining very pale greenish white with a darker green rather wavy spinal stripe, somewhat interrupted posteriorly; BL 2.1-3.0 mm. On undersides of leaves of Rubus parviflorus in California, USA, where it is anholocyclic, overwintering on young foliage at apices of older shoots (original description). 2n=18.

Amphorophora parviflori Hill Apterae are pale green; BL 3.1-4.3 mm. On Rubus parviflorus, and sometimes in small numbers on certain other Rubus spp., in British Columbia, Canada. 2n=12.

Amphorophora pawtincae Hottes Apterae are pea-green; BL 2.5-3.7 mm. On leaves, stems and flowers of Primula parryi in western USA (Colorado, Oregon) (Palmer 1952).

Amphorophora rossi Hottes & Frison Apterae are whitish green with a darker green spinal stripe; BL 1.9-3.0 mm. On Geum spp., especially G. canadense, in north-east USA and eastern Canada, and also recorded from G. macrophyllum in western USA (Washington, Oregon; A. Jensen colln – aphidtrek.org). Monoecious holocyclic; the original description included the fundatrix. 2n=46*.

Amphorophora rubi (Kaltenbach) Apterae are shiny pale yellowish green to green with pale siphunculi (see influentialpoints.com/Gallery); BL 2.2-4.7 mm. On undersides of leaves of Rubus spp., especially fruticosus s. lat., in Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), west Siberia (Stekolshchikov & Khureva 2020), south-west and central Asia, Mongolia (Kadyrbekov & Aoitzhanova 2005; Kadyrbekov 2017a), and introduced to Australia and New Zealand. Monoecious holocyclic with alate males, or anholocyclic in regions with mild winters. Records of A. rubi on raspberries mostly refer to A. idaei in Europe and to A. agathonica in North America. 2n=20.

Amphorophora rubicumberlandi Knowlton & Allen Colour of apterae in life unrecorded; BL 2.6-3.7 mm. On canes of wild and cultivated black raspberries (Rubus occidentalis, R. leucodermis) in north-western USA (Oregon, Washington).

Amphorophora rubitoxica Knowlton Plate 24g Apterae are shiny deep green, or yellow-green with a darker green spinal stripe, with siphunculi dusky-dark; BL 2.3-3.5 mm. On young shoots and under leaves of many Rubus spp., and causing lesions on leaves of R. occidentalis, but not colonising R. idaeus. Western North America. Life cycle has not been studied. 2n=30.

Amphorophora scabripes Miyazaki (Fig.43d) Apterae are pale green with two longitudinal dark green pleural stripes, and black-tipped siphunculi; BL 4.8-5.7 mm. On Thelypteris (= Lastrea) quelpaertensis in Japan. Smaller yellow aphids (BL 3.6-4.8 mm) with dark hind femora on Matteuccia intermedia (= Pentarhizidium intermedium) in China were described as a subspecies, A. scabripes ssp. galba by Su et al. (2014), but are perhaps a separate taxon closer to the A. ampullata group (q.v.).

Amphorophora sensoriata Mason Apterae are very pale bluish green with dark antennae, and siphunculi dusky towards apices; BL 2.8-3.5 mm. On stems of Rubus occidentalis, dropping off readily when disturbed. north-eastern North America, as far south and west as Kansas. 2n=72.

Amphorophora stachyophila Hille Ris Lambers (Fig.50c) Apterae are rather shiny pale green, rarely greenish pink; BL 2.5-3.5 mm. On stems and undersides of leaves of Stachys spp, esp rigida, in western North America (California, Oregon, Washington, British Columbia). 2n=12.

Amphorophora stolonis Robinson Apterae are pale green; BL 2.6-3.1 mm. On stems or at ends of runners of Rubus pubescens in Manitoba, Canada. Monoecious holocyclic with alate males (original description). 2n=48.

Amphorophora tigwatensa Hottes Apterae are pinkish brown or greenish, with antennae, legs and siphunculi mainly dark brown, cauda paler; BL 2.8-4.1 mm. On undersides of leaves and young stems of Rubus procerus (Leonard 1974), and also on R. occidentalis (A.Jensen colln – aphidtrek.org). In western North America. 2n=40*.

Amphorophora tuberculata Brown & Blackman (Fig.30d) Apterae are shining deep green; BL 2.2-3.4 mm. On leaves and flowerheads of Geranium macrorrhizum in south-east England and Bulgaria; specific to this host, and presumably indigenous to montane areas of continental Europe where this plant is native. Monoecious holocyclic, with both apterous and alate males (Brown & Blackman 1985). The low chromosome number has allowed detailed studies of this aphid’s spermatogenesis (Blackman 1985) and sex determination (Blackman & Hales 1986). 2n=4.

Amphorophora urtica Essig (Fig.55a) Apterae are dark red to greyish; BL 3.2-3.6 mm. On undersides of larger leaves of Urtica spp. in western North America (California, Utah, British Columbia). Life cycle is unknown, but presumably anholocyclic in California, as apterous and alate viviparae were abundant in December-January (Essig 1942).

Amphorosiphon Hille Ris Lambers

Aphidinae: Macrosiphini

One European species related to Amphorophora but with ante- and post-siphuncular sclerites and a long, very hairy rostrum.

Amphorosiphon pulmonariae (Börner) Plate 24h Apterae are pale to dark green with black siphunculi and posterior dorsal abdominal markings; BL 2.5-3.0 mm. On undersides of leaves and petioles of Pulmonaria spp. in Europe (Netherlands, Germany, Bulgaria, Czech Republic, Hungary, Ukraine). Monoecious holocyclic with apterous males (Hille Ris Lambers 1949).

Anaulacorthum A.K. Ghosh & Raychaudhuri

Aphidinae: Macrosiphini

A genus for two oriental species close to Aulacorthum, but with 4 hairs on the first tarsal segments.

Anaulacorthum fagopyri A.K.Ghosh & Raychaudhuri (Fig.28a) Apterae are pale brown; BL 2.3-2.8 mm. On undersides of young and old leaves of Fagopyrum cymosum in West Bengal, India. Alatae are undescribed.

Anaulacorthum zhangjiajiense (Zhang) Apterae are pale red with black dorsal markings; BL c.2.6 mm. On Sonchus oleraceus in Hunan Province, China. Described as a subspecies of Neomyzus circumflexus, and transferred to this genus by Eastop & Blackman (2005). Su & Qiao (2010b) provided a redescription.

Anoecia Koch

Anoeciinae

A distinctive holarctic genus of uncertain taxonomic position, with about 24 species, many of which are poorly known. Some have host alternation from Cornus to roots of Poaceae (in one case, Onagraceae). Others are entirely subterranean on Poaceae or Cyperaceae. Adult apterae of most species on grass roots are medium‑sized greenish grey or grey in colour, with a sclerotic dorsal abdominal plate. They have compound eyes and are fairly mobile when disturbed. Immatures are paler, white or cream in colour. The alate sexuparae, distinguished by their dark posteriodorsal abdominal patch and the large black pterostigmal spot on the forewing, are a common sight on Cornus leaves in the autumn, alighting and depositing ant-attended clusters of small, yellow and/or brown sexual morphs. The oviparae lay their eggs on the bark of the trunk. Different species utilise the native Cornus in North America, Europe and eastern Asia. Accounts are available for central Europe (Zwölfer 1958), north-west Europe (Heie 1980b), the UK (Blackman et al. 2019b), the Iberian peninsula (Nieto Nafría & Mier Durante 1998), India (Chakrabarti et al. 1982, A.K. Ghosh 1988), Japan (Sorin 1999a), China (J. Yang et al. 2008) and North American Rocky Mountain region (Palmer 1952). Halbert (1991) keyed the North American species. This is a difficult group in need of further revision. In the BMNH collection there are undescribed species from Cornus mas in Italy, and from Cornus spp. in Korea. Aclitus obscuripennis Foerster may be a specialised aphidiine parasitoid. See also Blackman & Eastop 2000 pp. 222-3.

Anoecia caricis Pergande Colour of aptera not noted; BL c. 1.4 mm. On roots of Carex sp. in USA.

Anoecia corni (Fabricius) Plate 5a, b Host-alternating between Cornus sanguinea and roots of Poaceae in Europe and North America; anholocyclic on roots of numerous species of grasses and some cereals in Europe, Asia, Africa, Argentina (Ortego 1998a) and North America. Apterae on grass roots are greenish grey to brown with sclerotized parts dark grey; BL 1.9-2.8 mm. Apterae in spring on Cornus are dark brown; fundatrices have reduced, 3-facetted eyes and 5-segmented antennae, whereas subsequent generations have large compound eyes and 6-segmented antennae. Both spring migrant alatae and the sexuparae returning to Cornus in autumn have a dark patch covering ABD TERG 3-6. BL of alata 1.9-3.0 mm. Wieczorek (2008a) described the reproductive system of the dwarf male, and Wieczorek & Świątek (2009) compared it with that of Glyphina betulae. See Knechtel & Manolache (1943) and Heie (1980) for further information. 2n = 6. An excellent photo of a sexupara of A. corni can be seen at: http://www.flickr.com/photos/dioctria/3929122241/, and see also influentialpoints.com/Gallery

Anoecia cornicola (Walsh) Host-alternating between Cornus spp. (amomum, stolonifera) and roots of Poaceae in North America, and also recorded from Brazil (BMNH collection) where it is presumably anholocyclic on grass roots. Apterae on grass roots (Andropogon, Digitaria, Eleusine, Muehlenbergia, Panicum, Poa, Setaria) are whitish or yellowish with cross-bands, wax-powdered; BL c. 1.7 mm. Alate including sexuparae produced on grasses in August-October have a dark dorsal abdominal patch, but this is apparently not present in spring migrants (Palmer 1952, as A. querci). BL of alata 2.0-2.5 mm. 2n=10.

Anoecia cornimaris Bozhko Apterae are pale brown; BL c. 2.4 mm. On Cornus mas and C. sanguinea in eastern Europe (Moldova, Ukraine). Life cycle is unknown.

Anoecia equiseti Halbert Apterae are cream-coloured with grey sclerotic areas; BL 2.1-2.4 mm. Late summer alatae commonly have vestigeal wings. In ant-attended colonies on roots of Equisetum laevigatum in Idaho, USA, and Manitoba, Canada (BMNH collection, leg. A. G. Robinson). Life cycle is uncertain, monoecious holocyclic or anholocyclic; oviparae were produced in late August, and eggs laid but no males were observed and eggs may have been inviable (Halbert 1991) .

Anoecia fulviabdominalis (Sasaki) Host-alternating between Cornus spp. (brachypoda, controversa) and roots of Poaceae in Japan; also anholocyclic populations occur more widely on roots of grasses, and it is a major pest of upland rice in east and south-east Asia. Apterae on grass roots are grey‑green; BL 2.1-2.5 mm. Apterae in spring on Cornus blackish, BL about 2 mm. Both spring migrant alatae and the returning sexuparae have dark dorsal abdominal patches. Tanaka (1961) gave an account (in Japanese) of its biology in Japan, summarized in Yano et al. (1983). Sorin (1999a) provided descriptions of the morphs produced on Cornus, and described fundatrices and emigrant alatae of a variant form as a subspecies, A. fulviabdominalis similis.

Anoecia furcata (Theobald) (= A. nemoralis Börner) Apterae are dark green to blackish brown; BL 1.4-2.4 mm. Alatae have a black dorsal abdominal patch. On roots of numerous grasses (Alopecurus, Arrhenatherum, Bromus, Deschampsia, Elymus, Festuca, Panicum, Phleum, Poa, Sieglingia), attended by ants. In Europe, Central Asia, India, China. Mainly anholocyclic on grass roots; part of the population may go through a sexual phase on Cornus sanguinea, but this has not been clearly established. Zwölfer (1957) and Paul (1977) studied variation in frequency and distribution of spatulate hairs. 2n=12.

Anoecia graminis Gillette & Palmer Apterae are dull black with dusky bands, wax-dusted; BL 1.6-2.5 mm. Alatae have no black dorsal abdominal patch. On roots of Hordeum sp. in Colorado, USA. Apparently monoecious holocyclic, with oviparae in September (Palmer 1952). Closely related to the European A. furcata. 2n=8.

Anoecia haupti Börner Apterae on grass roots are greenish to brown with sclerotic parts dark grey; BL 1.9-2.6 mm. Alatae have a dark dorsal abdominal patch; BL 1.4-2.2 mm. On roots of grasses (Bromus, Eragrostis, Hordeum, Poa, Setaria) in Europe (Germany, Spain, Portugal, Italy, Turkey), and there is also a record from an undetermined grass species in China (Inner Mongolia; J. Yang et al. 2008). Heteroecious holocyclic with a sexual phase on Cornus sanguinea (Zwölfer 1957), but the Cornus-feeding morphs and sexuparae are undescribed. 2n=8.

Anoecia himalayensis Chakrabarti & Maity Colour of apterae in life unrecorded, probably greenish or brown with grey sclerotic parts; BL 1.8-2.0 mm. On roots of unidentified grass species in northern India (Himachal Pradesh, Uttar Pradesh). Spring colonies comprising apterous fundatrigeniae and immature spring migrant alatae, in crinkled and crumpled leaves of Cornus macrophylla at 2550m altitude in Pakistan, were provisionally assigned to this species by Naumann-Etienne & Remaudière (1995).

Anoecia ilicicola Sorin Alatae described under this name from Ilex nipponica in Japan by Sorin (1999) are presumably vagrant alate sexuparae from populations on grass roots. Alatae identified as the same species have been collected in China on Cornus (=Bothrocaryum) controversa (J. Yang et al. 2008), as well as being recorded as presumed vagrants on other plants (Artemisia, Kalopanax).

Anoecia japonica Sorin Fundatrix, spring migrant alata, sexupara and ovipara are described from Cornus controversa and Cornus sp. in Hokkaido, Japan (Sorin 1999a). Sexuparae, presumably vagrants, were also collected on Pteryocarya sp. Both spring migrants and sexuparae have a black dorsal abdominal patch. Appearance in life and populations on secondary hosts (presumably grasses) are unrecorded.

Anoecia krizusi (Börner) Apterae are blue-green with dark grey dorsal sclerotisation; BL 2.1-2.5 mm. Alatae are unknown. On roots of Elymus (Agropyron) caninum, with tentative identifications also from Agrostis sp., Dactylis glomerata, Lolium perenne and Sorghum bicolor. In Germany, ?UK (Stroyan 1964), and also recorded from China (J. Yang et al. 2008). Monoecious holocyclic on grass roots (Zwölfer 1957, as Neanoecia). It seems closely related to A. haupti.

Anoecia major Börner Apterae are brown with extensive dark grey sclerotisation; BL 2.2-3.0 mm. Alatae (BL 2.3-3.2 mm) have a large black dorsal abdominal patch. On roots of grasses, especially Phalaris arundinacea (also Brachypodium pinnatum, Calamagrostis). Europe (UK, Sweden, Netherlands, Germany, Spain). Probably with a sexual phase on Cornus; not yet found on Cornus in nature, but Börner (1950) transferred sexuparae to C. sanguinea and obtained 2-3 generations of fundatrigeniae. Very close to A. corni and hybrids may occur. 2n=8 (2n=7 in possible hybrids with corni; Blackman 1980).

Anoecia mirae Narzikulov Apterae are brown, with broad dark abdominal cross-bands; BL c. 2.7 mm. On Sorghum halepense in southern Tajikistan. Probably a synonym of A. major.

Anoecia oenotherae Wilson Apterae are yellowish, with dusky cross-bands; BL c.1.5 mm. On roots of Oenothera biennis, apparently widely distributed in North America. Alate sexuparae produced on Oenothera roots in autumn are small (1.4-1.8 mm), with 0-4 secondary rhinaria on III, and may have dark cross-bands on ABD TERG 3-7, but never a solid black patch. Migration occurs to Cornus spp. (amomum, paniculata, stolonifera). Records of other Anoecia spp. from Oenothera roots may be due to misidentification. See also Palmer (1952).

Anoecia (Paranoecia) pskovica Mordvilko Apterae are greyish white; BL 2.4-2.7 mm. Alatae are unknown. On roots and subterranean runners of Cyperaceae (Carex, Eriophorum). In Europe (UK, Denmark, Sweden, Germany, northern Russia) and Central Asia (Kazakhstan; Kadyrbekov & Aoitzhanova 2005) . Monoecious holocyclic with oviparae and apterous males from mid-October. Usually ant-attended, but eggs are covered in protective wax and not collected and stored by ants (Zwölfer 1957).

Anoecia radiciphaga Pal & Raychaudhuri Colour of apterae in life is unrecorded; BL 1.7-2.1 mm. On roots of Eragrostis nigra and unidentified grass species in north-east India (West Bengal), and also recorded from various grasses and cereals including Alopecurus aequalis, Avena fatua, Imperata koenigii (= cylindrica) in China (J. Yang et al. 2008).

Anoecia setariae Gillette & Palmer Apterae are pale orange-yellow with brownish head and dusky cross-band on posterior abdomen; BL 1.6-2.0 mm. Alatae commonly have vestigeal wings. In ant-attended colonies on roots of grasses (Echinochloa, Setaria), apparently widely distributed in North America (Smith & Parron 1978). Probably monoecious holocyclic; ants were observed to gather eggs in September (Halbert 1991).

Anoecia similiradiciphaga Qiao & Jiang Apterae are presumably dark brown, BL 1.5-2.0 mm. On roots of Triticum aestivum in China (J. Yang et al. 2008), and apterae and alatae are also recorded from (roots of?) Cyanotis vaga and Spiraea salicifolia, which are unlikely hosts. Very similar to A. fulviabdominalis, the differences requiring further confirmation.

Anoecia stipae Mamontova Apterae are pale brown with blackish head, legs and tips of antennae, and dark cross-bands on abdiominal tergites 7 and 8; BL c. 2 mm. On roots of Stipa capillata in chalky ground in Ukraine.

Anoecia takahashii Sorin Apterae are presumably blackish brown, BL about 1.8 mm. Spring migrant alatae have a black dorsal abdominal patch (Sorin 1999a; the illustration is stated to be of the sexupara, but presumably this is in error, as that morph is not described) . On Cornus sp. in Japan.

Anoecia tanakai Sorin Only the sexupara is described, BL 2.1-2.5 mm, with an extensive black dorsal abdominal patch. In autumn on Cornus spp. in Japan (Sorin 1999a). Sexuparae, presumably vagrants, were also collected on Pterocarya rhoifolia.

Anoecia vagans Koch (= Anoecia willcocksi Theobald; Zwolfer 1957) Host-alternating between Cornus sanguinea and roots of Poaceae. Fundatrices on Cornus are blackish; BL c.5.0-5.8 mm (Depa & Mróz 2014). All their progeny are spring migrant alatae (BL 2.3-2.8 mm) which, unlike those of A. corni, have no dark posterior abdominal patch. Apterae on grass roots are yellowish, pale green or olive green; BL 2.1-2.7 mm. They feed on roots of numerous grass and cereal species (Aegilops, Agropyron, Arrhenatherum, Bromus, Cynodon, Elymus, Eragrostis, Festuca, Hordeum, Lolium, Panicum, Poa, Triticum). Alatae including sexuparae produced on grass roots have a black dorsal abdominal patch (BL of alate virginoparae 2.1-2.5 mm, of sexuparae 2.5-3.9 mm; Heie 1980b). In Europe, Tunisia, Egypt, Israel, Iran, Turkey, India, east Siberia, and alatae apparently of this species have been trapped in USA (South Carolina, Florida; Skvarla et al. 2017). The male genitalia were described and illustrated by Wieczorek et al. (2012). 2n=12.

Anoecia zirnitsi Mordvilko Apterae are greyish green or brownish green; BL 1.5-2.1 mm. Alatae are unknown. On thin roots of various grasses (Agrostis, Brachypodium, Calamagrostis, Festuca, Lolium). In Europe (UK, Denmark, Sweden, Finland, Germany, Poland, Latvia, west Russia), the Middle East and Central Asia (Kazakhstan; Kadyrbekov 2017a). Monoecious holocyclic with oviparae in October-December, eggs being stored by Lasius flavus (Zwölfer 1957). Males unknown; Zwölfer’s suggestion that oviparae may be parthenogenetic requires further investigation.

Anomalaphis Baker

Greenideinae: Cervaphidini

A genus for two Australian species living on Myrtaceae (one on Agonis and the other on Leptospermum). The antennae of both apterae and alatae are 5-segmented, and abdominal tergites 7 and 8 each bear slender hair-bearing processes. The siphunculi are cylindrical or tapering, with a ring of 3-8 hairs on the distal half, and the cauda is broadly rounded.

Anomalaphis casimiri Carver Apterae are small (BL 1.3-1.7 mm) and brown. Distinguished from the only other described species by the presence of polygonal reticulation on the siphunculi distal to the ring of hairs, and by dorsal hairs of aptera mostly arising from tubercles. Apterae and alatae occurring together with intermediate forms on Leptospermum scoparium var. Red Damask in New South Wales, Australia (Carver 1971). Appearance of colonies in life and biology are not recorded.

Anomalaphis comperei Pergande ex Baker Plate 5f Apterae are small, dark brownish or greyish green with a yellowish spinal stripe; BL 1.2-1.6 mm. Alatae have shiny black thoracic lobes and a black dorsal abdominal patch. In Western Australia on Agonis flexuosa; this may be the only true host, although the original records of this aphid were from Acacia and Eucalyptus. Monoecious holocyclic, with oviparae in February (Carver 1971).

Anomalosiphum Takahashi

Greenideinae: Cervaphidini

Nine species associated with woody, mostly shrubby Connaraceae, Leguminosae and Xanthophyllaceae in east and south-east Asia. The hosts of two species are unknown. The apterae have 4-segmented antennae, bear slender hair-bearing processes on abdominal tergites 7 and 8, and have a cauda tipped with a stylus. Alatae have 5-segmented antennae, much smaller processes on the posterior tergites, and the caudal process often indistinct. Both oviparae and males are alate in the one species (A. mendeli Quednau & Martin) of which the sexuales are known, obtained by canopy fogging from an unknown host. The male genitalia of this species were described and illustrated by Wieczorek et al. (2012). Agarwala & Martin (1994) revised the genus, and Quednau & Martin (2006) provided keys to apterae and alatae.

Anomalosiphum indigoferae A.K. Ghosh, M.R. Ghosh & Raychaudhuri Apterae in life are dark brown to black, BL 1.1-1.4 mm. Alatae are larger, BL 1.4-1.7 mm. Described from young twigs of Indigofera sp. and Mimosa sp. in India (A.K. Ghosh et al. 1971e). There are also records from Vigna sp. and Phyllanthus sp.; the latter record is of alatae only, probably vagrants. Recorded fom Sikkim and West Bengal. The life cycle is unknown. 2n=18.

Anomalosiphum murphyi Martin & Agarwala Plate 11a, b Colour in life unrecorded, apterae probably dark-pigmented; BL 1.2-1.6 mm. On Rourea sp. and Xanthophyllum stipitatum in Singapore, and on ?Sindora sp. in Sarawak. Alatae were collected in Singapore in January. The life cycle is unknown (Martin & Agarwala 1994).

Anomalosiphum philippinense Martin & Agarwala Colour of aptera in life is unknown, probably dark; BL about 1.3-1.4 mm. Alatae are larger, 1.5-1.8 mm. On Dalbergia ferruginea (= stipulacea) in the Philippines, and alatae have been trapped in Papua New Guinea (Martin & Agarwala 1994). Apterae and alatae were collected in February-April. The life cycle is unknown.

Anomalosiphum pithecolobii Takahashi Apterae are probably green in life; BL 1.3-1.4 mm. Alatae are greenish yellow. Described from alatae collected on young leaves and shoots of Pithecellobium (= Archidendron) lucidum in Taiwan. Subsequently apterae believed to be the same species were described from Canton on Albizia lebbek, although the alatae from Albizia showed some differences from the Taiwanese specimens (Takahashi 1936a). Specimens previously thought to be this species from the Philippines and Papua New Guinea were distinguished as A. philippense (Martin & Agarwala, 1994)

Anomalosiphum scleroticum Qiao & Zhang Only the alata is described, BL about 1.6 mm, collected in April from Dalbergia hupeana in southern China (Guanxi) (Qiao & Zhang 2001a).

Anomalosiphum takahashii Tao Apterae are pale yellow; BL 1.2-1.5 mm. Described fom an unknown host in China, and recently collected from Dalbergia millettii in Zhejiang province (Q. Liu et al. 2013). Martin & Agarwala (1994) provided a redescription of the aptera.

Anomalosiphum tiomanensis Martin & Agarwala Apterae are pale green; BL 1.3-1.5 mm. On new apical growth of Dalbergia torta (= D. candenatensis) in west Malaysia. Also collected from China on Dalbergia sp. in Hong Kong (Quednau & Martin 2006), and it is also recorded from Phyllanthus emblica in Guanxi province (Q. Liu et al. 2013). Apterae and alatae were collected in February. The life cycle is unknown.

Anthracosiphon Hille Ris Lambers

Aphidinae: Macrosiphini

A holarctic genus for two species feeding on runners of Potentilla, with subapical reticulation on the siphunculi, extensive dark dorsal sclerotisation and ventral spiculosity on the head. Closely related to Macrosiphum, the differences possibly being due to the different microhabitat.

Anthracosiphon crystleae Smith Apterae are reddish brown, sclerotic areas shiny, with black antennae and siphunculi; BL 1.8-2.4 mm. Alatae have 11-16 secondary rhinaria on ANT III. On runners of Potentilla canadensis and Potentilla sp. in eastern North America (North Carolina, Pennsylvania, Quebec). Presumably monoecious holocyclic, with alate males (original description).

Anthracosiphon hertae Hille Ris Lambers Plate 26i Apterae are shiny black dorsally, lead-coloured with rosy hue ventrally, appendages blackish; BL 2.3-2.8 mm. Alatae have 20-34 secondary rhinaria on ANT III. On runners of Potentilla spp. (anserina, palustris) in northern Europe. Monoecious holocyclic, with alate males and oviparae in September (original description and Prior 1971).

Antimacrosiphon Zhang

Aphidinae: Macrosiphini

One species in China with strong similarity to Sitobion and possibly not justifying separate generic status.

Antimacrosiphum bullacaudatum Zhang Apterae colour in life unrecorded, probably pale with dark siphunculi; BL about 2.2 mm. Feeding on Phyllanthus emblica in Yunnan, China (G. Zhang & Qiao 1998c).

Anuraphis Del Guercio

Aphidinae: Macrosiphini

A palaearctic genus of 10 species with fundatrices that typically crumple or roll the leaves of Pyrus in spring into “pseudogalls”. The progeny of the fundatrix are all alate and migrate to subterranean parts of Umbelliferae/Apiaceae or Compositae/Asteraceae, where they are ant-attended. The name Anuraphis has been applied more widely in the past to include species currently placed in Brachycaudus and Dysaphis. See Heie (1992) and Blackman (2010) for accounts of the three species in north-west Europe, Barbagallo & Cocuzza (2003) for morphometric studies and a key to western palaearctic species, and Kolesova (1970, 1972) for ecological studies of four species in Crimea.

Anuraphis cachryos Barbagallo & Stroyan Apterae are very dark brown to black, BL 2.0-3.0 mm. On main root and leaf bases below soil level of Cachrys spp. Also recorded from Echinophora tenuifolia and Ferula communis. In southern Italy (where it is apparently anholocyclic), and also recorded from Iran and Turkey.

Anuraphis capparidis Nevsky Apterae are brown; BL c. 1.9 mm. On Capparis spinosa in Central Asia. Presence of marginal tubercles on abdominal tergites 1 and 7 indicates that this is not an Anuraphis; perhaps it is an immature Aphis sp.

Anuraphis catonii Hille Ris Lambers Fundatrices fold leaves of Pyrus communis in spring, their all-alate progeny migrating to form colonies at ground level on Pimpinella spp. Apterae on the secondary host are bright yellowish or greenish, with brown legs and antenae; BL 1.6-2.4 mm. Alatae are much darker, with a large dark dorsal patch. Kolesova (1972) studied its ecology in Crimea. Secondary host populations occur throughout much of Europe, and in south-west and Central Asia (Kazakhstan; Kadyrbekov 2017a), but the pear-feeding generations are reported only from Crimea and Switzerland (Lampel & Meier 2007). 2n=26 (Kuznetsova 1975, by implication).

Anuraphis cortusae Nevsky Apterae are pale green with black eyes and dark tips to siphunculi; BL c. 1.9 mm. On Cortusa matthiola in Central Asia.

Anuraphis farfarae (Koch) Pear-Colt’s Foot Aphid. The large (BL 2.7-3.7 mm) plump-bodied brown fundatrices roll or fold leaves of Pyrus communis in spring, producing green progeny that all develop into alatae. These migrate to basal and subterranean parts of Tussilago, or sometimes they may occur on Petasites or Hieracium. Apterae on secondary hosts are dark greyish green often tinged with red laterally and with a yellowish spinal stripe, and with dark antennae, legs and siphunculi; BL 2.2-3.1 mm. Immatures are paler, yellowish white, reddish, or pale green with a yellowish white spinal stripe. Alatae have a broad dark patch on abdominal tergites 4-6, almost solid in spring migrants but smaller and with a large window in alatae produced on the secondary host. The oviparae on pear in autumn are relatively small and pale (BL 1.6-1.8 mm) with hind tibiae swollen along the entire length and bearing numerous scent glands. Males are relatively large (BL 2.3-2.6 mm) with secondary rhinaria distributed III c.140-150, IV 42-51, V 13-29, base VI 0-1. Part of the population probably also overwinters parthenogenetically on Tussilago roots (Shaposhnikov & Sharov 1977). In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), west Siberia (Stekolshchikov et al. 2008a) and south-west and Central Asia; a report of its occurrence in North America is unsupported (Foottit et al. 2006). 2n=12.

Anuraphis ferulae Shaposhnikov Colour of apterae in life unknown: BL c.3.3 mm. On sheaths of lower leaves of Ferula spp. in Tajikistan. Probably migrating from “pseudogalls” on Pyrus (Shaposhnikov 1995), but the primary host generations have not been described.

Anuraphis pyrilaseri Shaposhnikov Apterae are pale ochre-yellow with pale green cross-bands; BL 2.3-3.1 mm. On basal parts of many genera of Apiaceae/Umbelliferae (Cachrys, Ferula, Laser, Laserpitium, Magydaris, Opopanax, Thapsia). In south, central and eastern Europe, Russia (northern Caucasus) and Kazakhstan (Kadyrbekov 2017a). Heteroecious holocyclic in eastern Europe (Crimea), where pseudogalls were found in spring on Pyrus communis (Kolesova 1972), and in Switzerland it is recorded from P. communis ssp. pyraster (Lampel & Meier 2007). Anholocylic on Ferula in southern Italy (Barbagallo & Stroyan 1982), and probably elsewhere. 2n=12.

Anuraphis shaposhnikovi Barbagallo & Cocuzza Apterae are yellowish-ochreous to brownish; BL 1.8-2.6 mm. On root collar and leaf bases of Magydaris pastinacea and Opopanax chironium in southern Italy. Closely related to A. subterranea, but with relatively shorter R IV+V and fewer spinal tubercles (Barbagallo & Cocuzza 2003).

Anuraphis subterranea (Walker, 1852) Pear-Hogweed Aphid Plate 9h Plump-bodied brown fundatrices (BL 2.6-3.0 mm) roll and crumple the leaves of pear trees in spring like A. farfarae, but their progeny are brown (see influentialpoints.com/Gallery), developing as alatae which migrate in May to basal parts of certain Umbelliferae/Apiaceae, especially Heracleum and Pastinaca. Apterae on secondary hosts are brown, BL 2.0-3.5 mm, and their immature progeny are pale brown. They form large ant-attended colonies inside lower leaf sheaths, at stem bases and on subterranean parts; the mutualistic relationship with Lasius on Heracleum was studied by Hansen et al. (2006). The return migration to pear occurs in September. Alatae have a similar dorsal sclerotic pattern to A. farfarae. Oviparae on pear are small (BL 1.4-1.8 mm), relatively pale, with strongly swollen hind tibiae bearing numerous scent glands. Males are relatively large (2.3-2.7 mm) with secondary rhinaria distributed III c.120-140, IV 28-57, V 5-12, base VI 0-2. Throughout Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), Iran, western Siberia and Kazakhstan (Kadyrbekov 2014f). Cited to have been found in Colorado, USA, but there are no North American specimens in USNM (Foottit et al. 2006). 2n=26.

Anuromyzus Shaposhnikov

Aphidinae: Macrosiphini

One species on Cotoneaster, related to Dysaphis but with extensive dorsal sclerotisation and differences in the siphunculi and cauda (Shaposhnikov 1959a).

Anuromyzus cotoneasteris (Shaposhnikov) (Fig.24a,b) Apterae are very shiny dark brown to black; BL 2.2-2.5 mm. Secondary rhinaria of alatae (in spring) are distributed III 63-80, IV 18-23, V 3-6. On undersides of curled leaves of Cotoneaster spp. Known from Georgia, southern Ukraine (BMNH collection, leg G. Shaposhnikov) and the south of France (BMNH collection, leg. G. Remaudière). Monoecious holocyclic, with oviparae and alate males in October (original description). 2n=12.

Aphanostigma Börner

Phylloxeridae

A genus differing from Phylloxera in the absence of spiracles from abdominal segments 2-5. Two species are described from Pyrus, and are of economic importance (see Blackman & Eastop 2000, pp.224-5).

Aphanostigma iaksuiense (Kishida) Very small yellow phylloxerids laying eggs in bark crevices of Pyrus pyrifolia in Japan and Korea, and also in Sichuan province, China, where its ecology was studied by Fang (1980).

Aphanostigma piri (Cholodkovsky) Apterae are very small (BL 0.8-1.0 mm), yellow, laying eggs in bark crevices of Pyrus communis, also attacking and sometimes destroying the buds, and in some pear varieties causing necrosis around the calyx of the fruit. In southern Europe, Ukraine, Georgia, Crimea, Israel, Lebanon and also now in Thailand. A sexual phase is reported from southern France and Crimea, but it is apparently anholocyclic in Israel.

Aphidounguis Takahashi

Eriosomatinae: Eriosomatini

A monotypic genus characterised by the single developed tarsal claw of apterae on the secondary host. Emigrant alatae from galls on Ulmus have short antennae, forewings with unbranched media and small siphuncular pores without associated hairs. A cladistic study (Sano & Akimoto 2011) has indicated a close relationship with another monotypic east Asian genus Schizoneurella, which despite some obvious morphological differences has a remarkably similar life cycle.

Aphidounguis mali Takahashi (fig. 117A) Host-alternates between Ulmus parvifolia and roots of Malus spp. in Japan. Alatae have also been collected from U. parvifolia in China (Hangzhou), and from U. japonica and U. davidiana var. mandshurica in Korea. It was recently reported from North Carolina, USA (Footit et al. 2006). The Ulmus galls consist of several rolled leaves on one shoot; initially only one of these is occupied by the fundatrix, but later its progeny migrate to fill the empty leaf-rolls (Akimoto 1983). Alate emigrants, BL 1.5-1.7 mm, have dark brown head, thorax, legs and antennae, and dark bands between the abdominal tergites. Apterae on roots of Malus are small white aphids secreting wax wool, to which the name Watabura nishyae Matsumura has been frequently but wrongly applied. The alatae collected on apple roots in October in China described as A. pomiradicola Zhang & Hu (G. Zhang et al. 1999) are likely to be the sexuparae of A. mali. Apterae found causing economic damage to apple roots in Yunnan, China, and identified as Schizoneurella indica (Qiao et al. 2005d), may also be this species. [Note: the record of A. mali from Cydonia in Blackman & Eastop (2000) is in error.]

Aphidura Hille Ris Lambers

Aphidinae: Macrosiphini

A palaearctic genus of about 24 species, mostly in southern Europe or central Asia. They feed mainly at stem apices of Caryophyllaceae, but are little studied and life cycles are mostly unknown. Some species may host-alternate from Rosaceae, as one species (A. bozkhoae) is described from Prunus, and two from Prinsepia. They are Myzus-like aphids, but the presence of a pair of mesosternal processes indicates a relationship with Brachycaudus subgenus Acaudus; other Brachycaudus-like characters are the tendency for extensive dorsal sclerotisation, and the subapical annular incision of the siphunculi. Kadyrbekov (2013) described six new species from Kazakhstan. Nieto Nafría et al. (2013a, 2014) described more new species, reviewed the genus and provided a key to apterae, which was updated by Barjadze (2019).

Aphidura acanthophylli Remaudière Apterae are pale; BL 1.4-1.7 mm. Alatae have secondary rhinaria distributed III 15-22, IV 3-11, V 0-4. This species differs from other Aphidura in having apterae with dorsal hairs arising from conical tubercles. On an Acanthophyllum sp. in Iran (Remaudière 1989b). Nieto Nafría et al. (2013a) provided a photomicrograph of a mounted specimen.

Aphidura alatavica Kadyrbekov Apterae have variable and often extensive black dorsal sclerotisation, with unsclerotised areas yellowish green, and brown siphunculi and cauda; BL 1.7-2.0 mm. Living on Cerastium cerastoides and C. holosteoides (= fontanum ssp. vulgare), forming loose colonies on stems under flowers. In south-east Kazakhstan (Kadyrbekov 2013).

Aphidura amphorosiphon Nieto Nafría, Mier Durante & Remaudière Colour of apterae in life unknown; BL 1.3-1.6 mm. Alatae have 16-21 secondary rhinaria on ANT III, and a black dorsal abdominal patch. On Silene sp. and Eremogone rigida in Iran (Nieto Nafría et al. 2013a, Barjadze et al. 2017b). An alate male was collected in mid-September.

Aphidura bozhkoae (Narzikulov) Apterae are shiny black aphids, often with black siphunculi, BL 1.7-2.0 mm, infesting apical leaves of Prunus spp. (but not P. avium or P. cerasus), which curl up and turn red (Narzikulov 1958, 1965). Kadyrbekov (2013) also recorded it from Aflatunia (= Louiseania) ulmifolia. In mountainous areas of central Asia (Tajikistan, Kazakhstan, Kirgizia, Georgia, Iran). A single aptera collected on Hordeum in Pakistan (leg. C.I.B.C., 13.ii.63) also seems to be this species. The variation of key features of the morphology of this species (dorsal sclerotisation, pigmentation of siphunculi, development of mesosternal processes) was discussed and illustrated by Nieto Nafría et al. (2014). The life cycle is unknown.

Aphidura corsicensis Nieto Nafría, Blackman & Martin Apterae are shining dark green; BL 2.0-2.1 mm. On Cerastium soleirolii in the mountains of Corsica at 1100m (Nieto Nafría et al. 2014). Other morphs and biology are unknown.

Aphidura delmasi Remaudière & Leclant Apterae are shining dark brown with black siphunculi and pale cauda; BL 1.3-2.0 mm. Alatae have 3-8 secondary rhinaria on ANT III only. On Silene italica and Silene sp. in southern Europe (France, Sicily, Greece). Monoecious holocyclic, with oviparae and apterous males occuring in alpine France in late October; these morphs and the fundatrix were described by Nieto Nafría et al. (2013a).

Aphidura gallica Nieto Nafría, Mier Durante & Remaudière Apterae with variable black dorsal sclerotisation and black siphunculi and pale cauda, general colour in life unknown; BL 1.5-2.3 mm. Alatae have 16-34 secondary rhinaria on ANT III. On Silene spp. (gallica, paradoxa) in France (Nieto Nafría et al. 2013a). The life cycle is unknown.

Aphidura graeca Nieto Nafría, Mier Durante & Remaudière Colour of apterae in life is unknown; BL c.1.8 mm. Alatae are undescribed. On Gypsophila sp. in Greece (Nieto Nafría et al. 2013a).

Aphidura gypsophilae Mamontova-Solukha Apterae are yellow, BL c.1.9-2.1 mm. Alatae have 10-15 secondary rhinaria on ANT III only. On undersides of leaves and stems of Gypsophila spp. Recorded from Ukraine, Hungary, Slovakia, Russia (west Siberia), Iran (Rezwani et al. 1994) and Kazakhstan (Nieto Nafría et al. 2013a). The life cycle is unknown.

Aphidura iranensis Nieto Nafría, Mier Durante & Remaudière Apterae with black dorsal sclerotisation, colour in life unknown; BL 1.1-1.3 mm. On an unidentified species of Prunus in Iran (Nieto Nafría et al. 2013a). The life cycle is unknown.

Aphidura jimoi Barjadze Colour of apterae in life is unknown; BL 1.2-1.6 mm. On a species of Dianthus in Georgia (Barjadze 2019). Other morphs and life cycle are unknown.

Aphidura libanensis Nieto Nafría, Blackman & Martin Apterae with black dorsal sclerotisation, colour in lfe unknown; BL c.1.8-1.9 mm. On Prunus prostrata in Lebanon (Nieto Nafría et al. 2014). Other morphs and life cycle are unknown.

Aphidura massagetica Kadyrbekov Apterae have extensive black dorsal markings with unsclerotised areas yellowish green; BL 1.9-2.4 mm. An alatiform aptera had 22-24 secondary rhinaria on ANT III. In loose colonies on peduncles of Silene lithophila and S. brahuica in south and south-east Kazakhstan (Kadyrbekov 2013).

Aphidura melandrii Kadyrbekov Apterae have extensive black dorsal sclerotisation with unsclerotised areas yellowish green, and brown siphunculi and cauda; BL 1.9-2.3 mm. Living on Silene latifolia (= Melandrium album), forming loose colonies on stems under flowers. In east and south-east Kazakhstan (Kadyrbekov 2013). Very similar to A. ornatella.

Aphidura mordvilkoi Shaposhnikov Apterae are green or brownish green; BL c.1.8 mm. On shoot apices and petioles of Prinsepia sinensis in east Siberia. A. prinsepiae Pashchenko is a synonym (Kadyrbekov 2013).

Aphidura naimanica Kadyrbekov Apterae have variable black dorsal sclerotisation with unsclerotised areas yellowish green; BL 1.6-2.2 mm. The one described alata has 26-27 secondary rhinaria on ANT III. In loose colonies on stem under flowers of Gypsophila spp., in east and south-east Kazakhstan (Kadyrbekov 2013).

Aphidura nomadica Kadyrbekov Apterae have a black dorsal abdominal patch, with unsclerotised areas green; BL 1.7-2.3 mm. In loose colonies on peduncles of Silene spp. in Kazakhstan (Kadyrbekov 2013).

Aphidura ornata Hille Ris Lambers Apterae are shining black with unsclerotized areas pale greenish grey, siphunculi black and cauda greenish yellow; BL 1.8-2.2 mm. In rather dense colonies on stem apices and peduncles of Silene spp., sometimes visited by ants. Europe (Italy, France, Switzerland, Hungary, Russia, Ukraine) and Iran (Rezwani 1990).

Aphidura ornatella Narzikulov & Winkler Apterae are dark brown to black with unsclerotized areas green; BL 1.5-2.6 mm. Kadyrbekov (2013) provided a redescription, and described an alata with 29-30 secondary rhinaria on ANT III. Living singly or in loose colonies on shoot apices and peduncles of Silene spp. and Saponaria sp., in Tajikistan, Kazakhstan and also in Iran (Rezwani 2010), Pakistan (Nieto Nafría et al. 2013a) and north-west India (David et al. 1970, as A. bharatia). Differences from A. bharatia noted by Kadyrbekov (2013) may be due to intraspecific variation.

Aphidura pakistanensis Nieto Nafría, Mier Durante & Remaudière Colour of apterae in life is unknown; BL 1.6-2.0 mm. Alatae have secondary rhinaria distributed ANT III 32-48, IV 2-10, V 0 (Barjadze 2019). On unidentified Dianthus sp(p). Described initially from Pakistan (Nieto Nafría et al. 2013a), and also now recorded from Iran (Barjadze et al. 2017b), and from Georgia (Barjadze 2019). Barjadze (2019) compared the morphology of apterae from different locations and discussed intraspecific variation.

Aphidura pannonica Szelegiewicz Apterae are green with black dorsal markings, black siphunculi and dark green cauda; BL 1.7-2.3 mm. Alatae have 14-22 secondary rhinaria on ANT III only. On Silene spp. in eastern Europe (Hungary, Slovakia, Ukraine), also in Kazakhstan (Kadyrbekov 2017a), and there is also a record from Gypsophila paniculata (Nieto Nafría et al. 2013a). The Ukrainian population was given subspecies status (A. pannonica ssp. cretacea Mamontova-Solukha), but it is difficult to see the justification for this. 2n=12.

Aphidura picta Hille Ris Lambers (= Aphidura mingens Pintera) Plate 11i Apterae are yellowish to green with pale brown head and prothorax, very variable shiny brown dorsal abdominal markings, and dusky to brown-black siphunculi; BL 1.2-2.1 mm. Alatae have 25-39 secondary rhinaria on ANT III only. On flower-stalks of Silene spp. and Dianthus spp. in southern and eastern Europe (Sicily, Greece, Bulgaria, former Yugoslavia, Israel, Turkey) and south-west and central Asia (Iran, west Siberia, Afghanistan, Tajikistan, Pakistan). Narzikulov & Umarov (1969) gave subspecies status (A. picta ssp. pamirica) to a population in western Pamir.

Aphidura prinsepiae Pashchenko Apterae are green or brownish green; BL c.1.8 mm. On shoot apices and petioles of Prinsepia sinensis in east Siberia (Pashchenko 1988a). Probably synonymous with A. mordvilkoi.

Aphidura pujoli (Gómez Menor) Apterae are greenish, with pale/dusky siphunculi and cauda; BL 1.5-2.1 mm. Alatae have 22-25 secondary rhinaria on ANT III only. On Dianthus spp., especially D. caryophyllus, in southern Europe (Spain, Portugal, southern France, Corsica, Italy, Switzerland) and Ukraine, and also tentatively recorded from Silene parviflora (Bozhko 1976). The record from Pakistan (Naumann-Etienne & Remaudière 1995) should be applied to A. pakistanensis (Nieto Nafría et al. 2013a). Monoecious holocyclic in Corsica, where oviparae and alate males were collected in October (BMNH collection, leg. F. Leclant).

Aphidura togaica Kadyrbekov Apterae are pale yellow with paired brown dorsal markings; BL 1.6-2.0 mm. Alatae have 22-36 secondary rhinaria on ANT III and 0-1 on IV. In loose colonies on upper and lower sides of leaves of Gypsophila perfoliata and G. altissima in south-east Kazakhstan (Kadyrbekov 2013), and also now recorded from G. polyclada in Iran (Barjadze et al. 2017b).

Aphidura urmiensis Nieto Nafría, Mier Durante & Remaudière Colour of apterae in life is unknown; BL 1.9-2.1 mm. Alatae are undescribed. On Spergularia marina in Iran (Nieto Nafría et al. 2013a). The life cycle is unknown.

Aphiduromyzus Umarov & Ibraimova

Aphidinae: Macrosiphini

One rather large species on Rosa in Central Asia with an unusual combination of dark dorsal pigmentation, pale siphunculi, blackish cauda, and numerous secondary rhinaria in apterae as well as alatae, with no clear relationship to other rose-feeding aphids (Umarov & Ibraimova 1967).

Aphiduromyzus rosae Umarov & Ibraimova Apterae are dark green or brown with ANT I, II and VI, femora and cauda blackish, siphunculi pale; BL c.2.8 mm. Apterae have 20-25 secondary rhinaria on ANT III only, and alatae have them distributed III 30-35, IV c.6, V c.2. Alatae have an extensive black dorsal abdominal patch. On Rosa sp. in Tyan-Shan Mountains, Central Asia.

Aphis Linnaeus

Aphidinae: Aphidini

The largest aphid genus containing more than 500 species, most of which occur in the northern hemisphere, but a few are native to South America, New Zealand, and Australia. Mostly they are small to medium‑sized aphids, often greenish but varying from pale to dark green, and sometimes yellow, brown, reddish, or black. Most species live on shrubs or herbs, with relatively few on trees. They usually live on the young foliage and often distort the leaves, but some species live on young twigs or at the base of the stem, or on the roots. Most species are attended by ants which may construct earthen shelters for the species living just above soil level. There are evident species groups associated with particular groups of plants, e.g. subgen. Bursaphis host-alternating between Ribes and Onagraceae, but there seems to be no overall pattern of host association and aphid morphology on which to base a subgeneric classification with world-wide applicability. However, phylogenetic studies using both molecular (Coeur d’acier et al. 2007b, Kim & Lee 2008) and morphometric methods (Kim et al. 2010a) have confirmed that certain sub-groupings apply within the genus across the palaearctic region, and Lagos et al. (2014) have shown that they also apply to North American species. The primary hosts of the relatively few Aphis species that host-alternate provide no evidence of primitive host associations. An interesting possible explanation might lie in the suggestion that northern Aphis species might be derived from southern hemisphere ancestors (von Dohlen & Teulon 2003).

Accounts are available for the British Isles (Stroyan 1984), Fennoscandia/Denmark (Heie 1986), Switzerland (Lampel & Meier 2007), the Iberian peninsula (Nieto Nafría et al. 2005a), India (A.K. Ghosh & L.K. Ghosh 2006), Mongolia (Holman & Szelegiewicz, 1971), east Siberia (Pashchenko 1992, 1993a-c, 1994a-c) and Japan (Takahashi 1966). Pashchenko (1997a,b) provided keys to the east Siberian species. Brown (1989) keyed alatae of the north-western European species. Cook (1984b) reviewed and keyed the North American Compositae/Asteraceae-feeding species. Lagos et al. (2014) analysed morphological and molecular data for Aphis spp. in midwestern USA and identified four main species groups. Lagos-Kutz et al. (2016) reviewed and keyed the species of the “asclepiadis group”, which have a primary host asssociation with Cornus. An interactive key to Aphis species of midwestern USA is available online at 3I – An Interactive Key to Aphis in Midwestern United States of America. (speciesfile.org). Remaudière (1994) keyed the South American species (but since then many new species have been described, particularly from Chile and Argentina; see for example Nieto Nafría et al. 2016b; Ortego et al. 2004). Rusanova (1948) reviewed and keyed species in Azerbaijan, and described several new species that have not subsequently been recognised. Subgenus Bursaphis was revised by Remaudière (1993b), and subsequently studied intensively by Rakauskas and co-workers (e.g. Rakauskas 1998b, Rakauskas et al. 2011). Martin (1991) provided a key to Toxoptera, now placed as a subgenus of Aphis, and Qiao et al. (2008) reviewed and keyed the species of Toxoptera in China. Xu et al. (2021) provided a key to Aphis spp. with stridulatory spines on the hind tibiae. With some reservations we follow Kadyrbekov (2001c) in transferring all the species previously placed in Aphis subgenus Absinthaphis to Xerobion. We have previously treated Protaphis as a subgenus of Aphis, but now give it separate generic status, in accordance with Aphid Species File.

Aphis aba Miller Apterae have deep brown head and thorax, paler brown to pale red abdomen with dark brown marginal spots; BL 1.2-1.4 mm. Alatae have secondary rhinaria distributed III 18-22, IV 4-6, V 2-3. On leaf petioles and midribs of Iva xanthifolia in Massachusetts, USA.

Aphis acaenaevora Mier Durante & Ortego Apterae are greenish yellow, lightly wax-dusted, with dark brown or black head, thorax, siphunculi and cauda; BL 1.8-2.4 mm. Apterae have 1-8 secondary rhinaria on ANT III, alatae are unknown. On Acaena splendens, forming colonies at base of stem at or slightly below ground level. In Argentina (Andean region of Patagonia). Monoecious holocyclic with oviparae and apterous males in late April (Mier Durante & Ortego 1998).

Aphis acaenovinae Eastop Apterae are bright apple-green with three rusty orange blotches on abdomen, one at base of each siphunculus and the other in between; BL 1.4-2.0 mm. Apterae have 0-5 secondary rhinaria on ANT III, alatae have them distributed III 3-8, IV 0-2. On Acaena ovina, living at ground level in rosette, attended by ants. In South-east Australia. 2n=8 (D.F. Hales, pers. comm.).

Aphis acanthoidis (Börner) Apterae are dark green; BL 1.1-1.6 mm. On leaf rosettes and developing flower stems of Carduus spp. In Netherlands, central and eastern Europe, Italy (Barbagallo & Massimino Cocuzza 2014), western Siberia, and Kazakhstan, where it is also recorded from Cousinia sp. and Jurinea sp. (Kadyrbekov 2017a),. A record from Carthamus (= Carduncellus) plumosus in Algeria (Laamari et al. 2013) needs further comfirmation.

Aphis acanthopanaci Matsumura Apterae have BL 2.0-2.3 mm and are dark green according to original description of specimens from Acanthopanax ricinifolium (= Kalopanax septemlobus) (Matsumura 1917). Midsummer specimens (from Aralia cordata) have BL about 1.3 mm and are yellow according to Takahashi (1966), but he possibly had A. spiraecola. Only known from Japan. Alatae are undescribed, and the life cycle is unknown.

Aphis acetosae L. Apterae are dark green to black; BL 1.5-2.7 mm. Alatae have secondary rhinaria distributed III 4-21, IV 0-2. On stems and inflorescences of Rumex spp., especially R. acetosa, often in large ant-attended colonies. It has also been found on Rheum spp. (Pashchenko 1988a). In northern, central and eastern Europe, Iran, Kazakhstan, east and west Siberia, and now also recorded from Algeria (Laamari et al. 2013). Also recorded from USA, but this needs confirmation (Footit et al. 2006).

Aphis (Toxopterina) achillearadicis Pashchenko Apterae are dark green; BL c.1.4 mm. In small dense colonies on roots of Achillea ptarmicifolia. East Siberia (Maritime Territory). Clearly it is closely related to A. vandergooti, and therefore we have transferred it to subgenus Toxopterina.

Aphis achyranthi Theobald Apterae are dull yellowish brown with dark brown siphunculi and cauda (see aphids of Karnataka website), immatures yellow with white wax (David, 1958); BL 1.2-1.7 mm. Alatae have secondary rhinaria distributed III 16-20, IV 9-12, V 3-5. Described from Achyranthes sp. in southern India (Theobald 1929), and there are samples from A. aspera in BMNH collection from Pakistan (leg. M.A. Ghani). David (1958) redescribed it from Punica granatum in southern India, colonising tender shoots, not attended by ants, and dropping readily when disturbed. 2n=8 (Kurl & Chauhan 1986b).

Aphis acrita Smith Apterae are greyish pulverulent, with dark siphunculi and cauda; BL 1.7-1.9 mm. On Sedum sp. in Ohio, USA. Monoecious holocyclic, with both apterous and alate males (original description).

Aphis acuminata Nieto Nafría & von Dohlen Apterae are shining brown to black; BL 1.2-2.1 mm. On stems of Adesmia sp. in Salta province, Argentina (Nieto Nafría et al. 2016c), and now also found on Adesmia atacamensis in Chile (Arica and Parinacota region). Nieto Nafría et al. (2021b) provide a redescription, including an alata. Sexual morphs are unknown.

Aphis adesmiae Delfino Apterae are shining black, without wax powder; BL 1.3-2.1 mm. On Adesmia boronioides in Argentina, forming dense colonies mainly on the stems and inflorescences (Delfino et al. 2009). It was recently also recorded from Chile (Nieto Nafría et al. 2018). Alatae were described by Nieto Nafría et al. (2016c). Probably monoecious holocyclic, but sexual morphs are unknown.

Aphis affinis Del Guercio Apterae are pale yellow (small specimens) to dark grey-green, often with narrow dark intersegmental cross bands, immatures dusted with wax powder (see aphids of Karnataka website); BL 0.6-2.1 mm. Alatae have secondary rhinaria distributed III 3-9, IV 0. On Mentha spp., usually clustered on stem apices, attended by ants. In Europe, southern Russia, Middle East, Central Asia, India and Pakistan. Very small apterae (BL 0.57-0.83 mm) collected on M. suaveolens in Azores (BMNH collection, leg. F. Ilharco) are probably this species. There are single records from two other labiates, Rosmarinus officinalis in Tuscany (Barbagallo et al. 2011) and Marrubium alternidens (= anisodon) in Iran (Mohktari et al. 2012). Monoecious holocyclic with apterous males (Tuatay & Remaudière 1964). Ilharco (1987) redescribed apterae and alate viviparae. 2n=8.

Aphis agastachyos Hille Ris Lambers Apterae are oval, shining dark green (see aphidtrek.org), BL 1.5-2.0 mm, curling leaves of Agastache urticifolia and Agastache sp. in western USA (Idaho, Utah, Oregon, Washington). Possibly this species also colonises Monardella odoratissima (A. Jensen, pers. com.). Alatae have large dark marginal and postsiphuncular sclerites. Apterous males and oviparae were collected along with viviparae in July (original description), indicating that the life cycle is abbreviated.

Aphis agrariae Bozhko Apterae are dark brown, lightly wax-dusted, with black dorsal abdominal cross-bands and black appendages; BL c.2.2-2.3 mm. On Euphorbia agraria, colonising stems, flower stalks and leaves, in Ukraine.

Aphis albella Nevsky Apterae are white or pale yellow, with appendages only slightly dusky; BL c.1.8-2.1 mm. On undersides of leaves of Cichorium intybus, and also recorded in smaller numbers from Portulaca, Hibiscus and Foeniculum officinale. In southern Kazakhstan/Uzbekistan. Apart from the absence of pigmentation, which might be due to a genetic aberration or unusual environmental conditions, this species could be A. gossypii.

Aphis alchemillae Börner Apterae are shiny, dark green to black, with pale legs and antennae; BL 1.2-2.2 mm. On Alchemilla spp. (conjuncta, glaberrima, vulgaris) colonising stems, petioles and undersides of leaves, and sometimes in inflorescences (Börner 1952). In Germany and Austria. [From examination of specimens in the BMNH collection there appear to be two Aphis species on Alchemilla under this name, one collected from ant shelters at the stem bases of an unidentified species in Austria (leg. R.N.B. Prior), and the other from A. conjuncta in Frankfurt Botanical Garden (leg. M. Mackauer). Further work is necessary to see whether either of these is alchemillae Börner.]

Aphis alienus Theobald Apterae are green to dark green; BL 1.0-1.9 mm. Alatae have secondary rhinaria distributed III 4-7, IV 0 -3. On rhizomes and basal parts of Teucrium spp., especially T. scorodonia, in ant shelters. Known from various countries in Europe (England, France, Spain, Portugal, Italy, Czech Republic), and it has also been found in Pakistan (Naumann-Etienne & Remaudière 1995). Most pre-1983 records of this species are as A. scorodoniae (see Stroyan 1984).

Aphis alstroemeriae Essig Colour of apterae in life unknown, probably dark; BL 1.6-2.1 mm. On an unidentified species of Alstroemeria (Chilean Lily) in Aconcagua Province, Chile. Alatae have secondary rhinaria distributed III 5-11, IV 0-2. Remaudière (1994) redescribed the paratypes in the BMNH collection.

Aphis althaeae (Nevsky) (= davletshinae Hille Ris Lambers of Blackman & Eastop 2006; see García Prieto et al. 2004) Apterae are pale green, BL 1.6-1.8 mm, on undersides of leaves and stems of Alcea and Althaea spp., often forming large colonies which curl leaves and stunt young growth. Alatae have secondary rhinaria distributed III 6-15, IV 0-4, V 0-1. Nevsky (1929) also recorded it from Malva and Convolvulus, and Kadyrbekov (2004b) found it on Lavatera thuringiaca. Specimens collected on Salvia sclarea in Afghanistan were described as a subspecies, A. althaeae spp. afghanica Narzikulov & Umarov. In Europe (Germany, Poland, Russia, Spain, Italy, Greece, ?Sweden) and south-west and central Asia (Iran, Turkey, Lebanon, Kazakhstan, Uzbekistan). Monoecious holocyclic (Börner & Heinze 1957).

Aphis amaranthi Holman Apterae are dull olive green or reddish brown, BL 1.5-2.1 mm. On Amaranthus spp., colonising the roots and stem bases, and sometimes on the leaves or in the influorescences, attended by ants. Alatae have no dorsal abdominal sclerotization except laterally and at bases of siphunculi, and secondary rhinaria distributed III 8-22, IV 0-8, V 0-2, those on III being distributed irregularly (not in a row). In Brazil, Argentina, Venezuela, Cuba, Costa Rica and USA (American records: California – BMNH collection, leg. H. Walker, 1975; Florida – Halbert et al. 2000; Georgia and Louisiana -trapped alatae in BMNH collection). Life cycle is unknown; most common in winter in Cuba. 2n=8.

Aphis angelicae Lee & Seo, nec. Koch 1854 Apterae are pale yellow; BL not recorded. On Angelica decursiva, in ant shelter on stem. In Korea. Perhaps not an Aphis; apart from the 5-segmented antennae, it seems to have many features in common with the species described from A. sinensis in China under the name Dysaphis angelicophaga (q.v). [Note: if these two should prove to be synonymous, then angelicophaga becomes the valid name, as Aphis angelicae is preoccupied.]

Aphis antherici Holman (= liliago Müller) Apterae dark bluish green, olive brown or reddish brown to black, with basal part of antennal flagellum, and tibiae for most of their lengths, whitish; BL 1.1-2.2 mm. Alatae have secondary rhinaria distributed III 8-21, IV 0-3. On basal parts of Anthericum spp., living in ant shelters. In Czech Republic, Slovakia, Germany and Poland. Monoecious holocyclic with apterous or brachypterous males. Holman (1966) and Müller (1968a, as A. liliago) both gave full accounts of this aphid.

Aphis apocynicola Holman Apterae are olive to brownish green with black head and siphunculi, and pale cauda; BL 1.9-2.4 mm. Large colonies were found on the terminal parts of the shoots of Apocynum scabrum (= A. venetum var. scabrum?)in Uzbekistan, and on A. lancifolium in Kazakhstan (Kadyrbekov & Aoitzhanova 2005). Records of A. apocyni Koch from Apocynum in Kazakhstan, Uzbekistan and Tadzhikistan are referable to apocynicola, and A. apocyni from China could be hot weather dwarfs of this species. The life cycle is unknown.

Aphis aquilonalis Stekolshchikov & Khruleva Apterae are dark brown-black or dark dirty yellowish brown with sparse wax pulverulence; BL 2.0-2.6 mm. On Rumex arcticus (= R. aquaticus ssp. arcticus) in north-east Siberia (Chukotka), on undersides of leaves and in inflorescences. The life cycle is abbreviated; a collection in late July included old fundatrices, apterous and alate viviparae, oviparae and apterous males. Stekolshchikov & Khruleva (20150 provided descriptions of all morphs.

Aphis araliaeradicis Strom Apterae are pale brown, BL c.1.6 mm. Apterae have 0-5 secondary rhinaria on ANT III, alatae have 8-11. Described from Aralia nudicaulis in Wisconsin, USA, forming ant-attended colonies on roots and stem-bases. Also recorded from California (Smith & Parron 1978). Presumably monoecious holocyclic, but males are undescribed.

Aphis arbuti Ferrari Apterae are wine red; BL 1.7-2.7 mm. Alatae have secondary rhinaria distributed III 10-19, IV 0-6, V 0. On woody Ericaceae (Arbutus unedo, Erica arborea) in southern Europe, and also reported from Turkey (Görür et al. 2011a). A member of the taxonomically difficult A. fabae group, and only recognisable as a distinct species by its colour and host association. Mier Durante & Nieto Nafria (1986) provided morphometric descriptions of apterae and alatae. Apparently monoecious and holocyclic (Barbagallo & Stroyan 1982), but sexual morphs are still undescribed.

Aphis arctiumi Bozhko Apterae are yellow-green, wax-dusted; BL c. 1.5 mm. On Arctium tomentosum in Ukraine.

Aphis (Bursaphis) argentina Nieto Nafría, Ortego & Mier Durante Apterae are shining black; BL 1.8-2.4 mm. Alatae have 7-10 small secondary rhinaria on ANT III only. On Adesmia ?papposa and A. papposa var.radicifolia in Argentina (Chubut, Mendosa; Nieto Nafría et al. 2021b).

Aphis argrimoniae (Shinji) Apterae are green, with pale legs and antennae, BL 1.2-1.8 mm. Alatae have dark antennae with secondary rhinaria distributed III 12-24, IV 9-17, V 4-8. On Agrimonia spp. in Asia (Japan, Korea, east and west Siberia). Bozhko (1976a) described a similar aphid from Agrimonia eupatoria in Ukraine as a subspecies, A. argrimoniae ucrainica.

Aphis armata Hausmann Apterae are black in life, resembling A. fabae (see influentialpoints.com/Gallery); BL 2.2-2.9 mm. Alatae have secondary rhinaria distributed III 9-24, IV 0-8, V 0. On flower stems of Digitalis spp., vigorously attended by ants. In Europe, from UK to Italy (Barbagallo & Patti 1998), and introduced to British Columbia, Canada (BMNH collection, leg. C.-k. Chan). Its distribution is uncertain, as morphology of parthenogenetic generations overlaps with that of long-haired members of the A. fabae group. Monoecious holocyclic, with oviparae having hind tibiae much less swollen than A. fabae. See Jacob (1947) for a detailed account. 2n=8.

Aphis artemisiphaga Holman Apterae are green or dark green, mottled, matt; BL 1.1-1.5 mm. At base of stem of Artemisia spp., attended by ants, in east Siberia.

Aphis artemisiphila Holman Apterae are green, similar to A. artemisiphaga; BL 1.5-1.8 mm. On roots of an unidentified Artemisia sp., attended by ants, in east Siberia. Holman (original description) noted that differences from A. artemisiphaga could be seasonal and require confirmation.

Aphis (Pseudoprotaphis) artemisivora Holman Apterae are dull green, lightly wax-dusted, with a dark dorsal pattern; BL 1.9-2.4 mm. In ant-attended colonies on roots of an unidentified Artemisia sp. in Mongolia.

Aphis asclepiadis Fitch (= A. helianthi Monell, = A. carduella Walsh) Plate 6i Apterae on secondary hosts are pale green, yellowish green to deep olive green mottled with yellowish green, with black siphunculi (see influentialpoints.com/Gallery and aphidtrek.org); BL 1.4-2.5 mm. Alatae have secondary rhinaria distributed III 15-40, IV 0-2. Colonies occur in late spring and summer on stems and leaves of many genera and species, especially Compositae/Asteraceae and Umbelliferae/Apiaceae, and also able to colonise Apocynaceae. Heteroecious holocyclic, with a sexual phase on Cornus spp. Spring populations twist and curl the leaves of Cornus (Hottes & Frison 1931, as helianthi); apterae on Cornus in spring are 1.5-1.8 mm, dark yellow to green, with siphunculi variably pigmented from green to black. Oviparae in autumn have considerably swollen hind tibiae. In USA and Canada, and also recorded from Brazil on Yucca (assuming A. yuccicola Wilson is a synonym), Cuba, Costa Rica (as helianthi; Villalobos Muller et al. 2010), Peru (Vilca Mallqui & Vergara Cobián 2011) and Argentina (Gonzáles Rodríguez et al. 2016). Records from Turkey (Akyürek et al. 2010, as helianthi) and Kazakhstan (Kadyrbekov 2017a, as carduella) need further confirmation. Lagos et al. (2014) using both morphological and molecular data identified a clade of 10 species as “the asclepiadis group” (asclepiadis, cornifoliae, decepta, impatientis, neogillettei, nigratibialis, salicariae, saniculae, thaspii, viburniphila), five of which have Cornus as their primary or only host. Lagos-Kutz et al. (2016) reviewed and keyed this group, and provided conclusive molecular evidence that A. carduella should be treated as a junior synonym of A. asclepiadis. 2n=8.

[This species, found on numerous host plants in North America and now also introduced to South America, has a confused nomenclatural history. The name A. helianthi Monell was widely used in the North American literature, especially for populations on Compositae/Asteraceae, until Cook (1984b) established its synonymy with the earlier name A. carduella. Earlier, Addicott (1981) had synonymised A. heraclella, the name previously often applied to populations on Umbelliferae/Apiaceae, with A. carduella (as helianthi) on the basis of host transfers and morphological studies involving Cornus, Umbelliferae/Apiaceae and Epilobium (Onagraceae), but not Compositae/Asteraceae. Robinson & Chen (1969) had previously obtained transfers from Cornus and Helianthus to Sium suave, but not to other Umbelliferae/Apiaceae.]

Aphis astericola Tissot Apterae are dark reddish brown with mainly dark appendages; BL c. 1.7 mm. Alatae have 7-9 secondary rhinaria on ANT III only. Only known from Aster sp. in Florida, USA.

Aphis astragali Ossiannilsson (Fig.16i) Apterae are dull black, powdered with whitish wax; BL 1.9-2.7 mm. On Astragalus spp. in Sweden and Finland, where it is also recorded from Oxytropis campestris, and where oviparae and alate males are produced (on both host genera) as early as July (Heikinheimo 1984). Also recorded from Kazakhstan (Kadyrbekov 2017a). A. masoni on islands in subarctic Canada is possibly a synonym. A record from Algeria (Laamari et al. 2013) needs further confirmation.

Aphis astragalicola Holman & Szelegiewicz Apterae are black, with middle parts of antennae and tibiae whitish; BL 0.9-1.2 mm. On Astragalus sp., “presumably living on the basal parts of the stems” (from original description), and A. danicus (Stekolshchikov et al. 2008a). In Russia (west Siberia and Transbaikalia), and also recorded from Turkey (Şenol et al. 2014c), Moldova and Kazakhstan (Kadyrbekov 2017a). Alatae and sexual morphs are unknown.

Aphis astragalina Hille Ris Lambers (Fig.16j) Colour of aptera in life unknown; BL c.2.4-2.5 mm. Described from a white-flowered leguminous plant in Montana, USA and Astragalus sp. in Alberta, Canada, and there are later records from Astragalus pectinatus in Alberta and from Hedysarum alpinum in Manitoba (Robinson 1991) and Alaska (BMNH collection, leg. A.G. Robinson).

Aphis aubletia Sanborn Apterae are undescribed, alatae have black head and thorax and dusky to pale brown abdomen, siphunculi concolorous with abdomen, and cauda “black on distal half, hirsute” (original description); BL of alata c.1.3 mm. Alatae have secondary rhinaria distributed III 7, IV 3, V 1. Colonising all the above-ground parts of Verbena aubletia (= Glandularia canadensis) in May in Kansas, USA. Not recorded since original description.

Aphis (Toxoptera) aurantii Boyer de Fonscolombe Black Citrus Aphid Plate 13c, d Apterae are oval shiny, reddish-brown or black, with black‑and‑white banded antennae and black siphunculi and cauda (see influentialpoints.com/Gallery); BL 1.1-2.0 mm. Immatures are brownish. Alatae are brown-black with a black pterostigma and, unusually for Aphidinae, a usually once-branched media; they have secondary rhinaria distributed ANT III 2-14, IV 0(-6). In dense, ant-attended colonies on shoots and undersides of young leaves of host plants, causing slight rolling, twisting or bending of mid-ribs. Large colonies produce an audible scraping sound when disturbed. Found on more than 120 plant species, particularly trees and shrubs, in numerous families, especially Anacardiaceae, Anonaceae, Araliaceae, Euphorbiaceae, Lauraceae, Moraceae, Rubiaceae, Rutaceae, Sterculiaceae, and Theaceae. It is an important vector or viruses of economically-important plants (Blackman & Eastop, 2000, p.354). Distributed throughout the tropics and subtropics including the Pacific Islands, and in glasshouses in temperate climates. Apparently it is almost completely anholocyclic everywhere; however two alate males were collected on Citrus in December 1936 in Palestine (leg. F.S. Bodenheimer), and one ovipara was recently found in a sample from Sambucus in Malta (Pérez Hidalgo & Mifsud 2011). Stary et al. (1988) reported on biocontrol measures in southern France, and Hussein & Kawar (1985) studied natural enemies in Lebanon. Broughton & Harris (1971) analysed the sound produced by A. (T.) aurantii, which is the only aphid with audible stridulation. Populations in which alatae have several secondary rhinaria on ANT IV have been recorded on Celtis in Japan (as Aphis celtis Shinji 1922c) and on Ficus in China (referred here to A. schlingeri). Alatae of A. aurantii normally have only 0-1 rhinaria on ANT IV. 2n=8.

Aphis austriaca Hille Ris Lambers Apterae are pale sea-green, darker marginally, with head and prothorax blackish green, very faintly pruinose; BL 1.6-1.9 mm. Alatae have secondary rhinaria distributed (1 specimen) III 17-20, IV 4, V 0. Described from roots of Silene sp., probably S. nutans, in Austria, and also recorded from Poland, feeding quite deeply on roots of S. otites (Osiadacz 2009). Ant-attended. Very closely related to A. sambuci, and possibly just a variant anholocyclic population of that species.

Aphis axyriphaga Pashchenko Apterae are dark dingy-green, BL c. 1.3 mm. The holotype aptera has a pair of dark irregular sclerotic patches on dorsal abdomen, leaving broad unsclerotised spinal band. Alata has 4 large secondary rhinaria on ANT III. On undersides of leaves of Axyris amaranthoides in east Siberia.

Aphis axyriradicis Pashchenko Apterae are dark emerald green, BL c. 1.7 mm. On roots of Axyris amaranthoides, attended by ants. Alatae are undescribed. In east Siberia.

Aphis baccharicola Hille Ris Lambers Colour of apterae in life unknown, BL 1.4-1.7 mm. Alatae have 14-22 secondary rhinaria on ANT III only. On Baccharis pilularis and Baccharis sp. in western USA (California, Oregon). A member of the A. carduella group, but with long, finely-pointed antennal hairs.

Aphis ballotae Passerini (= balloticola Szelegiewicz of Blackman & Eastop 2006; see García Prieto et al. 2004) Apterae are dark grey-blue to mottled green (see also influentialpoints.com/Gallery); BL 1.0-2.0 mm. Alatae have secondary rhinaria distributed III 3-12, IV 0-3, V 0. On stems and undersides of leaves of Ballota spp. causing slight downward leaf-curl in early summer (Stroyan, 1984). Also recorded from Dracocephalum, Leonurus and Marrubium spp. Alatae have secondary rhinaria distributed III 3-12, IV 0-3. Monoecious holocyclic, with alate males (Börner 1950; Tuatay & Remaudière 1964). Throughout Europe (except Scandinavia), and eastward to Crimea, Iran and Turkey. There is also a record from Algeria (Laamari et al. 2013). A member of the A. frangulae complex, difficult to distinguish except by biology and host plant.

Aphis berberidorum Ortego & Mier Durante Apterae are brown, with antennae and legs partly pale; BL 1.5-2.4 mm. Alata has 6-12 secondary rhinaria on ANT III only. On Berberis spp. in the southern Andes (Chile and Argentina). Males are alate (Ortego & Mier Durante 1997). Apparently a member of the craccivora group.

Aphis berlinskii Huculak Apterae are matt brownish green to dark green, siphunculi dark, cauda also dark but somewhat paler than siphunculi; BL 1.1-1.4 mm. Alata has 6-8 secondary rhinaria on ANT III only. On Viburnum spp., forming colonies on roots and underground runners, or tented over by ants. Only known from Poland. Monoecious holocyclic, with oviparae and apterous males in late August-September (original description).

Aphis berteroae Szelegiewicz Apterae have a dark green abdomen, rest of body blackish green or blackish; BL 1.25-1.6 mm. Alata has 3-5 secondary rhinaria on ANT III. In small, ant-attended colonies on roots of Berteroa incana, feeding on smaller roots 30-50 cm below the surface. Only known from Poland. Monoecious holocyclic, with apterous males (original description).

Aphis biobiensis Nieto Nafría & Mier Durante Apterae are shining black; BL 1.3-2.3 mm. Alatae have 6-9 rhinaria on ANT III and 0-1 on IV (0-3 on ANT III in alatiform apterae). On Adesmia viscida in Chile (Biobío region) (Nieto Nafría et al. 2016c), and now also reported (and redescribed) from A. papposa in Argentina (Mendosa province; Nieto Nafría et al. 2021b).

Aphis blackmani Kadyrbekov Apterae are pale greenish with red eyes, with only tips of appendages darkened; BL 1.7-2.2 mm. Feeding on the roots of a species of Tragopogon in Kazakhstan (Almaty region; Kadyrbekov 2019b). Other morphs and biology are unknown.

Aphis boydstoni Pike Apterae are reddish brown to blackish, with dark appendages; BL 1.6-2.9 mm. Alatae have secondary rhinaria distributed III 22-31, IV 0-2. On Ceanothus velitinus in open conifer forests in mountainous regions of north-west USA (Washington, Oregon, Idaho, Nevada) and Canada (British Columbia). Monoecious holocyclic with oviparae collected in September (Pike et al. 2004).

Aphis bozhkoae Eastop & Blackman (replacement name for A. eupatorii Bozhko; Eastop and Blackman 2005) Apterae are yellowish with brown appendages; BL c. 1.9-2.0 mm. Alata has secondary rhinaria distributed III 23-27, IV 12-15, V 4. On Eupatorium cannabinum in Ukraine (Bozhko 1976a, as A. eupatorii).

Aphis brachychaeta Holman Apterae are reddish brown; BL 1.5-2.2 mm. Alata has secondary rhinaria distributed III 5-9, IV 0-2. On (probably) Stellera chamaejasme, colonising upper parts of roots and at bases of stems, attended and sheltered by ants. In Mongolia. Presumed to be monoecious holocyclic.

Aphis brachysiphon Narzikulov Apterae are dark brown; BL 1.3-1.7 mm. On Clematis orientalis in Central Asia (Iran, Tajikistan, Kazakhstan, Uzbekistan, Afghanistan; Remaudière & Remaudière 1997). Generic placement is open to question; Kadyrbekov (2005b) placed this species in Brachyunguis.

Aphis breviseta Holman Apterae are bright yellowish green; BL 1.1-1.9 mm. Alata has secondary rhinaria distributed III 3-6, IV 0(-1). On Potentilla spp., living on basal parts of stems and rhizomes, under earthen sheaths built by ants. In steppe localities in eastern Europe. Monoecious holocyclic, with apterous males (original description).

Aphis brevitarsis Szelegiewicz Apterae are dark green with black siphunculi and cauda; BL 1.8-2.1 mm. On stem of a Mentha sp. in Mongolia.

Aphis brohmeri Börner Apterae are dark purplish brown, BL 1.5-2.6 mm. Immatures have no discrete pleural wax spots (cf. fabae). Alata has secondary rhinaria distributed III 9-18, IV 0-3. Colonies occur low down on stems and leaf sheaths of Anthriscus sylvestris, attended by ants. There are also records from some other Umbelliferae/Apiaceae, e.g. Heracleum and Pastinaca (Chumak 2004), Carum and Seseli (Ivanoskaya 1977), and Aegopodium (Kadyrbekov 2012a), although slide-mounted specimens may easily be confused with A. fabae, and it is possible that some natural hybridisation occurs. Widely distributed in northern and central Europe, and in Iran, western Siberia and Kazakhstan. Monoecious holocyclic, with apterous males (Heie 1986).

Aphis brotericola Mier Durante Apterae are black, often covered with whitish wax powder; BL 1.2-2.2 mm. Alata has secondary rhinaria distributed III 5-16, IV 0-5. Forming dense colonies on leaves of Euphorbia spp. in Spain, Italy (Barbagallo & Patti 1998), France, Greece (Papapanagiotou et al. 2012), Turkey, Iran (Barjadze et al. 2017b) and Morocco. Monoecious holocyclic in Spain, with alate males (García Prieto et al. 2001).

Aphis brunellae Schouteden Apterae are yellowish or whitish, sometimes with a mid-dorsal suffusion of dark green (see influentialpoints.com/Gallery); BL 1.0-1.6 mm. Alatae have secondary rhinaria distributed III 6-9, IV 2-5, V 0-1. On stems, in bracts of inflorescences and on undersides of leaves of Prunella spp., attended by ants. Widely distributed in Europe, and in western Siberia (Altai; Stekolshchikov et al. 2008a). Monoecious holocyclic, with apterous males; oviparae have markedly swollen hind tibiae bearing c.100 scent glands. In spite of clear biological differences, the mitochondrial COI and COII sequence of a sample identified as A. brunellae in Italy was indistinguishable from that of A. gossypii (Cocuzza et al. 2009).

Aphis brunnea Ferrari Apterae are brown-black; BL 1.7-2.5 mm. Alatae with secondary rhinaria distributed ANT III 4-13, V 0-3. On shoot apices of Ononis spp. (especially natrix) in the Mediterranean area and adjacent countries; France, Spain, Italy, Austria, Slovakia, Bulgaria, Rumania, Israel, Jordan and Lebanon. In Spain it has been found on Anthyllis cytisoides (Nieto Nafría et al. 2005). The life cycle is unknown, perhaps anholocyclic in southern Italy (Barbagallo & Stroyan 1982). 2n=8.

Aphis bupleuri (Börner) Apterae are brown-black covered with greyish-white wax powder; BL 1.3-1.9 mm. Alatae have secondary rhinaria distributed III 17-41, IV 6-15, V 3-10. On stems, lateral shoots and inflorescences of Bupleurum spp. in Europe (France, Spain, Germany, Poland, Italy, Hungary, Czech Republic, Rumania, Ukraine). One of a group of closely related wax-covered Aphis species on Bupleurum, the others being caroliboerneri, funitecta and talgarica. Nieto Nafría et al. (1986) tabulated morphological differences, and Kadyrbekov (2001a) provided a key. Monoecious holocyclic with alate males, and oviparae with strongly swollen hind tibiae bearing more than 200 scent glands. A. bupleurisensoriata Bozhko is a synonym (Kadyrbekov 2001a).

Aphis cacaliasteris Hille Ris Lambers Apterae are plump-bodied, matt grey-black with black appendages; BL 2.0-3.1 mm. Apterae have secondary rhinaria, distributed III 0-20, IV 7-15, V 0-3, and alatae have them distributed III 23-40, IV 12-20, V 0-3. On Senecio spp. and related genera of tubuliferous Compositae (Adenostyles, Doronicum) in south and central Europe, and recently recorded from western Siberia (Altai Republic; (Stekolshchikov & Novgorodova 2013). Monoecious holocyclic, with oviparae and alate males in July-September. Lampel (1984) gave a full account.

Aphis calaminthae (Börner) Apterae are pale to dark green with brownish head; BL 1.0-1.1 mm. Alatae have secondary rhinaria distributed ANT III 5-7, IV 0-2. In loosely rolled leaves and inflorescences of Clinopodium spp. formerly placed in Calamintha and Acinos (acinos, menthifolium, grandiflorum, sylvaticum), and also recorded from Hyssopus officinalis. Jörg & Lampel (1988) noted differences in the morphology of samples from Acinos arvensis (= Clinopodium acinos) and Hyssopus officinalis. Ant-attended. In southern Germany, Austria, Switzerland, Hungary and Czech Republic. Lampel & Meier (2007) recorded it also from Italy and Poland. Life cycle and sexual morphs are unknown.

Aphis caliginosa Hottes & Frison Small, brownish aphids with dark legs and antennae, in ant-attended colonies on ends of branches of Cornus racemosa in eastern North America (Illinois, Pennsylvania, North Carolina, New Jersey). Life cycle is unknown (Robinson & Chen 1969).

Aphis callunae Theobald Apterae are small, plump-bodied, blackish, brownish violet or dark brown, with wax powder on dorsum arranged in hexagonal fields, and dark brown to black appendages (see influentialpoints.com/Gallery); BL 1.0-1.5 mm. Immatures are greenish. Alatae have secondary rhinaria distributed ANT III 3-7, IV 0-3. On terminal shoots and inflorescences of old, straggly plants of Calluna vulgaris, well-camouflaged and easily overlooked. In western, northern and central Europe, and also recorded from British Columbia, Canada. Monoecious holocyclic on Calluna, with oviparae and alate males in September in northern Germany (alate, brachypterous. and apterous males were obtained in culture; Müller 1976b).

Aphis calycerarum Nieto Nafría, Ortego & Mier Durante Apterae are dark green to black covered with a faint ashen wax powder, with antennae and legs mainly pale, darkening towards apices; BL 1.3-1.9 mm. Alatae have secondary rhinaria distributed ANT III (1-)4-8, IV 0-3. On two species of Calyceraceae, Calycera spinosa and Boopis anthemoides, in Argentina (Nieto Nafría et al. 2021c).

Aphis (Zyxaphis) canae Williams (Fig.12v) Apterae are green with darker head and dorsal abdominal patch or mottling, and white wax markings including a transverse band on anterior abdomen; BL c.1.6 mm. On stems and leaves of Artemisia and Seriphidium spp. in western USA (see Palmer 1952). There is some confusion in the literature with A. hermistonii and A. oregonensis, but the three species seem separable by the key characters given.

Aphis carduicola Stekolshchikov Apterae are bluish green with end of abdomen yellowish green, or pale yellowish green; BL 1.1-1.9 mm. Alatae have secondary rhinaria distributed III 3-7, IV 0-1. At base of stem and on petioles of basal leaves of Carduus spp. in southern Russia (Astrakhan province) and Georgia (Stekolshchikov 2005), and also recorded from Turkey (Şenol et al. 2014c). Monoecious holocyclic; fundatrices were collected in April-May on C. uncinatus.

Aphis cari Essig Apterae are transparent white, yellow or pale yellowish green with distinctive dark green dorsal markings, including a transverse bar between the siphunculi; BL c.1.7 mm. Alatae have secondary rhinaria distributed III 15-50, IV 0-4. On stems of Carum kelloggii, and also recorded from Angelica tomentosa, in California, USA. Only known from the original description, and it seems possible that these are hot weather dwarfs of Aphis carduella (q.v.).

Aphis caroliboerneri (Remaudière) Apterae are grey covered with dense waxy secretion; BL 1.0-1.3 mm. Alatae have secondary rhinaria distributed III 7-16, IV 1-5, V 0-3. In small colonies on flowerheads of Bupleurum fruticosum. Only known from France and Spain.

Aphis carrilloi Ortego, Mier Durante & Nieto Nafría Apterae are dark green, with varable dorsal sclerotisation; BL 1.5-2.8 mm. Monoecious holocyclic on Gunnera magellanica in Chile (Maule). Fundatrices, apterous fundatrigeniae, apterous viviparae and one alate vivipara were collected in January-February (Ortego et al. 2013).

Aphis carverae Hales, Foottit & Maw Apterae are black with antennae and legs partly whitish; BL 0.9-1.5 mm. Alatae are unknown. In small colonies on growing tips of Epilobium billardierianum in Australia (New South Wales). Monoecious holocyclic, with oviparae and apterous males in March-April (Hales et al. 2014)

Aphis caryopteridis Holman Apterae are dark green with dark siphunculi and cauda; BL 1.0-1.5 mm. Alatae have 3-5 secondary rhinaria on ANT III. On stems of Caryopteris mongholica, ant-attended. In Mongolia.

Aphis catalpae Mamontova Apterae are rich yellow to yellow-green, with a dark green spot on dorsal abdomen in life; BL c.1.6-1.7 mm. On undersides of leaves, causing strong wrinkling, and along petioles, of Catalpa spp. (bignonioides, speciosa), in Ukraine; also reported from Hungary, shrivelling leaves of C. bignonioides and producing much honeydew (Ripka 2001), Italy (Cocuzza et al. 2009), Turkey, Iran, Uzbekistan (Davletshina 1964), Kazakhstan (Kadyrbekov 2017a) and east Siberia (Pashchenko 1997). The far eastern population is dark green and has longer siphunculi. Holocyclic on Catalpa with alate males (Davletshina 1964), but reported to migrate facultatively to herbs and grasses in Ukraine (Mamontova 1953). [The host association is a mystery as the two Catalpa species from which it is recorded are native to North America, yet the aphid is only recorded from Eurasia. Populations on Catalpa in Italy identified as this species have mitochondrial COI and COII sequence data very close to that of A. gossypii (Cocuzza et al. 2009).]

Aphis ceanothi Clarke Apterae are dull reddish or amber-brown, with black head and black markings posterior to siphunculi; BL 1.0-2.3 mm. Alatae have secondary rhinaria distributed III 11-36, IV 0-5, V 0-1. On flower stems and leaves of Ceanothus spp. in western USA and Canada; also found (in California) on Noltea africana (BMNH collection, leg. E.O. Essig). Monoecious holocyclic; oviparae and apterous males were collected in September in Manitoba, Canada (BMNH collection, leg. A.G. Robinson).

Aphis celastrii Matsumura (synonym of spiraecola acc. to Moritsu 1983). Apterae are “dark brown with a greenish shade” according to the original description. There is doubt about the identity of this species. The name has since been applied to yellow-green aphids on Celastrum, Deutzia, Ilex and Polygonum in Japan (Higuchi & Miyazaki 1969) and Korea (Lee et al. 2002c) which, according to the description given by Takahashi (1966), and slides in the BMNH collection determined as celastrii by Takahashi, are not clearly distinct from A. spiraecola. Oviparae were collected on Deutzia by Takahashi (1966). 2n=8.

Aphis cephalanthi Thomas Apterae are dark purplish brown to almost black, wax-dusted (see influentialpoints.com/Gallery); BL 0.9-1.7 mm. Alatae have secondary rhinaria distributed III 9-15, IV 0-5. On leaves, stems and flowerheads of Cephalanthus occidentalis, widely distributed in North America, and also in Cuba (as Schizaphis sp.; Holman 1974). A record from Impatiens balsamina in South Africa (Millar 1990) perhaps needs additional confirmation. Well-developed marginal tubercles are present on most segments and, atypically for an Aphis, the tubercles on abdominal tergite 7 are posterior to, and on a level with, the spiracles. Oviparae and alate males occur in late September (in New York; BMNH collection, leg. L.L. Pechuman).

Aphis cephalariae Barjadze Apterae are black except for pale ANT III, IV and basal half of tibiae; BL 1.6-2.7 mm. Alatae have secondary rhinaria distributed III 19-31, IV 0-8, V 0-2. Living in dense, ant-attended colonies on stems of Cephalaria gigantea in Georgia (Barjadze 2011b), and now also recorded from Dipsacus sp. in Turkey (Özdemir & Barjadze 2015). Life cycle is unknown but probably monoecious and holocyclic.

Aphis cerastii (Börner) Apterae are dark grey, BL 1.2-1.6 mm. On subterranean parts of Cerastium arvense in Germany. Males are apterous.

Aphis cercocarpi Gillette & Palmer (= Aphis coweni Palmer, = Aphis holodisci Robinson; new synonymies by Jensen 2021) Apterae are shining greenish black to lead grey with antennae and legs mainly pale, and siphunculi and cauda black; BL 1.0-2.3 mm. Alatae have secondary rhinaria distributed III 15-34, IV 0-13, 0-1. Heteroecious holocyclic, migrating from primary hosts Holodiscus spp. to form ant-attended colinies on the secondary hosts, Veratrum spp.; this has been confirmed experimentally by host transfers (Jensen 2021). However apterous males are produced in some locations, indicating that there is either life cycle variation or that two cryptic species may be involved. In western USA and Canada, and apparently also on various Veratrum spp. in east Siberia (Pashchenko 1988a, as A. veratri). The original description of A. cercocarpi and subsequent records from Cercopus montanus in western USA seem to be based on host misidentification (Jensen 2021). [There is another undescribed Aphis sp. on Veratrum in North America; apart from the discriminants in the key to apterae on Veratrum, other differences between this species and A. cercocarpi are tabulated by Jensen (2021)].

Aphis (Toxoptera) chaetosiphon (Qiao) Colour of apterae in life is unrecorded; BL 1.7-2.0 mm. On new shoots of Camellia oleifera, attended by ants. In Guangxi Province, China (Qiao et al. 2008).

Aphis chetansapa Hottes & Frison Apterae are yellowish-brown to yellowish-red with darker antennae and legs and blackish siphunculi, and dark markings on posterior abdominal tergites. Alatae have only 4-8 secondary rhinaria confined to the third antennal segment, dark wing veins, and often have dark tranverse bands across last three abdominal tergites (Hottes & Frison 1931). In tightly curled terminal leaves of Pyrus sp. and Prunus sp., recorded from Illinois, Missouri and Oregon, USA. The description of this species closely resembles that of A. longicauda. Life cycle is unknown.

Aphis chilopsidi Davletshina Apterae are pale green to dark greenish brown, wax-dusted, with dark siphunculi and pale cauda; BL c.1.6-1.7 mm. In small colonies on leaves and petioles of Chilopsis linearis in Uzbekistan.

Aphis chloris Koch Apterae are pale green or lemon yellow, or in at lower temperature olive green, with dark head, apices of antennae and legs, and siphunculi (see also influentialpoints.com/Gallery); BL 1.0-1.9 mm. Alatae have 5-9 secondary rhinaria on ANT III and 0-2 on IV. On Hypericum spp., usually living in ant-attended colonies at stem bases just below soil surface, occasionally on aerial parts. Widespread in Europe, across Asia to China, and introduced for control of Hypericum perforatum to South Africa (Durr 1983), Australia (Briese & Jupp 1995) and Canada (Foottit et al. 2006). Monoecious holocyclic, with apterous males. Wilson (1938) made one of the earliest studies of induction of sexual morphs using this species. Aphis citrina Nevsky, described from Central Asia as a subspecies of A. chloris and subsequently given full species status, seems likely to be a synonym. 2n=8.

Aphis (Zyxaphis) chrysothamni Wilson (Fig.21j) Apterae have wine-coloured head and pale green to pinkish abdomen with (usually) a dark spot anteriorly, and some wax at the end; BL 1.0 (summer dwarfs) – 2.4 mm. Alatae have secondary rhinaria distributed III 4-10, IV 0-5, V 0.1. On Chrysothamnus nauseosus in western North America. Oviparae and apterous males appear in late September in Utah (BMNH collection, leg. G.F. Knowlton).

Aphis (Zyxaphis) chrysothamnicola (Gillette & Palmer) (Fig.21k) Apterae are dusky reddish brown or dull blackish green with dark bands on posterior abdomen; BL 1.7-2.1 mm. Alatae have 3-7 secondary rhinaria on ANT III. On leaves and stems of Chrysothamnus nauseosus in western USA (Rocky Mountain region). Oviparae and apterous males in October.

Aphis ciceri Müller (Fig.16h) Apterae are dark green, appearing black, shiny on dorsum; BL 2.2-2.6 mm. On lower part of stem of Astragalus cicer, just above the soil surface, attended by ants. Only known from Germany (Jena), Iran (Rezwani et al. 1994) and Kazakhstan (Kadyrbekov 2017a). Oviparae and apterous males were obtained in September-October (original description).

Aphis cimicifugae Holman Apterae are grass green with black siphunculi and cauda; BL 1.8-2.3 mm. Alatae have 3-10 secondary rhinaria on ANT III. On Actaea (= Cimicifuga) spp., on upper parts of stems and in inflorescences, attended by ants (Pashchenko 1994b). In east Siberia.

Aphis cinerea Nieto Nafría & Ortego Apterae are black, thickly coated with whitish grey wax; BL 1.7-2.5 mm. Alatae have 3-8 secondary rhinaria on ANT III. In dense colonies on Lathyrus macropus in Argentina (Nieto Nafría & Ortego 2002). Life cycle and sexual morphs are unknown.

Aphis cirsiioleracei (Börner) Apterae are brownish grey, lightly wax-dusted; BL 1.2-1.6 mm. On basal parts, and later in upwardly rolled leaves of Cirsium spp., in central and eastern Europe (Germany, Switzerland, Czech Republic, Poland; also in northern Italy), and also recorded from Kazakhstan (Kadyrbekov 2014f), where there is a record from Cousinia microcarpa. Monoecious holocyclic according to Börner (1952). It has been suggested that A. viridescens Bozhko, described from Cirsium elodes in Ukraine is a synonym, although Bozhko’s species was pale green in life.

Aphis cirsiphila Pashchenko Colour of apterae in life unknown; BL c.2 mm. In dense colonies on leaves and stems of Cirsium spp. in east Siberia. A record from Turkey (Görür et al. 2017) needs further confirmation. Monoecious holocyclic.

Aphis cisticola Leclant & Remaudière Apterae are yellow to bluish green, with light dusting of wax, dark apices to legs and antennae, and black siphunculi and cauda; BL 1.0-1.8 mm. Alatae have 3-7 secondary rhinaria on ANT III, 0(-2) on IV. On young growth of Cistus spp., especially C. salvifolius, forming dense colonies on leaves shoots and flower stems, attended by ants. In southern France, Corsica, Italy and Spain. Barbagallo & Stroyan (1982) tabulated morphometric data for comparison with A. lupoi.

Aphis (Toxoptera) citricidus (Kirkaldy) Tropical Citrus Aphid Plate 8c Apterae are shining, very dark brown to black, usually larger than A.(T.) aurantii, and with antennae less distinctly banded (see influentialpoints.com/Gallery); BL 1.5-2.4 mm. Immatures are brown. Alatae have a shiny black abdomen, black ANT III, forewing with pale pterostigma and media twice-branched; they have secondary rhinaria distributed ANT III 6-20, IV 0-4. (Specimens in alcohol colour the fluid deep red.) In ant-attended colonies on young growth of host plants, rolling leaves and stunting shoots. Mainly on Rutaceae, especially Citrus, but occasionally large colonies develop on young growth of other trees and shrubs (see also Blackman & Eastop 2000, p. 355). In southern Africa, southern Asia, Australia, New Zealand, Pacific islands and subtropical and warm temperate parts of South America (but records from Chile seem to be unfounded – see Nieto Nafría et al. 2015a). More recently it has spread to important citrus-growing areas in Central America, the Caribbean and southern USA (Yokomi et al. 1994, Halbert & Brown 1996), and to Madeira, Portugal and north-western Spain (Hermoso de Mendoza 2008), but not yet to the Mediterranean region and Middle East. It thrives in moist warm climates and seems able to tolerate colder conditions than A.(T.) aurantii, for example occurring at higher altitudes, but it is not found in regions with long hot dry seasons. Apparently it is anholocyclic everywhere except Japan, where there is a sexual phase (Komazaki 1988). Michaud (1998) reviewed the literature on this aphid, which includes laboratory studies of population parameters (Galatoire 1983, Takanashi 1989); field studies in Japan (Komazaki 1988) and Venezuela (Rondón et al. 1983 – including natural control by fungus and predators); and a review of transmission of citrus tristeza virus (Roistacher & Bar-Joseph 1987). Zhao et al. (2021) found differences in host selection and performance among seven common Citrus species in south-east China. Shang et al. (2021) studied the role of long non-coding RNAs in wing development. An exotic aphidiid parasitoid Lipolexis oregmae has been used for biological control in Florida (Persad et al. 2007). Qureshi (2010) examined the potential impact of climate change on populations on citrus in Florida. Nieto Nafría et al. (2005b) established that citricidus is the correct specific epithet for this aphid. 2n=8.

Aphis clematicola Pashchenko Apterae are yellow-green, wax-dusted; BL c. 1.5 mm. On Clematis fusca in east Siberia, forming large dense colonies on undersides of leaves, attended by ants.

Aphis clematidis Koch Apterae are dark olive green with a blackish spinal stripe, marbled with white wax powder; BL 1.7-2.3 mm. Smaller, paler specimens may occur in midsummer. Alatae have 5-9 secondary rhinaria on ANT III. On young shoots, flower stems and leaves of Clematis spp. in central and southern Europe. Pigmentation and length of siphunculi show great variation, and there has been confusion with A. vitalbae; reliable differences from that species are the longer pointed hairs on the appendages and the longer hind tarsal segment II (Tashev 1964b). Monoecious holocyclic (Börner 1952). 2n=8.

Aphis clematiphaga Pashchenko Apterae are bright green and dark green, without wax markings; BL c. 2.2 mm. On stems, flower shoots and undersides of leaves of Clematis spp. in east Siberia. Monoecious holocyclic. Visited by ants.

Aphis clerodendri Matsumura Apterae are straw-coloured to bright yellow green or olive-green; BL 1.1-1.9 mm. On Clerodendrum spp. in east Asia (Japan, Korea, Vietnam, Laos, Taiwan, Philippines, Papua New Guinea) and Australia (Carver et al. 2003), forming loose leaf pseudogalls. Alatae have 3-8 secondary rhinaria on ANT III. Very close to A. gossypii, and the characters that separate the two species in east Asia may not be reliable for other parts of the world. Takahashi (1966) described a subspecies, A. clerodendri amamiana, specific to C. yakusimense (= C. trichotomum var. fargesii) in Japan, but this seems indistinguishable from A. gossypii. 2n=8.

Aphis cliftonensis Stroyan Apterae are rather bright pale green; BL 0.8-1.2 mm. On roots of Helianthemum nummularium, in ant shelters. In England and Sweden (BMNH coll. VFE). Monoecious holocyclic with apterous males (Stroyan 1984).

Aphis clinepetae Pashchenko Apterae are pale yellow-green or dark green, usually wax-dusted; BL c. 1.5 mm. Alatae have secondary rhinaria distributed III 9-18, IV 2-6, V 0-2. In small, dense colonies on stems and flowers of Nepeta manchuriensis, and more rarely on Clinopodium chinense, in east Siberia. [Aphids from Calamintha grandiflora in Crimea (BMNH collection, leg. J. Holman) have apterae intermediate in morphology between this species and clinopodii, but include an alata with secondary rhinaria distributed III 8 and 11, IV 3 and 3, so are possibly this species.]

Aphis clinopodii Passerini Apterae are blackish green; BL 0.9-1.2 mm. Alatae have 2-5 secondary rhinaria on ANT III. Curling and distorting terminal leaves of Clinopodium spp. in Europe (England, France, Spain, Italy, Germany, Poland, Austria, Czech Republic, Ukraine). One of a closely-related group of European species on Lamiaceae, and very similar to A. serpylli (but with a longer antennal terminal process) and A. origani (but with shorter siphunculi). Monoecious holocyclic. Several colonies in south-east England were all attended by the ant Myrmica sabuleti (Wood-Baker 1979). 2n=8.

Aphis clydesmithi Stroyan Apterae are dull green; BL 1.1-1.3 mm. Alatae have 2-3 secondary rhinaria on ANT III. On Sanicula canadensis, living at base of stem and on root collar, attended by ants. In North Carolina, USA.

Aphis coffeata Mamontova Apterae are coffee brown with a pale cauda; BL c. 1.8 mm. Secondary rhinaria in alata (one specimen) distributed III 12-13, IV 0-2. In colonies on stems and in flowerheads of Melampyrum nemorosum and M. pratense in Ukraine, Byelorussia and western Russia. It was redescribed by Stekolshchikov et al. (2007b).

Aphis comari Prior & Stroyan Apterae are blue-green, very dark green or bluish black; BL 1.0-1.9 mm. Alatae have 4-8 secondary rhinaria on ANT III. On shoots or leaves of Comarum palustre growing in sphagnum bogs, sometimes in ant shelters built from plant debris. In northern and north-west Europe (UK, Denmark, Finland, Kola Peninsula, Poland), and also recorded from Kazakhstan on Sibbaldianthe (= Potentilla) bifurca and Potentilla ?virgata (= dealbata) (Kadyrbekov 2009b, 2014d, 2017a). Monoecious holocyclic, with apterous and brachypterous males (original description).

Aphis commensalis Stroyan Apterae 1.4-1.7 mm, dark grey-green to blackish-green, heavily clothed in mealy greyish wax powder (see influentialpoints.com/Gallery), living in late spring and summer within empty leaf-edge pocket-galls of the psyllid Trichopsylla walkeri on leaves of Rhamnus cathartica in Europe, and also in Kazakhstan (Kadyrbekov 2017a). Earlier in the year fundatrices amd fundatrigeniae cause rolling and twisting of leaves (see Pérez Hidalgo et al. 2010). Small apterous males and oviparae mature within the gall in September-October, and eggs are laid within the gall (Stroyan 1952, or for a fuller account, Böhm 1963). Alatae are unknown.

Aphis comosa (Börner) Apterae are brown with shining black dorsum, antennal flagellum and middle sections of tibiae pale; BL 1.5-2.4 mm. Alatae have 4-9 secondary rhinaria on ANT III. On Melilotus albus and Lathyrus pratensis in Europe (France, Germany, Austria, Italy, Spain). Also recorded from Latvia, Poland and Romania by García Prieto et al. (2004), but their record of its occurrence in UK appears to be erroneous. Some records may apply to A. pseudocomosa (q.v.). Redescribed by Hoffmann (1972).

Aphis conflicta Nieto Nafría, Ortego & Mier Durante Apterae are shing black or dark reddish brown, with some whitish wax powder, antennae and tibiae being predominantly yellow; BL 1.2-2.3 mm. Monoecious holocyclic on Ochetophila (= Discaria) trinervis in Argentina, and also recorded from species in several other genera of Rhamnaceae (Colletia, Condalia, Discaria, Trevoa), and also on Tristerix (Loranthaceae), in Chile and Argentina. Nieto Nafría et al. (2008) described all morphs including fundatrices (collected in November), oviparae and apterous males (early April). A sample collected in January at Neuquén (Argentina) included small (aestivating?) apterae. López Ciruelos et al. (2017b) reviewed morphological variation, host plants and distribution.

Aphis confusa Walker Apterae are pale yellow, yellowish green, green or dark green, depending on location on host; BL 1.0-2.3mm. Alatae have secondary rhinaria distributed III 3-7, IV 0-1. On Dipsacaceae (Dipsacus, Knautia and Scabiosa, occurring both as larger green or dark green aphids on upper parts of stems and inflorescences, or in summer as smaller and paler yellow or yellower green specimens on undersides of leaves, at base of stems or on roots (Heie, 1986). Muir (1959) recorded it from Succisa pratensis. Ant-attended. Widely distributed in Europe, in Central Asia (Kadyrbekov 2017a), and Remaudière et al. (2006) reported its occurrence in Turkey, Iran and Mongolia. Monoecious holocyclic, with apterous males. Kanturski & Bezdĕk (2019) provided a detailed redescription including sexual morphs. A very variable species, and there is no reliable way of distinguishing it morphologically from the equally variable polyphagous A. gossypii, which may sometimes occur on Dipsacaceae. A. gossypii is, however, unlikely to occur on basal parts of the plant, and is less assiduously ant-attended. In A. gossypii there is also usually more contrast between the pigmentation of siphunculi and cauda.

Aphis conspicua Nieto Nafría & Mier Durante Apterae are matt yellowish grey to matt black; BL 1.1-1.9 mm. In compact groups on stems of Baccharis spp. (linearis, salicifolius). Alatae seem rare; the one known specimen has 6-7 secondary rhinaria on ANT III and 0-1 on IV. In Argentina (Neuquén) and Chile (Maule). Sexual morphs are unknown (Nieto Nafría et al. 2019c).

Aphis coprosmae Laing Apterae are in life covered in fluffy white wax); BL c. 1.6 mm. Alatae are remarkable in having only 0-1 secondary rhinaria on ANT III. On Coprosma spp. in New Zealand (Cottier 1953, Teulon et al. 2013). Monoecious holocyclic; eggs are laid on stem of plant and hatch in September. DNA analysis has revealed an nnusually high level of genetic diversity for a single species (Teulon et al. 2013).

Aphis cordubensis Ortego, Mier Durante & Nieto Nafría Apterae are dark green to greenish black, slightly powdered with white wax; BL 1.4-1.7 mm. On cultivated specimens of Abutilon megapotamicum in Córdoba, Argentina, and possibly in other South American localities where this plant is cultivated (Ortego et al. 2021). Monoecious holocyclic, with oviparae and alate males collected with apterous and alate viviparae in April and included in the original description.

Aphis coreopsidis (Thomas) Plate 6c Apterae 1.5-1.8 mm, yellow to green with darker antennae, legs and siphunculi (see influentialpoints.com/Gallery), on stems and leaves of new shoots of Nyssa sylvatica in spring. Apparently host-alternating in North America (Illinois) between Nyssa and secondary hosts in Compositae/Asteraceae (Bidens, Clibadium, Eupatorium, Sonchus), Malvaceae (Hibiscus, Sida) and Lamiaceae (Blephilia, Nepeta). Oviparae in autumn on Nyssa are small (1.1-1.2 mm), yellowish-green; males are alate, about 1 mm, dark green and brown (Hottes & Frison 1931). The fundatrices on the primary host are still undescribed. Presumed anholocyclic populations occur on secondary hosts (especially Bidens pilosa) in Central and South America and Hawaii, and alatae identified as this species have been trapped in Ghana, Uganda (BMNH collection) and Saudi Arabia (Hussain et al. 2015a).

Aphis coridifoliae Mier Durante & Ortego Apterae are pale or greenish yellow powdered with white wax, with brown-black siphunculi and cauda, and frequently with dark brown intersegmental markings on dorsal abdomen; BL 1.0-1.7 mm. Alatae have 4-8 secondary rhinaria on ANT III. In small dense colonies on Baccharis coridifolia, and perhaps on other Baccharis spp., in Argentina (Mier Durante & Ortego 1999).

Aphis cornifoliae Fitch Apterae are small (1.0-1.4 mm), dark brown to dull greenish-black, on both upper and undersides of leaves of Cornus spp., living without host alternation; males are apterous (Palmer 1952). Widely distributed in USA and Canada. Robinson & Chen (1969) discussed differences from A. carduella (as helianthi). A member of the asclepiadis group (Lagos et al. 2015).

Aphis cornuta Blackman & Brightwell Apterae are bright yellow to pale yellow-green with pale green head and prothorax, black siphunculi and a dusky cauda (see influentialpoints.com/Gallery); BL 0.9-1.8 mm. living without host alternation at stem bases of Picris (= Helminthotheca) echioides, attended by ants. Oviparae and apterous males were collected in September (Blackman et al. 2019a). Only known from southern England.

Aphis coronillae Ferrari Apterae are dark brown to brownish green with grey-black dorsal shield (not wax-powdered; see influentialpoints.com/Gallery); BL 1.3-2.2 mm. Alatae have secondary rhinaria distributed III 3-9, IV 0-2. Living on basal parts of certain Leguminosae/Fabaceae (Coronilla, Glycyrrhiza, Securigera, Trifolium, Medicago), sheltered by ants. Widely distributed in Europe, and also in west Siberia (Stekolshchikov et al. 2008a) and Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic, with males (mostly apterous, occasionally brachypterous or alate) and oviparae appearing in September. Hoffmann (1968) studied host relationships and described populations on Medicago lupulina as a subspecies (A. coronillae arenaria Hoffmann, as scaliai ssp. arenaria). Stroyan (1984) discussed morphological variation in relation to geography and host plant. A. yangbajaingana Zhang, in Tibet on Medicago, is very similar but has shorter, thicker siphunculi.

Aphis coronopifoliae Bartholomew Apterae are green, BL 1.4-1.7 mm. Alatae have secondary rhinaria distributed III 11-17, IV 2-7. On Cotula coronopifolia in California, USA. It seems likely, as suggested by Cook (1984a), that this species is A. mimuli (q.v.).

Aphis (Bursaphis) costalis Cook Colour of aptera in life unknown, probably pale green or yellowish green; BL 1.5-1.8 mm. On Mimulus spp. in California, USA (Hille Ris Lambers 1974b, as mimuli Oestlund). It is a member of the oenotherae group, which typically host-alternate between Ribes and Onagraceae. An aphid in central Washington identified as this species is believed to migrate from Ribes aureum to Mimulus and Veronica, but there are possibly other similar species in that region, including one that is monoecious on Ribes cereum (see aphidtrek.org).

Aphis cottieri Carver Apterae are brown-black, shining; BL 2.2-2.7 mm. Alatae are without secondary rhinaria. On leaves and stems of Muehlenbeckia spp. in New Zealand. Oviparae in April-June, males are unknown (Carver 2000, Teulon et al. 2013). Podmore et al. (2018) reported on genetic variation in mitochondrial COI sequence.

Aphis craccae L. Apterae are black, covered in grey wax powder (see influentialpoints.com/Gallery); BL 1.9-2.8 mm. Alatae have 5-13 secondary rhinaria on ANT III. Living in often dense, ant-attended colonies on terminal growth, flowers and seed-pods of vetches (Vicia spp., especially V. cracca). Records from other hosts may be based on confusion with other “black-backed” Aphis species. Widely distributed in Europe, eastward to China, Japan and Korea, and also in north-eastern USA and Canada. Oviparae and alate males appear in September (Heie 1986).

Aphis craccivora Koch Plate 6d (Fig.41k) Apterae are shining black, with immatures lightly dusted with wax, usually ant-attended (see influentialpoints.com/Gallery); BL 1.4-2.2 mm. Alatae have 2-10 secondary rhinaria on ANT III. Colonising young growth of numerous plants, particularly Leguminosae/Fabaceae; plants in other families tend to be colonised more in the dry season. A major pest of leguminous crops (Blackman & Eastop 2000, p. 227). Worldwide, but particularly common in warm temperate and tropical regions. Anholocyclic almost everywhere, but a sexual phase with alate males has been reported from Germany, India and Argentina. Rhopalosiphum esculentum Raychaudhuri & Raychaudhuri, described from Manihot esculenta in southern India, is a possible synonym (see also Skvarla et al. 2019). Wongsa et al. (2017) have studied variation in COI sequence of populations of A. craccivora on leguminous crops in Thailand. A long-haired variant on Robinia in east Asia has been distinguished as a subspecies, A. craccivora ssp. pseudoacaciae Takahashi (1966). Mehrparvar et al. (2012) found morphometric differences between populations in Iran on different host plants, a population from Robinia again being the most distinct. The discovery that populations on Medicago and Robinia have different endosymbionts (Brady & White 2013) provides further confirmation that two distinct taxa are involved. Mitochondrial DNA (COI) analysis of numerous samples of A. craccivora group by J. Wang et al. (2011) also supports the existence of a separate taxon on Robinia. All 37 mitochondrial genes of the form on Robinia have been sequenced (Song et al. 2016). [If populations on Robinia need a new name then A. robiniae Macchiati 1885 is available.] 2n=8.

Aphis crassicauda Smith & Eckel Apterae are reddish tan to light brown, dusted with wax; BL 1.1-2.1 mm. Alatae have 7-14 secondary rhinaria on ANT III. On Viburnum spp. in North Carolina, USA. Monoecious holocyclic, with oviparae and apterous males in September-October (original description).

Aphis crepidis (Börner) Apterae are dark bluish green to yellow-green, not wax-powdered (see influentialpoints.com/Gallery); BL 1.15-1.95 mm. Alatae have 3-7 secondary rhinaria on ANT III. On Crepis spp., living in ant shelters at base of plant, and also once found on Picris echioides (in Italy: BMNH collection, leg. H.L.G. Stroyan and D. Hille Ris Lambers). In west, central and south Europe, including Iberian peninsula, and also recorded from Iran (Mokhtari et al. 2012). Monoecious holocyclic; oviparae in September, males apparently undescribed. 2n=8.

Aphis crinosa (Paik) Apterae yellowish brown with white wax secretion (Shinji 1923, as Pterocomma ligustri); BL 2.3-2.7 mm. On trunks and branches of Ligustrum spp., and also recorded from Lonicera and Viburnum. In Japan and Korea (Lee et al. 2002).

Aphis curtiseta Holman Apterae are green to dark green; BL 1.0-1.7 mm. On Hieracium spp., living mostly on basal parts of plants, and in early summer on inflorescences. Ant-attended. In Austria, Czech Republic, Slovakia, Bulgaria, Moldova and Ukraine. Monoecious holocyclic, with apterous males (Holman 1998).

Aphis cuscutae Davis Apterae pale green, thinly wax-dusted; BL 1.5-2.1 mm. Alatae have antennal segment III swollen with numerous tuberculate secondary rhinaria. Described from Cuscuta epithymum growing on alfalfa in Utah, USA, and also found on Cuscuta sp. growing on Rhus laurina in California (BMNH colln, leg. R.C. Dickson).

Aphis cuyana López Ciruelos & Ortego Apterae are caramel to brown with “dorsal bright spots” (absent in midsummer dwarfs); BL 1.3-2.3 mm. On Gymnophyton polycephalum in Argentina (López Ciruelos et al. 2017a). Alatae have secondary rhinaria distributed ANT III 6-12, IV 0-4.

Aphis cytisorum Hartig Apterae are black, dark brown or very dark green, usually densely covered in grey wax powder, ant-attended (see influentialpoints.com/Gallery); BL 1.5-2.8 mm. On young shoots and petioles, and later on inflorescences and developing seed pods, of woody Leguminosae/Fabaceae (Laburnum, Cytisus, Spartium, Sophora, Calicotome). In Europe eastward to Russia and Turkey, Iran, North Africa, China (as A. sophoricola Zhang); also widespread in North America, and recorded from Peru (Vilca Mallqui & Vergara Cobián 2011) and Argentina (see Mier Durante et al. 2012). Alatae have secondary rhinaria distributed III 4-9, IV 0, V 0 in A. cytisorum s. str., and III 5-9, IV 0-2, V 0-1 in A. cytisorum ssp. sarothamni Franssen, which occurs on Cytisus scoparius in Europe. Stroyan (1984) compared morphometric parameters of the two subspecies. Apterae are almost impossible to separate, but oviparae of ssp. sarothamni have more swollen hind tibiae than cytisorum, with many more scent glands. Males are alate. There is a complex of closely-related species on woody Leguminosae/Fabaceae in Europe, including others with very similar apterae on Cytisus and Chamaecytisus (A. pseudocytisorum, A. zweigelti). 2n=8 (both subspecies).

Aphis danielae Remaudière Apterae are black; BL 1.6-2.5 mm. On Lycium spp. in Chile and Argentina. In Chile colonies also occur regularly on flower buds of a cactus, Echinopsis chilensis (Heie et al. 1996). Monoecious holocyclic, with oviparae and apterous males in April (Remaudière & Ortego 1998).

Aphis dasiphorae Holman Apterae are reddish brown, powdered with greyish wax; BL 1.2-2.4 mm. Alatae have 4-10 secondary rhinaria on ANT III. In dense colonies on terminal parts of shoots of Potentilla (as Dasiphora) spp. in eastern Russia, Kazakhstan (Kadyrbekov 2012a) and Mongolia. Apparently monoecious holocyclic.

Aphis decepta Hottes & Frison Apterae are yellow with dark dorsal markings posterior to siphunculi, dark brown siphunculi and a brown cauda; BL 1.4-2.2 mm. Alatae have secondary rhinaria distributed III 37-48, IV 0-4. On undersides of leaves of Pastinaca sativa (a European plant) in north-eastern USA (Illinois, New York, Pennsylvania). Native hosts are probably Heracleum spp.; it is recorded from H. lanatum in Manitoba, Canada (Rojanavingse & Robinson 1976) and from H. maximum in Illinois (Lagos et al. 2014). A member of the asclepiadis group.

Aphis dianthiphaga Pashchenko Apterae are dark emerald green, BL c.1.4-1.5 mm. In small to medium-sized colonies feeding at bases of stems of Dianthus repens in east Siberia.

Aphis difabioae Nieto Nafría, Ortego & Mier Durante Apterae are matt black, without wax powder; BL 11.3-1.6 mm. On Adesmia papposa (var. papposa) in Argentina (Neuquén province; Nieto Nafría et al. 2021b). Alatae and other morphs are unknown.

Aphis diluta Pashchenko Apterae are pale or bright green; BL c.1.8 mm. Alatae have secondary rhinaria distributed III 11-25, IV 9-17, V 5-12. On stems and under leaves of Potentilla supina, forming large, loose colonies, and sometimes causing leaf deformation. Possibly also on Geum aleppicum. In east Siberia.

Aphis dlabolai Holman Colour of apterae in life unknown, probably black dusted with whitish wax; BL 1.8-2.4 mm. Alatae have 6-13 secondary rhinaria on ANT III only. On Euphorbia sp. in Iran.

Aphis droserae Takahashi Apterae are greenish black, with appendages mainly dark; BL 1.0-1.2 mm. On flowers and undersides of leaves of Drosera spp. in China and Taiwan (Tao 1991), and now recognised to occur also in southern Germany (Barjadze et al. 2017b).

Aphis duckmountainensis Rojanavongse & Robinson Apterae are black, BL 2.0-2.4 mm. Apterae have secondary rhinaria distributed III 3-26, IV 10-20, V 0-6, and alatae have them distributed III 44-53, IV 14-22, V 3-8. On terminal growth of Senecio pauperculus (= Packera paupercula) in Manitoba, Canada.

Aphis ecballii Rusanova Apterae are yellow-brown or dirty yellow, with dark siphunculi and yellow cauda; BL 1.9-2.1 mm. Alatae have secondary rhinaria distributed III 8-13, IV 1-3. In colonies on leaves and shoots of Ecballium in Azerbaijan. Probably a member of the frangulae group; the description does not allow apterae to be differentiated from those of A. gossypii.

Aphis egomae Shinji Apterae are pale yellow to yellow-green or orange, with black siphunculi; BL small, unrecorded. Alatae have 3-6 secondary rhinaria on ANT III. On Perilla frutescens and Salvia splendens, feeding on undersides of young leaves and causing severe leaf-curl. Moritsu (1983) also recorded it from Physostegia sp. In Japan, and also in South Korea (Lee et al. 2002c).

Aphis elatinoidei Nevsky Apterae are pale green, BL 1.2-1.6 mm. On Elatinoides elatine in Uzbekistan (see also Davletshina 1964).

Aphis elegantula Szelegiewicz Apterae are green mottled with darker green, with black siphunculi and cauda; BL 1.75-2.1 mm. Alatae have secondary rhinaria distributed III 3-6, IV 0-1, V 0-1. On Polygonum angustifolium in Mongolia (Szelegiewicz 1964a), and also recorded from south-west Siberia (Kurgan province; see Novgorodova & Stekolshchikov 2013).

Aphis elena Lagos-Kutz & Voegtlin Apterae are dark green, dusted with white wax; BL 1.3-1.6 mm. Alatae have secondary rhinaria distributed ANT III 5-6, IV 1-2. On Pycnanthemum virginianum in Illinois, USA (Lagos-Kutz et al. 2017). Very similar in morphology to A. gossypii, but DNA analysis indicates a closer relationship with another North American Lamiaceae feeder, A. monardae. The life cycle is unknown.

Aphis (Bursaphis) epilobiaria Theobald Apterae are reddish brown to very dark blackish brown or blackish green, with a striking pattern of dense pleural wax bands across the dorsum, except for a spindle-shaped mid-dorsal clear area (see influentialpoints.com/Gallery); BL (1.3)-2.1-2.7 mm. Alatae have secondary rhinaria distributed III 7-13, IV 2-8, V 0-2. On shoot apices and inflorescences of Epilobium hirsutum, and occasionally other Epilobium spp., in Europe (Stroyan 1984; Heie 1986), and also reported to occur in Turkey, Iran, Kazakhstan and Pakistan. Monoecious holocyclic, with oviparae and alate males in September. Rakauskas (2008) found an Oenothera sp. colonised by A. epilobiaria including sexual morphs and eggs in autumn in the Czech Republic, but fundatrices hatching the following spring did not complete their development when reared on Oe. biennis. 2n=8.

Aphis (Bursaphis) epilobii Kaltenbach Apterae are brown or greenish black, rather more uniformly wax-powdered than A. epilobiaria (see influentialpoints.com/Gallery), and smaller; BL 1.3-2.1 mm. Alatae have secondary rhinaria distributed III (5-)9-20, IV 2-10, V 0-4. On upper parts of stems, flowerheads and undersides of upper leaves of Epilobium montanum, and more rarely other species such as E. lanceolatum. Usually without ant attendance. Widespread in Europe, also reported from Iran, Kazakhstan (Kadyrbekov 2017a), and introduced to North America (Foottit et al. 2006). Monoecious holocyclic, with oviparae and alate males in September. 2n=8.

Aphis epipactis Theobald Apterae are very dark green, almost sooty black; BL c.1.3-1.7 mm. On Epipactis spp. in Europe. There are no satisfactory distinguishing characters, and possible synonymy has been suggested with A. fabae (Börner 1952) or, more likely because of the long hairs and frequent presence of marginal tubercles on abdominal tergites 2-4, A. ilicis (Eastop & Hille Ris Lambers 1976) or A. viburni (Stroyan 1984); Epipactis possibly acts as a reserve host for various members of the A. fabae group.

Aphis equiseticola Ossiannilsson Apterae are light green or dark green, BL 1.4-2.0mm. Alatae have secondary rhinaria distributed III 8-11, IV 0-5, V 0. On Equisetum silvaticum in northern Europe (Sweden, Germany). Presumably monoecious holocyclic, with alate males in early August (Heie 1986).

Aphis (Pseudoprotaphis) erigerontis Holman Apterae are dark green dusted with bluish grey wax; BL 1.5-1.9 mm. Alatae have secondary rhinaria distributed III 27-32, IV 10-15, V 7-13, VI base 3-4. In ant shelters on root collar and roots of Erigeron acer in Czech Republic, and found on Conyza canadensis in Poland (Osiadacz & Hałaj 2017); also reported to occur in Finland, Poland, Slovakia, Spain (García Prieto et al. 2004), Iran (Rezwani 2010) and Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic, with apterous males (original description).

Aphis eryngiiglomerata Bozhko Apterae are greenish black, dusted with wax powder; BL 1.6-2.2 mm. Alatae have secondary rhinaria distributed III 5-15, IV 0-3. In ant-attended colonies on Eryngium spp. in eastern Europe (Czech Republic, Poland, Ukraine, Turkey), Kazakhstan (Kadyrbekov 2017a) and south-west Siberia.

Aphis esulae (Börner) Apterae are pale brown, sometimes blackish dorsally especially behind and between siphunculi; BL 1.8-2.3 mm. Alatae have 9-12 secondary rhinaria on ANT III. On stems and inflorescences of Euphorbia esula, and possibly on other Euphorbia spp., in Netherlands (Barjadze et al. 2017b), Austria, Bulgaria, Hungary, west Siberia (Stekolshchikov et al. 2008a), Turkey (Akyürek et al. 2019) and Kazakhstan (Kadyrbekov 2011b). A single aptera from northern England (BMNH collection, leg. Airy Shaw) also appears to be this species. A member of the difficult A. euphorbiae group, distinguished in Börner’s very brief original description by its short hairs and association with E. esula. Szelegiewicz (1962b) provided a redescription on the basis of specimens from Bulgaria, and his interpretation is followed here. However, populations of A. euphorbiae group aphids with short hairs are not restricted to E. esula, and this plant may also be colonised in western Europe by aphids with typical A. euphorbiae morphology, so further work is needed.

Aphis etiolata Stroyan Apterae are pale yellowish , whitish or bluish green; BL 1.4-2.0 mm. Alatae have 3-5 secondary rhinaria on ANT III. On roots of Rumex acetosella in ants’ nests, rather deep in soil. In Europe (England, Netherlands, Sweden, Finland, Poland, Czech Republic, Slovakia). Monoecious holocyclic, with apterous males.

Aphis eucollinae López Ciruelos & Ortego Apterae are yellow, sometimes with brown dorsal markings, and with appendages and siphunculi brownish; BL 1.3-2.3 mm. Alatae have 5-9 secondary sensoria on ANT III only. On Euphorbia collina (= portulacoides ssp. collina) in Argentina. Monoecious holocyclic, with oviparae and apterous males in April (López Ciruelos et al. 2016). There is a close morphological similarity to the palaearctic species A. esulae.

Aphis eugeniae van der Goot Apterae are orange-yellow to brownish-orange with black siphunculi and cauda; 1.3-2.1 mm. Alatae have 5-10 secondary rhinaria on ANT III, and 0-1 on IV. Resembling A. spiraecola, but may be distinguished by the characters given in the key to polyphagous aphids, and also by the presence of modified, peg-like hairs on the hind tibiae. Most commonly on Euphorbiaceae, with a preference for woody species (Glochidion, Breynia, Phyllanthus), but plants in other families may be colonised, e.g. Compositae/Asteraceeae (Chromolaena, Clibadium), Convolvulaceae (Convolvulus), Dipsacaceae (Dipsacus), Malvaceae (Hibiscus), Myrtaceae (Eugenia) and Rosaceae (Pyrus, Eriobotrya). Throughout east and south-east Asia (westward as far as Pakistan; Naumann-Etienne & Remaudière 1995), in Australia (Eastop 1966, as A. hardyi). It is also now in Hawaii (Foottit et al. 2012) and New Caledonia (Mille et al. 2020), and (since 2011) in Florida, USA, where large damaging colonies are reported on Eriobotrya japonica (Skvarla et al. 2017). A record from Tunisia (Ben Halima-Karmel 2012) needs further confirmation. Calilung (1976) gave an account of this species in the Philippines. Pseudotoxoptera pyrisucta Qiao & Zhang, described from Pyrus pyrifolia in China (G. Zhang et al. 1999d), is likely to be a synonym. 2n=8.

Aphis evonymi Fabricius Apterae are various shades of chocolate brown, often with transverse bars of wax dust on anterior abdominal tergites (see influentialpoints.com/Gallery); BL 1.7-2.9 mm. Alatae have secondary rhinaria distributed ANT III 18-28, IV 0-7, V 0. Monoecious and holocyclic in Europe on Euonymus europaeus, curling the leaves. It is also recorded from east Siberia, on E. maximowiczianus (Pashchenko 1997). Males, and usually the gynoparae also, are apterous. Hybridisation with A. fabae occurs readily in the laboratory, and may occur in the field, so subspecies status might be more appropriate for this aphid (Müller & Steiner 1986). Heie (1986) provided a general account, and discussed the confused application of this name (as A. euonymi; note that correct spelling is evonymi) in the literature. 2n=8.

Aphis eupatorii Passerini Apterae are dark green, or whitish yellow in midsummer dwarfs; BL 0.9-2.0 mm. Alatae have secondary rhinaria distributed III 4-10, IV 0-4. In ant-attended colonies on young growth and undersides of leaves of Eupatorium cannabinum in Italy. Monoecious holocyclic, fundatrices having been found in early June (Barbagallo & Stroyan 1982). A member of the frangulae group, close to frangulae s. str. (q.v.) which, according to Börner (1952), has Eupatorium cannabinum as a secondary host in Germany. No reliable morphological discriminants are available.

Aphis euphorbiae Kaltenbach Apterae are blackish brown, shiny or wax-dusted, with tibiae pale brown except at apices, and siphunculi and cauda black (see influentialpoints.com/Gallery); BL 1.3-2.4 mm. Alatae have secondary rhinaria distributed III 7-18, IV 0-5. On upper parts of stems of Euphorbia spp., especially (in Europe) E. cyparissia. Sexual morphs have not been described, and it seems to be mainly or entirely anholocyclic. In Europe (but rare in north, and apparently not in Iberian peninsula – see García Prieto et al. 2004); in western Siberia, south-west and central Asia, and introduced to USA (California, but an early record from Nebraska is not this species; see Foottit et al. 2006). A member of a taxonomically difficult group of closely-related species on Euphorbia. Records of A. euphorbiae from Africa and Australia, as well as many of those from southern Europe and the Middle East, seem to be referable to A. tirucallis.

Aphis euphorbicola Rezwani & Lampel Apterae are dark brown or greyish brown, lightly wax-dusted; BL 1.0-1.7 mm. Alatae have secondary rhinaria distributed III 6-12, IV 1-2. On undersides of leaves and shoots of Euphorbia denticulata in Iran.

Aphis exploratus Rusanova Apterae are greenish yellow or dark orange, siphunculi yellowish with darker apices, cauda yellow; BL 1.5-1.9 mm. Alatae have secondary rhinaria distributed III 6-8, IV 0-3. On leaves, flowers and fruits of Paliurus in Azerbaijan. Possibly this was A. nasturtii.

Aphis exsors Rusanova Apterae are brown or green, with dark siphunculi and cauda; BL 1.6-2.3 mm. Alatae have secondary rhinaria distributed III 9, IV 3-4, V 2. Described as occurring on various Compositae/Asteraceae (e.g. Conyza canadensis, Dahlia, Sonchus) in Azerbaijan. This species could not be included in keys on the basis of the published description.

Aphis fabae Scopoli Fig. 4 Plate 6e Apterae are dull black, sometimes with white wax markings, usually ant-attended (see influentialpoints.com/Gallery); BL 1.5-3.1 mm. Alatae have secondary rhinaria distributed III (3-)7-33, IV 0-14, V 0-3. Curling leaves of Euonymus europaeus in spring (also Philadelphus coronarius and/or Viburnum opulus, depending on subspecies – see below), and migrating to a wide range of secondary hosts, including young growth of some trees, and many crops. In Europe there seems to be a complex of sibling species or partially interfertile subspecies, of which four are known to overwinter on Euonymus; they are A. fabae sensu stricto (which in host transfers will colonise Vicia faba), A. fabae cirsiiacanthoidis Scopoli (which will transfer to Cirsium arvense), and two that are dealt with here as separate taxa; A. solanella (which will transfer to Solanum nigrum) and A. evonymi, which stays through the summer on Euonymus. A. fabae sensu stricto, A. fabae cirsiiacanthoides and A. solanella will also sometimes use Viburnum opulus as their primary host. Another closely related form, A. fabae mordwilkoi Börner & Janich, has a sexual phase on Viburnum opulus, or sometimes Philadelphus coronarius, but not on Euonymus, and migrates particularly to Arctium spp. and Tropaeolum majus. This subspecies also occurs in Canada, where it was studied (as barberae Robinson) by Robinson (1980) and Barber & Robinson (1980), and has a sexual phase on Viburnum trilobum. Oviparae on primary hosts in autumn are small, with strongly swollen hind tibiae, and males are alate. Anholocyclic populations of aphids of the A. fabae group (especially its short-haired close relative A. solanella) occur on secondary hosts in southern Europe, south-west Asia, Africa, Indian subcontinent, Korea (Kim et al. 2006b), North and South America, Hawaii and Auckland Isles. Accounts of this group in Europe were given by Stroyan (1984), Heie (1986) and Müller (1988), and Blackman & Eastop (2007) discussed the taxonomic problems. Hybridisation and reproductive isolation between members of the complex have been studied by Thieme & Dixon (1996), Raymond et al. (2001) and Tosh et al. (2004), and Görür et al. (2005) studied the role of phenotypic plasticity in host race formation and speciation. Béji et al. (2015) studied mitochondrial DNA (COI) variation in A. fabae-group populations in Tunisia. Sandrock et al. (2011) conducted microsatellite DNA analyses of A. fabae s. str. collected at different latitudes in Europe and demonstrated a relationship between climate, population structure and life cycle. See also Blackman & Eastop (2000, 2006). 2n=8 (all subspecies).

Aphis farinosa Gmelin Apterae 1.6-2.5 mm, dull pale to mid green with cauda distinctly darker than the long pale siphunculi, in dense colonies on young shoots of Salix spp. in spring and early summer, attended by ants (see influentialpoints.com/Gallery). Alatae have secondary rhinaria distributed III 5-11, IV 0-2. Monoecious, with green oviparae and orange males appearing from July onwards, although occasionally populations of viviparae persist until August or September (Stroyan 1984, Heie 1986). It occurs throughout northern temperate parts of the world, and in Argentina. A nominal subspecies, A. farinosa yanagicola Matsumura, occurs commonly on Salix spp. (daphoides, rorida) in Japan, and is also recorded from China (Qiao & Jiang 2005); it is distinguished by having long, fine hairs on the third antennal segment (about twice the middle diameter of the segment), and it produces sexuales in the autumn (Takahashi 1966). Populations on Salix gracilistyla in Siberia have short antennal hairs like European farinosa, but a longer last rostral segment; R IV+V 1.27-1.45 × HT II, as compared with 1.07-1.25 × HT II (Holman 1987). 2n=6.

Aphis feminea Hottes Apterae red with black head, pro- and mesothorax, antennae, legs, siphunculi and cauda, lightly dusted with white wax. It forms dense colonies on young twigs and small branches of Prunus serotina in north-eastern USA (Hottes & Frison 1931).

Aphis (Zyxaphis) filifoliae (Gillette & Palmer) (Fig.12s) Apterae are brownish amber to dark brown or blackish, with lighter brown markings and a reticulate pattern of white wax; BL c.1.1 mm. On leaves and stems of Artemisia and Seriphidium spp., and also reported to occur on Chrysothamnus nauseosus, in western USA. Monoecious holocyclic; oviparae and apterous males were collected in early October (see Palmer 1952).

Aphis filipendulae Matsumura Apterae are dark greenish brown, yellow-green or green, powdered with wax; BL c. 1.0-1.3 mm. On flower stems and lower leaf surfaces of Filipendula spp., causing wrinkling of leaves. In Japan (Hokkaido), east Siberia (Takahashi 1966; Pashchenko 1988a) and Kazakhstan (Kadyrbekov 2012a).

Aphis (Bursaphis) fluvialis Martin Apterae are bluish green with a slight wax bloom: BL 1.0-1.7 mm. On shoot apices and flower stalks of Epilobium hirsutum in Africa (west Sudan), attended by ants. Probably anholocyclic. Martin (1982) discussed the possible origins of the aphid and its host from Europe. 2n=9.

Aphis foeniculivora Zhang Apterae are green, BL c. 1.4 mm. On Foeniculum vulgare in China. Apart from a longer antennal terminal process, this species could not be distinguished by its published description from A. gossypii.

Aphis folsomii Davis Apterae are reddish brown (see influentialpoints.com/Gallery); BL 1.6-1.8 mm. On Parthenocissus spp., forming dense clusters along midribs of leaflets and petioles, attended by ants (Williams 1911, as Aphis parthenocissi). Widely distributed in USA east of Rocky Mountains.

Aphis forbesi Weed Apterae are dark bluish green, sometimes with yellowish mottling (see influentialpoints.com/Gallery); BL 1.0-1.9 mm. Immatures are yellowish green. Alatae have black dorsal abdominal cross-bands and only 1-4 secondary rhinaria on ANT III. In ant-attended colonies on shoots and at bases of leaf petioles of Fragaria spp., and also sometimes on the roots. Native to North America, and introduced into Europe about 1928; also in Kazakhstan, west Siberia, Japan, Australia (Brumley 2020), Central and South America. Monoecious holocyclic with apterous males; Marcovitch (1925) discovered the photoperiodic induction of sexual morphs in aphids while working with this species. 2n=8.

Aphis frangulae Kaltenbach Apterae are various shades of yellow, green, blue-green, brown or greenish black, sometimes mottled (see influentialpoints.com/Gallery); BL 0.9-2.4 mm. Alatae have secondary rhinaria distributed III 3-16, IV 0-8, V 0-3. A complex of species/subspecies utilising a wide range of herbaceous plants, some of which have a sexual phase in Europe on Rhamnus frangula, curling its leaves in spring, and the name is also applied to aphids utilising other Rhamnus spp. (davurica, ussuriensis) as primary hosts in east Asia (Pashchenko 1997). [A. rhamnicola (q.v.) described from R davurica in Korea is also not clearly distinct.] Parthenogenetic overwintering of members of the group also occurs on the evergreen R. alaternus in the Mediterranean region (Swirski & Amitai 2001), and an aphid on Sageretia in Pakistan has also been identified as A. frangulae (Naumann-Etienne & Remaudière 1995). Stroyan (1984) reviewed the group and discussed the taxonomic problems. In Europe, A. frangulae sensu stricto migrates from R. frangula to Epilobium angustifolium, also occurring frequently on Capsella bursa-pastoris and Lysimachia vulgaris, and A. frangulae beccabungae Koch migrates from the same primary host but preferentially colonises Veronica beccabunga, also occurring on Solanum tuberosum and various Lamiaceae. A. f. beccabungae has particularly been reported to occur on various labiates (Dracocephalum, Scutellaria, Ziziphora) in the Kazakhstan part of West Tien-Shan (Kadyrbekov 2005c). However, summer populations on labiates are practically indistinguishable from those of other species/subspecies in the complex (e.g. balloticola, capsellae, lamiorum, nepetae) that are monoecious holocyclic on the same plant genera, and populations outside Europe (e.g. in south-west and central Asia), can usually only be identified as frangulae (sensu latore). A third European form only known from Germany, A. frangulae testacea Thomas, may be monoecious on R. frangula. Populations of these three subspecies possibly show colour differences in life (see Heie 1986). There are no fully reliable morphological discriminants, perhaps because natural hybridisation may tend to confuse secondary host associations and life cycle categories, and there are also geographical differences, which may change with time. For example, Stroyan (1984, p.124) noted a frangulae-like aphid that had recently been collected in England on Hypericum; this seems to be the same as an Aphis sp. collected in Syria and Lebanon prior to 1954 (BMNH collection, leg. A.L. Talhouk). Alatae in populations on Nepeta from Spain (leg. H.L.G. Stroyan) and USA (Idaho, Utah; leg. G.F. Knowlton) all have a large number of secondary rhinaria distributed over ANT III, IV and V, and may be a distinct species. The cosmopolitan pest A. gossypii (q.v.) is closely related, and introduces a further complication to the taxonomic treatment of the group, as it can be conveniently treated as a subspecies of frangulae in Europe (Stroyan 1984, Heie 1986). However, this becomes untenable when world-wide populations are taken into account (see below and Blackman & Eastop 2017), and recent work comparing mitochondrial DNA sequences (Cocuzza et al. 2009, Carletto et al. 2009a) has shown clear differences between the frangulae and gossypii groups. Cocuzza & Cavalieri (2014) have gone on to compare a mitochondrial DNA (COI) sequence in numerous samples of frangulae-group aphids collected from Lamiaceae. They found generally very low levels of variation, and in particular suggested that three nominal species (lamiorum, stachydis, symphyti) should be regarded as synonyms of frangulae, but this does not take into account the different life cycles of these species. Further molecular work is needed using different genetic markers. 2n=8.

Aphis franzi Holman Colour of apterae in life unknown; BL 1.1-1.7 mm. Alatae are undescribed. On Seseli elatum ssp. austriacum in Austria, and reported also to occur in Kazakhstan on several other Seseli spp., with records also from Peucedanum morisonii and Silaum alpestre (Kadyrbekov 2009a, 2012a, 2017a); also in west Siberia (Kadyrbekov 2014g – but there on Linaria, unlikely to be a true host).

Aphis fraserae Gillette & Palmer Apterae are yellowish green, mottled with darker green, often blackish, with dark siphunculi and cauda; BL c.2 mm. Apterae have 0-15 secondary rhinaria on ANT III, and alatae have 18-28 on III and 1-7 on IV. On flower stalks of Frasera speciosa in western USA (Palmer 1952). Distinguished from polyphagous Aphis spp. by the long R IV+V and short ANT PT.

Aphis fuentesi Nieto Nafría & Ortego Apterae are matt dark green to matt black; BL 1.0-1.7 mm. Alatae have secondary rhinaria distributed III 4-13, IV 0-3. In compact groups on stems of several Baccharis spp. in Argentina (Neuquén, Chubut, Rio Negro) and Chile (Maule). Oviparae and males were collected in April (Nieto Nafría et al. 2019c).

Aphis fukii Shinji Apterae are dark grey-brown or black, with black siphunculi and cauda; BL 2.1-2.5 mm. Alatae have secondary rhinaria distributed III 20-32, IV 5-11, V 1-7. Curling and twisting leaves of Petasites spp. in east Asia (Japan, Korea, Sakhalin, Taiwan). Also recorded (from Japan) on Senecio kaempferi (= Farfugium japonicum). 2n=8.

Aphis fumanae Remaudière & Leclant Apterae are very dark green to black, shining; BL 0.9-1.3 mm. Alatae have 3-8 secondary rhinaria on ANT III. In small colonies on shoots and flower peduncles of Fumana spp. in southern France. Monoecious holocyclic with apterous males (original description). Colonies were attacked by the aphidiid parasitod Lysiphlebus fabarum Marsh.

Aphis funitecta (Börner) Apterae are densely wax-covered, BL 1.6-2.4 mm. On Bupleurum spp. in Germany, Poland, Kazakhstan (Kadyrbekov 2017a) and Russia (Altai Krai). Monoecious holocyclic with apterous males, and oviparae having unswollen hind tibiae (Nieto Nafría et al. 1986).

Aphis galiae Ivanoskaya Apterae are broadly oval, greenish brown with black antennae, legs, siphunculi and cauda; BL c. 2.3-2.4 mm. Described from Populus in western Siberia, and also recorded from P. nigra in Kazakhstan (Kadyrbekov 2017a), but there are also records from Betula nana (in west Siberia), and from Betula sp. in the Kuril Islands (Ivanoskaya 1977, Krivoluckaya & Ivanoskaya 1966), indicating a host range that warrants further investigation (see also Stekolshchikov & Khureva 2020). The life cycle is unknown.

Aphis galiiscabri Schrank (Fig.29a) Apterae are greenish black, heavily wax-powdered, BL 1.4-2.2 mm. Alatae have secondary rhinaria distributed III 5-14, IV 0-3, V 0-1. On upper parts of Galium spp., causing shortening of stems so that leaves and flowers of young shoots become close-set. It is also reported from some species of Asperula and Rubia. In Europe, west Siberia, Turkey, Iran and Central Asia (Kazakhstan); Nieto Nafría et al. (2005a) reported it also in Mongolia and Canada. Populations in southern Italy show some morphological differences (Stroyan 1984). Fundatrices have been collected in May (BMNH collection), sexual morphs are apparently unknown.

Aphis gallowayi Robinson Apterae are dark brown to black, BL 1.6-2.2 mm. On Astragalus pectinatus in Alberta, Canada. Very similar to A. astragalina, and which has been found on the same plant.

Aphis gaultheriae López Ciruelos, Brown & Nieto Nafría Colour in life unknown, BL of aptera 1.8-2.1 mm. On Gaultheria mucronata in southern Chile (Chiloé). Other morphs and life cycle are unknown (López Ciruelos et al. 2018).

Aphis genistae Scopoli Apterae are black, coated with wax meal (see influentialpoints.com/Gallery); BL 1.4-2.6 mm. Alatae have 4-8 secondary rhinaria on ANT III. On Genista spp. in Europe, eastward to Ukraine and Turkey, and introduced to North America (Foottit et al. 2006). Sometimes with ants. Monoecious holocyclic, with oviparae and alate males in September 2n=8.

Aphis gentianae (Börner) Apterae are dark green, BL 1.2-1.6 mm, at stem bases in ant shelters and on subterranean parts of Gentianaceae (Blackstonia, Centaurium, Gentiana) in central, southern and eastern Europe, south west Siberia (Novgorodova & Stekolshchikov 2013), and recently found in Wales (Baker 2009a). A redescription of apterae and alatae was provided by Barbagallo & Massimino Cocuzza (2014). Sexual morphs were found in Poland in September (Szelegiewicz 1975a). Infestation of Gentiana cruciata stems by A. gentianae significantly deterred oviposition by the butterfly Maculinea alcon (Árnyas et al. 2009).

Aphis gerardiae (Thomas) Apterae are blackish (see influentialpoints.com/Gallery), BL 1.9-2.9 mm. On Agalinis and Aureolaria spp. in eastern USA (Florida, Illinois, Indiana, Kansas, Kentucky, New York, Pennsylvania). Life cycle seems not to have been studied.

Aphis gerardianae Mordvilko Apterae are greyish black to black, wax-dusted; BL 1.0-1.7 mm. Alatae have 8-11 secondary rhinaria on ANT III. On stems and flower stalks of Euphorbia sequieriana (incl. gerardiana) in central and eastern Europe (Jörg & Lampel 1988), and eastward to Central Asia, where Kadyrbekov (2005b) recorded it from several other Euphorbia spp. in Kazakhstan. In Hungary it is attended by the ant Plagiolepis pygmaea (Szelegiewicz 1966a).

Aphis glareosae Bozhko Apterae are dark brown, wax-dusted; BL c. 2 mm. Alatae have secondary rhinaria distributed III 10-12, IV 1-3. On Euphorbia glareosa, and also on E. pannonica (BMNH collection, leg. J. Holman), in eastern Europe (Hungary, Czech Republic, Bulgaria, Rumania, Ukraine, Turkey). Oviparae have been collected on E. glareosa in October (BMNH collection, leg. J. Holman).

Aphis globosa Pashchenko Apterae are pale green with some dark abdominal markings, BL c. 1.8 mm. On underground stems of Artemisia gmelinii in east Siberia. Biology is unknown.

Aphis glycines Matsumura Apterae are pale yellow or greenish yellow with pale head and basal antennal segments, and siphunculi dark except at their bases (see influentialpoints.com/Gallery); BL 1.2-1.7 mm. Alatae have 4-9 secondary rhinaria on ANT III. On stems and undersides of leaves of various Leguminosae/Fabaceae, particularly Glycine spp., and a major pest of soybean (Blackman & Eastop 2000, p.229); also recorded from Amphicarpaea, Desmodium, Phaseolus and Pueraria. In east and south-east Asia, and introduced to USA in about the year 2000 (Ragsdale et al. 2004), and then to Canada (Hunt et al. 2003). It has also been present in eastern Australia since at least 2000 (Fletcher & Desborough 2000), and is also in New Caledonia (Mille et al. 2020). Heteroecious holocyclic in China and Japan with Rhamnus spp. (davurica, japonica) as primary hosts (Wang et al. 1962, S. Takahashi et al. 1993, Kim et al. 2010b). In North America Rhamnus cathartica, R. alnifolia, R. frangula and R. lanceolata are accepted as primary hosts (Voegtlin et al. 2004, 2005; Yoo et al. 2005; Lagos-Kutz & Hartman 2021). Its introduction into USA stimulated the setting-up of a soybean aphid suction trap network (reviewed by Lagos-Kutz et al. 2020), and led to intensive studies of all aspects of its biology; see Tilmon et al. (2011) for a general review, and Orantes et al. (2012), Jun et al. (2013) and Giordano et al. (2020) for subsequent studies of genetic diversity in US and Asian populations. Naaum et al. (2014) have developed a specific PCR assay with potential for field identification. H. Li et al. (2018) developed methods of instar identification. The complete genome sequence has been published (Wenger et al. 2020), as well as that of the mitochondrial genome (Song et al. 2019). 2n=8 (Mandrioli et al. 2019b).

Aphis gnetuma Qiao Apterae are reddish brown to dark brown, covered with white wax powder; BL 1.7-1.9 mm. On undersides of leaves of Gnetum parvifolium in Guangxi, China (Y. Xu et al. 2021). Other morphs and life cycle are unknown.

Aphis gossypii Glover Plate 7c Apterae vary in colour from dark blackish green or green mottled with dark green (larger specimens in favourable conditions; see influentialpoints.com/Gallery) to very pale whitish yellow (small specimens in crowded colonies or hot conditions); BL 0.9-1.8 mm. Alatae have secondary rhinaria distributed III (3-)6-12(-15), IV 0(-2). On a very wide range of host plants, its polyphagy being particularly evident during the dry season in hot countries. A major pest of cotton and cucurbits, and in glasshouses in cold temperate regions (Blackman & Eastop 2000, p.230). Distributed almost worldwide, and particularly abundant and well-distributed in the tropics, including many Pacific islands. Anholocyclic in warm climates, and mainly so in Europe. Molecular studies indicated that occasional (“cryptic”) sex may be occurring in southern France (Thomas et al. 2009), and subsequent work (Thomas et al. 2012) found evidence of recent sexual reproduction in spring migrant alatae alighting on melon plants in three French regions, although the primary hosts of these alatae were unknown and some confusion with members of the A. frangulae group seems possible.

Host alternation and a sexual phase occur more regularly in parts of east Asia (e.g. Japan, Takada 1988; China, Zhang & Zhong 1982a) and also in North America (Kring 1959), with several unrelated plants utilised as primary hosts (incl. Catalpa bignonioides, Hibiscus syriacus, Celastrus orbiculatus, Rhamnus spp. and Punica granatum). Apterae of spring populations on primary hosts are usually greenish. Males are always alate. However, Komazaki & Toda (2008) obtained wingless gynoparae from some holocyclic clones in Japan, and Zhang & Zhong (1982a) noted monoecious holocyclic populations on cotton and Hibiscus in China. Likewise there are aphids identified as A. gossypii that overwinter as eggs on twigs of Lycium barbatum in China (Yan et al. 2017).

Molecular studies indicate the occurrence of host-associated populations or races, some of which may have a world-wide distribution (Charaabi et al. 2008, Carletto et al. 2009b, L.Wang et al. 2016, Luo et al. 2016, R.H. Miller et al. 2019); but see also Lokeshwari et al. (2015). Genetic diversity of A,. gossypii populations in a cotton agroecosystem in northern Cameroon has been studied by Brévault et al. (2008), and Nam et al. (2019) investigated the genetic structure of populations on peppers in Korea using microsatellites. Najar-Rodríguez et al. (2009) demonstrated clear differences in both behavioural and physiological mechanisms of host detection and acceptance of Malvaceae and Cucurbitaceae by alatae of clones originating respectively from cotton and pumpkin in Australia. Y.-C. Zhang et al. (2017) found differences in gene expression profiles between cotton-specialised and cucurbit-specialised lineages of A. gossypii in China.

Some populations regarded as A. gossypii certainly seem to be functioning as distinct species; for example, populations with a sexual phase on Rubia cordifolia in Japan that seem to be isolated from those on other primary hosts (Inaizumi 1981), populations reproducing sexually on Hibiscus in Korea that would not colonise vegetable crops including cucumber (Y.-H. Kim et al. 2008), and a form feeding on Compositae/ Asteraceae in Europe and north Africa that can produce sexual morphs under certain conditions (A. parvus Theobald? – see Margaritopoulos et al. 2006). However, F. Zhang & Liu (2012) demonstrated interbreeding between populations with different life cycles and host associations in China.

Populations producing eggs on roots of various plants (Ixeris, Lactuca, Marrubium, Viola) in China identified as A. gossypii (e.g. Chu 1949) may be other closely-related monoecious holocyclic species, such as A. violaeradicis (q.v.). Taxonomic implications of the genetics, biology and distribution of A. gossypii were reviewed by Blackman & Eastop (2007, 2017). A. gossypii has been often been treated as a subspecies of A. frangulae in the European aphid taxonomic literature, but recent molecular work has shown that these two taxa are phylogenetically distinct (see under A. frangulae). Favret & Miller (2011) have designated a neotype for A. gossypii from material freshly collected in South Carolina, and published its DNA barcode. The complete mitochondrial genome has been published (S. Zhang et al. 2016), and a high quality draft of the whole nuclear genome is now available (Quan et al. 2019) . 2n=8.

Aphis grandis Juchnevitch Apterae have dark brown head and prothorax, rest of thorax and abdomen yellow with broad brownish cross-bands and marginal spots, appendages mainly dark brown; BL 2.8-3.1 mm. Alatae have 23-27 secondary rhinaria on ANT III. In ant-attended colonies on green shoots of Spiraea spp. in Kazakhstan. Apparently migrating in June to an unknown secondary host.

Aphis grata Pashchenko Apterae are brown, pulverulent; BL 1.3-1.9 mm. Alatae have 8-9 secondary rhinaria on ANT III. In small dense colonies on undersides of leaves and flower stems of Spiraea spp. in east Siberia.

Aphis gratiolae Bozhko Apterae are yellowish with greenish mottling; BL c.1.2 mm. Alatae have 4-5 secondary rhinaria on ANT III. On Gratiola officinalis in Ukraine.

Aphis gregalis Knowlton Apterae are bluish green to yellowish green with darker head and pronotum, and variably-developed dorsal abdominal dark markings; BL 1.2-1.9 mm. Secondary rhinaria of alata ditributed III 2-7, IV 1-2. On Chrysothamnus spp. in western USA (western slopes of Rocky Mountains), often in large colonies on shoot tips, leaves and flower-stems, causing stunting and distortion of growth. Oviparae and apterous males in October (original description). Closely related to A. ornata, and without any clear distinguishing characters in slide-mounted specimens.

Aphis grosmannae Börner Colour of apterae in life unknown, probably dark green; BL 1.8-2.5 mm. Alatae have 6-12 secondary rhinaria on ANT III. On stems and umbels of Peucedanum spp. in central and eastern Europe (Germany, Poland, Czech Republic), Kazakhstan and west Siberia. There are also records from other Umbelliferae/Apiaceae (Angelica, Cicuta, Heracleum, Libanotis, Seseli), and from Parasenecio (= Cacalia) hastatus (Ivanoskaja 1977; probably not a normal host). Szelegiewicz (1964b) provided a redescription.

Aphis (Bursaphis) grossulariae Kaltenbach Apterae are dull greyish green to dark green or yellow-green, with pale siphunculi (see influentialpoints.com/Gallery); BL 1.2-2.1 mm. Secondary rhinaria of alata ditributed III 7-19, IV 3-11, V 0-5. Host-alternating between Ribes spp., mainly R. grossularia, and Epilobium spp., especially E. montanum. Certain other Onagraceae (Clarkia, Fuchsia) may also be utilised as secondary hosts. Colonies on gooseberry cause considerable leaf-curl. On Epilobium the aphids feed on the flowerheads, and in Europe may form mixed colonies with A. epilobii. In Europe, Central Asia, Pakistan (Naumann-Etienne & Remaudière 1995), across Siberia and in China (Jiang et al. 2011b). A record from USA (Oregon) is unconfirmed (Foottit et al. 2006). Rakauskas (1999, 2000) and Turčinavičienė (2000) reported on experimental hybridisation with the closely-related species A. schneideri and A. triglochinis, and Rakauskas (2003b) reported morphological evidence of natural hybridisation in Europe with A. schneideri. Turčinavičienė et al. (2006) used molecular techniques to study phylogenetic relationships with other palaearctic Bursaphis. 2n=8.

Aphis gypsophilae Holman Apterae are shining dark green; BL 1.6-2.1 mm. Secondary rhinaria of alata ditributed III 10-14, IV 1-3, V 0-3. On Gypsophila dahurica in Mongolia.

Aphis haroi Nieto Nafría Apterae are matt black; BL 2.1-3.1 mm. Apterae have secondary rhinaria distributed III 0-14, IV 2-18, V 0-5; alatae have them distributed III 36-39, IV 18-19, V 4-5 (Nieto Nafría et al. 2005a). On Senecio pyrenaicus (= tournefortii) in Spain. Monoecious holocyclic with apterous males (original description).

Aphis hasanica Pashchenko Apterae are black, with waxy coating; BL c.1.9 mm. Alatae are undescribed. On Thalictrum simplex in east Siberia. Ant-attended.

Aphis healyi Cottier Apterae are dark green to black, paler laterally, often with greenish tinge, siphunculi and cauda black; BL c. 2.0-2.2 mm. Alatae have 0(-1) secondary rhinaria on ANT III. On terminal parts, especially new growth, flowers and seed-pods, of Carmichaelia spp. (odorata, subulata) in New Zealand. 2n=8*. Oviparae and males occur in March-April (Teulon et al. 2013). Podmore et al. (2019) reported COI sequence variation between populations.

Aphis hederae Kaltenbach Apterae are dark brown or dull black (see influentialpoints.com/Gallery); BL 1.4-2.5 mm. Secondary rhinaria of alatae distributed III 8-28, IV 0-13, V 0-4. On young shoots and foliage of Hedera helix, sometimes on other Araliaceae (Aralia, Schefflera), and also on Cuscuta (Convolvulaceae). Widespread in Europe, Tunisia, west and south-west Asia, Korea (Kim et al. 2006b), southern Africa, New Zealand, and also common and widespread in North America and temperate South America. Monoecious holocyclic; males may be apterous or alate [Possibly a species complex; two other closely-related taxa have been described, A. boerneri Franssen and A. pseudohederae Theobald, but these are treated here as variant populations of A. hederae.] 2n=8.

Aphis hederiphaga Takahashi Apterae are dark green or brown; BL 2.0-2.2 mm. Secondary rhinaria of alatae distributed III 6-8, IV 0-2. In large colonies on new growth and leaves of Hedera spp. in Japan, and also in Korea (Lee et al. 2002c).

Aphis hedysari Holman Apterae are blackish brown with basal parts of antenna and most of tibiae whitish; BL 2.4-2.7 mm. Secondary rhinaria of alatae distributed III 6-11, IV 0-4. On Hedysarum austrosibiricum in Russia (Transbaikalia), living in dense colonies on terminal plants of host.

Aphis heiei Holman Apterae are bluish green to dark green; BL 1.3-1.8 mm. On stems and leaves of Hieracium umbellatum. So far only known from sandy areas on west coast of Jutland, Denmark (Holman 1998).

Aphis helianthemi Ferrari Apterae are bright pale green, or dark bluish green (“ssp. thermophila Börner”), or blackish (“ssp. obscura Bozhko”), the more pigmented specimens having dusky/dark head, antennae, femora, siphunculi and cauda, and dark to black distal sections of tibiae; BL 0.9-1.9 mm. Alatae have 2-6 secondary rhinaria on distal part of ANT III. On upper parts of stems and flowers of Helianthemum spp., attended by ants. Lampel & Meier (2007) also recorded it from Halimium spp. Widely distributed in Europe. A. helianthemi ssp. thermophila, described from H. canum, has been treated as a separate species by several authors, but there are no good distinguishing characters for slide-mounted specimens, and the colour variants are not consistently associated with particular species of host plant. The key couplet provided by Remaudière & Leclant (1972) will not work for some British populations of helianthemi (BMNH collection), nor for some Spanish thermophila (Nieto Nafría 1976). We are therefore treating all these populations as one variable species.

Aphis heraclicola Shinji Apterae are pale green with dark siphunculi and cauda; BL c. 1.3-1.4 mm. Alatae have secondary rhinaria distributed III 10-14, IV 4-8. On Angelica polymorpha and Heracleum lanatum in Japan. This species could not be included in the keys to apterae on Angelica and Heracleum on the basis of the information available, although alatae should be recognisable by the distribution of rhinaria on ANT III-IV.

Aphis (Zyxaphis) hermistonii Wilson (Fig.12u) Apterae are dark grey-green with white powdery markings and thick black siphunculi; BL 0.9-1.4 mm. On leaves and young shoots of Artemisia filifolia and Seriphidium tridentatum in western USA. Records from Chrysothamnus may be misidentifications. There is some confusion in the literature with A. canae and A. oregonensis, but the three species seem separable by the key characters given.

Aphis herniariae Mamontova-Solukha Apterae are brownish with dark siphunculi and cauda; BL c. 1.1 mm. Alatae have not been described. On Herniaria besseri (= incana) in Ukraine.

Aphis hieracii Schrank Apterae are bluish to dark green, with dark head, siphunculi and cauda; BL 0.8-2.8 mm. Alatae have secondary rhinaria distributed III 5-10, IV 0-3. In ant-attended colonies on stems and leaves of Hieracium spp. in northern, eastern and central Europe, west and east Siberia (Pashchenko 1992) and Kazakhstan (Kadyrbekov 2012a). North American records seem to be unsubstantiated (Footit et al. 2006). Monoecious holocyclic with apterous males. Dwarf apterae occur in midsummer (Holman 1998).

Aphis hillerislambersi Nieto Nafría & Mier Durante Apterae are dark olive-green to black covered in greyish wax; BL 1.3-2.0 mm. Alatae have secondary rhinaria distributed III 5-15, IV 0-3, V 0(-2). On Euphorbia spp., recorded from Spain, France, Italy, Croatia (BMNH collection, leg. H.L.G. Stroyan), Balearics, Canary Isles, Algeria (Peréz Hidalgo et al. 2012b), Madeira (BMNH collection and Ilharco, 1974, as A. paralios Hille Ris Lambers, nomen nudum – although Aguiar et al. (2013) noted some differences between Spanish and Madeiran specimens), Turkey (BMNH collection, leg. B. Kovanci) and Saudi Arabia (Aldryhim & Khalil 1996). The Spanish populations were found to be anholocyclic (original description).

Aphis hiltoni Essig Apterae are pale green with dark head and dorsal abdominal markings, partially covered with white powdery wax; BL c.1.3 mm. Alatae have 4-6 large secondary rhinaria on ANT III. Described from dense colonies on apical twigs of Artemisia californica in California, USA, but there are apparently no subsequent records.

Aphis hispanica Hille Ris Lambers Colour of apterae in life unknown, probably dark and wax-dusted; BL 1.0-1.3 mm. Alatae have 6-7 secondary rhinaria on ANT III (Nieto Nafría et al. 2005a). On Euphorbia sp. in Spain (original description), and Tuatay & Remaudière (1965) decribed oviparae and alate males from an unidentified Euphorbia sp. in October in Turkey. It has also been found in Iran (Rezwani 2010) and Corsica (Lampel 2001).

Aphis holmani Kanturski & Bezdĕk Appearance of apterae in life is uknown; BL 1.5-2.0 mm. Alatae have secondary rhinaria distributed III 4-6, IV 0-1. On Knautia drymeia in central and south-east Europe (Czech Republic, Bulgaria, Romania). Sexual morphs occur in September (Kanturski & Bezdĕk 2019). [This is the species referred to as A. knautiae in Holman & Pintera (1981), where it is a nomen nudum. A holmani Kanturski & Bezdĕk is a secondary homonym of A. holmani Stekolshchikov, and will require replacement.]

Aphis holmani Stekolshchikov Apterae are brown, lightly wax-powdered; BL 1.6-2.3 mm. On root collar of Epilobium parviflorum in southern Russia (Astrakhan province), sheltered and attended by ants (Stekolshchikov 2005). Life cycle is unknown, but probably monoecious holocyclic.

Aphis (Bursaphis) holoenotherae Rakauskas Apterae are dull green, yellow-green or yellow; BL 1.3-2.0 mm. Alatae have secondary rhinaria distributed III 8-15, IV 3-7, V 0-2. In northern and eastern Europe. Populations in northern Europe are monoecious holocyclic on Oenothera spp., with alate males (Rakauskas 2007, 2008). Anholocyclic overwintering may also occur. This species is very similar to the North American A. oenotherae (q.v.), and was originally regarded as an introduced population of that species. Oenothera is a nearctic genus, although with a long history of cultivation in Europe, so A. holoenotherae must be either a recently evolved species or one as yet unrecognised in its country of origin.

Aphis horii Takahashi Apterae are blackish-green, BL 2.1-2.7 mm, clustered on stems and leaf petioles of Sambucus spp. in Japan. Paik (1965) also recorded this aphid (as Sappaphis euscaphis) in Korea from Euscaphis japonica (Staphyleaceae). Closely related to A. sambuci, but differing in the presence of long, fine hairs on the head and antennae. Host alternation to roots of Caryophyllaceae, Polygonaceae, etc., as in A. sambuci is not recorded in the literature, but specimens possibly of this species have been collected in Korea on Rumex coreanus (Lee et al., 2002c). A. horii was originally described from Cirsium dipsacalepsis (Takahashi 1923), but Takahashi (1966) regards this host record as doubtful. 2n=8 (Chen & Zhang 1985b).

Aphis humuli (Tseng & Tao) Apterae are yellowish green with black eyes and blackish brown siphunculi; BL c.1.2 mm. Alatae have 8-14 secondary rhinaria on ANT III. On Humulus japonicus (= scandens?) in China and Taiwan (BMNH collection, leg. C-C. Tao).

Aphis hyperici Monell Apterae are pale reddish to purple, wax-covered; BL 1.0-1.7 mm. Alatae have secondary rhinaria distributed ANT III 6-13, IV 0-5, V 0-4. On young twigs and undersides of leaves of Hypericum spp. in Illinois and Missouri, USA, and recently found in California (G.W.Watson, pers. comm. 2011) and New Mexico (aphidtrek.org). Monoecious holocyclic; Lagos et al. (2012) redescribed apterae and alatae, and also described the fundatrix, ovipara and alate male. Sexual morphs occur in October-November. 2n=8.

Aphis hypericiphaga Pashchenko Apterae are yellowish green, yellow or yellow-orange, with bands of wax; BL c. 2 mm. Alatae have secondary rhinaria distributed III 6-11, IV 0-2. On undersides of leaves of Hypericum spp. in east Siberia, and North Korea (Lee et al. 2002c).

Aphis hypericiradicis Pashchenko Apterae are yellow-green with dark green anterior and posterior parts of body, black siphunculi and green cauda (immatures almost white); BL c. 1.4 mm. In large ant-attended colonies on stem bases and roots of Hypericum spp. in east Siberia, and now also recorded from Kazakhstan (Kadyrbekov 2017a).

Aphis hypochoeridis (Börner) Apterae are bright yellow to pale greenish yellow, with orange spots at bases of the dark siphunculi, and a dusky cauda (see influentialpoints.com/Gallery); BL 0.7-1.6 mm. Alatae have secondary rhinaria distributed III 5-8, IV 0-4. In ant-attended colonies at base of stem, on root collar and on undersides of radical leaves of Hypochaeris spp. It is widely distributed in Europe, and was introduced many years ago to Japan (Sugimoto 2020a, Sasaki 2021). Monoecious holocyclic; fundatrices in April (BMNH collection), sexual morphs have not been observed.

Aphis ichigo Shinji Apterae are bluish green; BL 1.8-2.1 mm. Alatae have secondary rhinaria distributed III 4-7, IV 0-2. On young shoots and leaves of Rubus spp. in Japan, Korea and east Siberia, and also found on Fallopia japonica (= Polygonum cuspidatum), which is perhaps the secondary host (Takahashi 1966). 2n=8.

Aphis ichigocola Shinji Apterae are brownish yellow with brown head and blackish siphunculi; BL 1.5-2.0 mm. Alatae have 4-6 secondary rhinaria on ANT III. On Rubus spp. in Japan, China and Korea, and also recorded from Fragaria spp. in China (Tao 1999) and Korea (Lee et al. 2002c). A new subgenus (Machilaphis) was erected for this species by L. Zhang & G. Zhang (2002b).

Aphis idaei van der Goot Apterae are yellow mottled with green or pale green to yellow, with long thin curved dusky or dark-tipped siphunculi; BL 1.0-2.0 mm. Alatae have 3-6 secondary rhinaria on ANT III, and 0-1 on IV (see also influentialpoints.com/Gallery). In dense ant-attended colonies at shoot tips of Rubus idaeus, causing severe leaf-curl, and later on undersides of leaves. Related cultivars, e.g. loganberry, and R. occidentalis, are also colonised. Throughout Europe, west Siberia, Iran, Central Asia (Kadyrbekov 2017a), and in New Zealand and North America (British Columbia; BMNH collection, leg. R. Stace-Smith). Monoecious holocyclic with apterous males. 2n=8.

Aphis ilicis Kaltenbach Apterae are dark olive-brown, reddish or greyish-brown (see influentialpoints.com/Gallery); BL 1.7-2.9 mm. Alatae have secondary rhinaria distributed III 8-28, IV 0-6, V 0-1. Living in dense colonies on young shoots and undersides of young leaves of Ilex aquifolium. Attacked leaves curl towards their undersides. Immatures often have white pleural wax markings (it is a member of the A. fabae group). Apparently monoecious, although it is unclear how colonies survive through the summer under natural conditions, as mature leaves are not colonised. Sexuales are recorded from July onwards; males are alate. A. ilicis occurs in most of Europe, eastward to Turkey. North American records require confirmation (Foottit et al. 2006). (See also Stroyan 1984, Heie 1986.) 2n=8.

Aphis illinoisensis Shimer Plate 6f Apterae are rather shiny, deep reddish-brown to almost black (see influentialpoints.com/Gallery); BL 1.6-2.1 mm. On Vitaceae (Ampelocissus, Cissus, Parthenocissus, Vitis) in eastern and central USA, central and South America, and introduced into Turkey and the Mediterranean region, where it is currently extending its distribution, and is now reported from Greece, Montenegro, Cyprus, Malta, Israel, Spain, Algeria, Tunisia, Libya, Italy, France (Barjadze & Ben Dov 2011, Havelka et al. 2011, Mifsud & Pérez Hidalgo 2011, Casirhagi 2021, Mouttet & Balmes 2021). It is also now in Saudi Arabia (Hussain et al. 2015a) and Iran (Heidarnia & Derakhshan 2018). Heteroecious holocyclic in USA (Virginia) with a sexual phase on Viburnum prunifolium (Baker 1917c), possibly anholocyclic elsewhere. There is a North American record of it feeding on Agalinis (=Aureolaria) laevigata (Favret et al. 2010). Havelka et al. (2011) discussed the potential role of native parasitoids in Mediterranean countries.

Aphis impatientis Thomas (= Aphis floridanae Tissot; Lagos-Kutz et al. 2018) Apterae are olive, purplish brown or brownish, dusted with white wax; BL 1.1-1.8 mm. Alatae have secondary rhinaria distributed III 7-17, IV 1-10, V 0. Host-alternating from Cornus spp. to Impatiens spp. in eastern USA and Canada. Lagos-Kutz et al. (2018) recorded Erechtites hieracifolius and Lactuca floridana as additional secondary hosts and described sexual morphs obtained by transfer to Cornus racemosa. Records from Turkey (Görür et al. 2009) and Algeria (Laamari et al. 2013) require further confirmation.

Aphis impatiphila Pashchenko Apterae are pale green; BL c. 1.7-1.8 mm. Ant-attended, on roots of Impatiens glandulifera in east Siberia.

Aphis impatiradicis Pashchenko Apterae are pale green; BL c.1.7 mm. Ant-attended, on roots of Impatiens glandulifera in east Siberia.

Aphis incerta Nevsky Apterae are green or yellow-green with black siphunculi and cauda: BL 1.4-1.8 mm. Alatae have secondary rhinaria distributed III 14-16, IV 8-10, V 0. On Medicago sativa in Uzbekistan. Males are alate (original description). This species could not be included in the key to aphids on Medicago on the basis of the published description.

Aphis indigoferae Shinji Apterae are brick-brown, covered in white wax; BL c. 1.2 mm. . Alatae have secondary rhinaria distributed III 8-10, III 1-4. On Indigofera sp. in Japan, and apparently unrecorded since its original description. Takahashi (1966) regarded this species as indeterminate, and it could not be included in the key to aphids on Indigofera on the basis of the published description.

Aphis (Zyxaphis) infrequens Knowlton in Knowlton & Smith Colour of apterae in life is not recorded; BL 1.5-1.75 mm. On Chrysothamnus parryi in Utah, USA. Alatae were collected from Seriphidium tridentatum, but these were possibly vagrants (Palmer 1952). Very similar to A. chrysothamnicola, and it needs to be confirmed as a distinct species.

Aphis ingeborgae Nieto Nafría & Brown Apterae are shining black, sometimes with a thin layer of whitish powder; BL 1.1-1.9 mm. Alatae have 3-7 secondary rhinaria on ANT III. On stems and proximal parts of leaves of Baccharis spp, especially B. linearis, often forming dense groups (Nieto Nafría et al. 2019c). In Chile, between Santiago and Los Lagos regions. Sexual morphs are unknown.

Aphis intrusa Ortego Apterae are shining dark brown: BL 1.3-1.9 mm. Apterae have 0-4 (-7) secondary rhinaria on ANT III and alatae have 10-13 (2 specimens). On Senecio subumbellata in Argentina (Ortego 1998a). Very close to A. craccivora; further collections are needed to confirm the distinction.

Aphis intybi Koch Apterae are black, wax-powdered (see influentialpoints.com/Gallery); BL 1.2-2.3 mm. Alatae have 2-9 secondary rhinaria on ANT III, and 0-1 on IV. On young growth of Cichorium intybus in spring, and later at stem bases in ant shelters. In Europe, Mediterranean region, west and central Asia east to Pakistan (Naumann-Etienne & Remaudière 1995). It has been intercepted at ports of entry to USA (Stoetzel & Russell 1991), but specimens in the Canadian National Collection indicate that it may be established in North America. It has also been introduced to Argentina (Mier Durante et al. 2012) and Chile (Nieto Nafría et al. 2018). Monoecious holocyclic on Cichorium, with apterous males (Tuatay & Remaudière 1964). Records from other plants are most probably misidentifications. Populations in the Mediterranean/ Middle East region differ somewhat from those in north and central Europe by having few if any marginal tubercles on abdominal segments 2-6, and longer hairs on abdominal tergite 8.

Aphis iranensis García Prieto & Nieto Nafría Apterae are green with dark head and siphunculi; BL 0.7-1.3 mm. Holocyclic and probably monoecious on Rhamnus pallasii in Iran (García Prieto & Nieto Nafría 2007). The oviparae (collected in October) have hind tibiae unswollen with very few scent glands, indicating that males are probably apterous.

Aphis ishkovi Kadyrbekov Apterae are pale brown tinged with grey, with dark head, siphunculi and cauda; BL 1.7-2.1 mm. In small ant-attended colonies on green shoots and in inflorescences of Myricaria bracteata in south-east Kazakhstan.

Aphis iteae (Tissot) Apterae are brown; BL. c.1.2 mm. Alatae have 5-9 secondary rhinaria on ANT III. On Itea virginica (= Cyrilla racemiflora) in Florida, USA.

Aphis jacetana García Prieto & Nieto Nafría Apterae are black, BL 1.3-1.8 mm. On young growth of Pimpinella saxifraga in Spain (Pyrenees). Alatae and life cycle are unknown (García Prieto & Nieto Nafría 2005).

Aphis jacobaeae Schrank Apterae are very dark green to dark brown or black (see influentialpoints.com/Gallery); BL 1.5-2.6 mm. Apterae have secondary rhinaria distributed III 0-12, IV 1-15, V 0-3, and alatae have them distributed III 17-38, IV 0-20, V 0-8. In compact colonies on stems, inflorescences and undersides of leaves of Senecio, Parasenecio and Tephroseris spp., especially Senecio jacobea, attended by ants, and sometimes in ant shelters at stem bases (Stroyan 1984). In western, central and eastern Europe, and also recorded from Kazakhstan (Kadyrbekov & Aoitzhanova 2005, Kadyrbekov 2009a). Monoecious holocyclic according to Börner (1952).

Aphis janischi (Börner) Apterae are black, wax-dusted; BL 1.8-2.7 mm. On leaf bases and stems of Cirsium oleraceum (rarely arvense) in Europe (Denmark, Finland, Sweden, Germany, Poland, Austria). It is also reported from several Cirsium spp. in Kazakhstan (Kadyrbekov 2009b, 2017a), and there are records from Cirsium incanum and Sonchus arvensis in Kirghizia (Kadyrbekov 2013c). Further studies are needed to show whether or not this species is biologically distinct from A. fabae s. lat., as there are no reliable morphological discriminants, especially from A fabae cirsiiacanthoidis which preferentially colonises Cirsium and also has a high incidence of abdominal marginal tubercles.

Aphis junelliae González Rodríguez & Nieto Nafría Apterae are green with a reddish patch between the siphunculi, which are brown, and a dusky cauda; BL 0.9-1.2 mm. Forming a small compact colony on a twig of Junellia crithmifolia in Argentina (San Juan province). The single known alata has 5 secondary rhinaria on ANT III (González Rodríguez et al. 2018). Other morphs are unknown.

Aphis jurineae Bozhko Apterae are blackish brown; BL 2.0-2.3 mm. Alatae have 4-6 secondary rhinaria on ANT III. Forming large colonies on stems of Jurinea cyanoides in Ukraine, and also recorded from J. multiflora in Kazakhstan (Kadyrbekov 2011b, 2017a).

Aphis kachkoulii Remaudiere Apterae are pruinose; in spring 1.2-1.5 mm and pale green, in autumn only 0.5-0.8 mm and pale yellow or whitish. In sometimes dense colonies on leaves of Ficus carica, and F. johannis (probably the original host), in Iran (Remaudiere 1989), and now also reported from Turkey (Görür et al. 2011a). Monoecious, with apterous males. Antennae are 5-segmented in both apterae and alatae, and also in males (sometimes 4-segmented in dwarf apterae).

Aphis kamtchatica Pashchenko Apterae are black, imm. greenish black; BL c.2.7 mm. Alatae have secondary rhinaria distributed III 20-27, IV 0-9. On young shoots, stems and undersides of leaves of Spiraea stevenii in east Siberia (Kamchatka). Ant-attended. Monoecious holocyclic, with fundatrices in late June (original description).

Aphis khrulevae Stekolshchikov & Buga Apterae are black with dark siphunculi and cauda; BL 2.0-2.3 mm. On Veronica longifolia, on shoot apices between flowers and on undersides of leaves that become rolled and crinkled. In north-west Russia, and in the north of central Siberia (Taimyr Peninsula). Monoecious holocyclic; fundatrices were collected in June-July (Stekolshchikov & Buga 2018).

Aphis klimeschi (Börner) Apterae are greyish green to dark greenish brown, not waxy; BL 1.2-2.3 mm. Alatae have secondary rhinaria distributed III 6-10, IV 0-5, V 0-2. In basal ant-shelters on Anthyllis vulneraria in Europe (not Italy or Iberian peninsula). Monoecious holocyclic with apterous males.

Aphis kobachidzei (Rusanova) Apterae are pale brown or reddish brown (BL 1.6-2.0 mm) on Typha angustifolia in Azerbaijan. They were described in the genus Brachysiphum and transferred to Aphis by Favret et al. (2017). The siphunculi are very short (less than half the length of the cauda), and there are marginal tubercles on ABD TERG 1-4 and 6-7. Monoecious holocyclic; an ovipara was part of the original collection.

Aphis kogomecola Matsumura Apterae are yellowish-green; BL c.1.5 mm. On Spiraea thunbergii in Japan, rolling leaves together. Not since recognised, and the fact that the cauda was not evident suggests that it might be a Brachycaudus, possibly B. helichrysi.

Aphis korshunovi Ivanoskaya Apterae are dull black, without wax, antennae and legs dark except for basal part of ANT III, femoral bases and middle parts of tibiae, siphunculi and cauda black; BL 0.9-2.2 mm. Alatae have secondary rhinaria distributed III 5-9, IV 0-1. On Veronica spp. in northern Europe (Denmark, Finland) and east to west Siberia and east Kazakhstan. Monoecious holocyclic with oviparae and apterous males in late August (Heikinheimo 1978, as A. pseudolysimachiae).

Aphis kosarovi Tashev, Popov & Berova Apterae are black, resembling A. fabae; BL 1.9-2.5 mm. On Digitalis spp. in eastern Europe (Bulgaria). Monoecious holocyclic; oviparae (with only slightly thickened hind tibiae) and alate males were collected in October. [Originally described as a subspecies of A. armata, but having much shorter hairs. The specimens in the original description (Tashev et al. 1970) were noted as lacking symbionts. Three apterae on a slide from Tashev in the BMNH collection have leg hairs mostly short, but with a few hairs long and distorted. This raises the possibility that the shortness of the hairs is due to nutritional deficiency in an asymbiotic population of A. armata.]

Aphis kurosawai Takahashi (Fig.12t) Apterae are green or yellow-green, wax-powdered, with dark siphunculi and cauda (see aphids of Karnataka website); BL 1.1-2.0 mm. A common species on leaves of Artemisia spp. in eastern Asia (India, Nepal, east Siberia, China, Vietnam, Laos, Korea, Taiwan, Japan). Holman (1987) lists morphometric data in comparison with other Artemisia-feeding species. Holman’s data for A. kurosawai from the east Siberia agree with BMNH specimens from Japan and Taiwan, but specimens from the Himalayas have a longer R IV +V (1.5-1.9 X HT II, as opposed to 1.36-1.6 X HT II), and may be a distinct form, for which the name A. lhasartemisiae Zhang (described from Tibet) is available. Bhagat (1981) reported an alate male from Kashmir, and Medda & Chakrabarti (1998) described the ovipara. Some specimens collected on Artemisia lavandulaefolia in China (BMNH collection, leg.V.F.Eastop) appear to be fundatrices. A. artemifoliae Shinji 1922 is a synonym according to Takahashi (1966). 2n=8.

Aphis laciniariae Gillette & Palmer Apterae are pale green (Hottes & Frison 1931, as A. zilora); BL 1.1-1.4 mm. Alatae have 4-8 secondary rhinaria on ANT III. On leaves and stems of Liatris spp. in USA. Oviparae and apterous males in Colorado in September-October (Palmer 1952).

Aphis lambersi (Börner) Apterae are dark green, purplish brown or almost black (see influentialpoints.com/Gallery); BL 1.3-2.2 mm. Alatae have secondary rhinaria distributed III 7-19, IV 0-7, V 0-2. On root collar and in basal leaf sheaths of Daucus carota, ant-attended. Also recorded from Pimpinella saxifraga (Nieto Nafría & Mier Durante 1985) and Conopodium majus (Nieto Nafría et al. 2005a), and a large sample of summer dwarfs from Foeniculum in the Italian Alps (BMNH collection, leg. R. Stäger) may also be this species. Throughout Europe, and also in the Middle East (Iran; Rezwani et al. 1994) and North Africa. Monoecious holocyclic with apterous males. 2n=8.

Aphis lamiorum (Börner) Apterae are dark blue-green to dark green (see influentialpoints.com/Gallery); BL 1.6-2.2 mm. On Lamium spp., causing downward leaf-curl of shoot apices. Alatae have secondary rhinaria distributed III 6-19, IV 0-7. Ant-attended. Monoecious holocyclic, with oviparae present in mid-August (BMNH collection, leg. V.F.Eastop) and having hind tibiae only slightly swollen, and apterous males (original description). Closely related to the frangulae group; spring and summer apterae are practically indistinguishable from those of the host-alternating A. frangulae ssp. beccabungae, which migrates to Lamium from Frangula (Stroyan 1984). Apparently widely distributed in Europe, but some of these records may be summer colonies of A. frangulae beccabungae.

Aphis lantanae Koch Apterae are dark greenish brown with variably developed dark dorsal sclerotic markings (see influentialpoints.com/Gallery); BL 1.5-2.1 mm. Alatae have secondary rhinaria distributed III 17-28, IV 1-11, V 0-3. In ant-attended colonies curling leaves, on young stems or under senescing leaves of Viburnum lantana. Widely distributed in Europe. Monoecious holocyclic with alate males. Stroyan (1984) gave subspecies status to an alpine form, A. lantanae coriaria Börner, found in Austria, Switzerland, Italy and Andorra (Nieto Nafría et al. 2005a), which, apart from the characters given in the key, has longer antennal hairs and apterous or alatiform males. Specimens collected on Sorbus aria in Bavaria, Germany, in 1933 were described as a new species under the name Doralis setacea Hille Ris Lambers, but this name was synonymised with A. lantanae by Eastop & Hille Ris Lambers (1976).

Aphis leontodontis (Börner) Apterae are greenish black (see influentialpoints.com/Gallery); BL 1.2-1.7 mm. Alatae have secondary rhinaria distributed III 5-10, IV 1-3, V 0-1. On Leontodon spp., colonising undersides of etiolated basal parts of leaves at soil level, often in ant shelters. In Europe, (Denmark, Sweden, Finland, Germany, Poland, Byelorussia, Czech Republic). Monoecious holocyclic. Records of A. taraxacicola from Leontodon should probably be referred to this species.

Aphis lhasaensis Zhang Apterae are presumably shining black; BL c. 2.3 mm. On Phaseolus vulgaris in Tibet. Possibly large specimens, or a variant population, of Aphis craccivora, which may sometimes have marginal tubercles consistently on abdominal tergites 2-4 (see Stroyan 1984). [This is now supported by DNA (COI) evidence – see J. Wang et al. 2011.]

Aphis lichtensteini Leclant & Remaudière Apterae are shining pale yellow to green, with dark apices to antennae and legs, and black siphunculi and cauda: BL 0.9-1.7 mm. Alatae have secondary rhinaria distributed III 4-11, IV 0-2. On Cistus spp., especially C. monspeliensis, around the Mediterranean (southern France, Corsica, Sardinia, Sicily, Spain, Algeria). It tends to form small colonies on leaves and along the flower stalks, causing some deformation in the form of spiral rolling of the leaves. Only rarely ant-attended. Apparently anholocyclic.

Aphis ligulariae Holman Colour of apterae colour in life unknown, presumably dark with antennae and tibiae mainly pale, siphunculi and cauda dark; BL 2.1-2.2 mm. On basal parts of Ligularia schmidti in east Siberia. Presumably monoecious holocyclic.

Aphis (Anthemidaphis) ligusticae Barbagallo & Stroyan Apterae are yellow-green to yellowish white or cream; BL 1.0-1.7 mm. Alatae are undescribed. On underground stolons of Achillea ligustica, attended by ants. Only known from Italy (Sicily). Monoecious holocyclic with oviparae in late September-November (original description).

Aphis liliophaga Holman Apterae are dull dark green; BL 1.7-2.2 mm. Alatae have secondary rhinaria distributed III 6-10, IV 0-1. On seed capsules of Lilium tenuifolium in Mongolia.

Aphis limonicola Pashchenko Apterae are black; BL c. 1.2 mm. On undersides of leaves of Limonium platyphyllum in Russia (on a plant that was shipped from Altai to Vladivostock).

Aphis lindae Danielsson Colour of apterae in life unknown; BL 1.2-1.6 mm. On basal part of stem of Tripleurospermum maritimum, growing between stones near seashore, not attended by ants. Only known from Sweden.

Aphis lini (Bozhko) Apterae are brown, wax-dusted, with black siphunculi and cauda; BL c. 2.0-2.1 mm. In dense colonies on stems and young shoots of Linum spp. in eastern Europe (Serbia, Ukraine).

Aphis linorum (Bozhko) Apterae are dark green to nearly black, with antennal flagellum, femora and tibiae whitish basally and black distally; BL 1.4-1.9 mm. In ant-attended colonies on the root collar of Linum spp. in eastern Europe (Slovakia, Romania, Ukraine). Monoecious, with oviparae and apterous males in October (Holman 1966, as Aphis lini).

Aphis lippiae González Rodríguez & Nieto Nafría Apterae are greenish yellow with dark brown siphunculi and pale cauda; BL 1.0-1.4 mm. In small groups on twigs of Lippia turbinata in Argentina (provinces Córdoba, Mendoza, La Rioja). Alatae have secondary rhinaria distributed III 4-12, IV 1-4, V 0-2. Other morphs and life cycle are unknown (González Rodríguez et al. 2018).

Aphis lithospermi Wilson Apterae are dark green with black dorsal markings, and black antennae, siphunculi and cauda; BL c. 2.1 mm. Alatae have secondary rhinaria distributed III 9-12, IV c.2. On Lithospermum pilosum (= L. ruderale?) in California and Oregon, USA.

Aphis longicauda (Baker) Apterae of variable colour, brown or yellowish-white with dark antennae and legs, and dark posterior abdominal markings; BL c.1.7-1.8 mm. On twigs and leaves of plum trees (Prunus domestica) in USA, recorded as very injurious to plum in Virginia, with twigs dying the year after attack (Baker 1920a), but little known since original description. Monoecious, with apterous males. The description resembles that of A. chetansapa.

Aphis longini Huculak Apterae are matt dirty green to olive brown; BL 1.7-2.3 mm. In large ant-attended colonies at bases of stems of Knautia arvensis in Poland. Monoecious, with oviparae and apterous males in October (original description). Kanturski & Bezdĕk (2019) provided a redescription.

Aphis longirostris (Börner) (= longirostrata Hille Ris Lambers) Apterae are dark green to dull bluish green (see influentialpoints.com/Gallery); BL 1.0-1.8 mm. Alatae are unknown. In ant shelters on root collars and etiolated leaf bases of Plantago spp., especially P. maritima, but not only in coastal habitats. In Europe, west Siberia, and Central Asia (Kadyrbekov 2004b). Monoecious with apterous males.

Aphis longisetosa Basu Apterae are pale whitish with legs pale, siphunculi and cauda dark; BL 1.2-1.7 mm. Alatae have 3-8 secondary rhinaria on ANT III (Chakrabarti & Raychaudhuri, 1975, as A. ruborum ssp. longisetosa). On undersides of leaves of Rubus spp. in north India, Pakistan, Nepal and Thailand. 2n=6 and/or 2n=8 (see references in Kar et al. 1990; the discrepancy requires further investigation).

Aphis longituba Hille Ris Lambers (= clematidis ssp. simalensis Kumar & Burkhardt) Apterae are pale green, mottled with darker green on abdomen, with antennae and legs pale, and siphunculi black-tipped; BL 1.3-1.6 mm. On young leaves of Clematis grata and Clematis sp. in Pakistan and India, and now also recorded from C.vitalba in Turkey (Görür et al. 2011b). The small cauda combined with dark-tipped siphunculi are distinctive. 2n=8.

Aphis loti Kaltenbach (Fig.41j) Apterae are warm dark brown, without wax; BL 1.2-2.1 mm. Alatae have secondary rhinaria distributed III 4-8, IV 0-2. On shoot apices and inflorescences of Lotus and Dorycnium spp., and Anthyllis vulneraria, sometimes ant-attended. Widely distributed in Europe, eastward to Turkey (Akyürek et al. 2011), Iran (Rezwani 2010) and Kazakhstan (Kadyrbekov 2017a). Monoecious with apterous males. Some records of A. craccivora on Lotus may be this species. 2n=8.

Aphis lotiradicis Stroyan Apterae are dirty greenish brown; BL 1.25-1.9 mm. Alatae have secondary rhinaria distributed III 4-8, IV 0-3. On basal parts of stem and roots of Lotus spp. in sandy soils, tented over by ants. In Netherlands, UK, Germany, Spain, Finland, Poland, Rumania, Serbia, Czech Republic and Kazakhstan, where it is also recorded from Medicago falcata (Kadyrbekov 2017a). Monoecious; oviparae and apterous males occur in October.

Aphis lugentis Williams Apterae are dark yellow-brown with greenish tinge, to dark olive or dull blackish green, with entirely dark appendages; BL 1.9-2.8 mm. Apterae have secondary rhinaria distributed III 0-11, IV 0-11, V 0-2; in alatae they are distributed III 19-39, IV 7-20, V 0-4. On leaves, stems and roots of Senecio spp., and an Erigeron sp., in USA (except the north-east), Mexico and western Canada. In recent years it has also been reported from South America (Peru, Chile, Argentina), from southern France and Tunisia (Martinez et al. 2014, Ortego et al. 2019), and from Australia (Petit et al. 2021) . Identification of Argentinian, Tunisian and Australian populations as A. lugentis is supported by DNA analysis (Ortego et al. 2019, Petit et al. 2021). It can form dense colonies, and is often attended by ants. In Argentina it has been found colonising some endemic Asteraceae, and has also been observed to be displacing native Senecio-feeding Aphis species (Ortego et al. 2019). Monoecious holocyclic, with oviparae and alate males in October (Palmer 1952, and as A. nyctalis). Although A. nyctalis seems to be a synonym, there are other closely-related North American species on Senecio with longer and finer hairs, including A. senecionis (q.v.) and an undescribed species lacking marginal tubercles. These all belong to a holarctic group of Senecio-feeders, characterised by apterae with dark tibiae and rhinaria on ANT IV, that needs further investigation.

Aphis lupinehansoni Knowlton Colour of apterae in life unrecorded; BL 1.7-2.4 mm. On Lupinus sp. in north-western USA.

Aphis lupini Gillette & Palmer Apterae are dusky olive to greenish black (see aphidtrek.org); BL 2-3 mm. On leaves and stems of Lupinus decumbens (= argenteus) in western USA . Monoecious holocyclic (Palmer 1952), with apterous males.

Aphis lupoi Barbagallo & Stroyan Apterae are green or yellow-green, wax-dusted, with head brownish; BL 1.0-1.8 mm. Alatae have 3-8 secondary rhinaria on ANT III. The original description provided a detailed morphological comparison with A. cisticola. On stems and undersides of leaves of Cistus spp. in Sicily, where it is probably anholocyclic (original description). Also recorded from Portugal (Nieto Nafría et al. 2005a). Sometimes visited by ants.

Aphis lupuli Rusanova Apterae are greenish yellow, with dark siphunculi and a yellow cauda; BL 1.6-1.7 mm. Alata (one specimen) has secondary rhinaria distributed III 8, IV 4. In large colonies on shoots of hops (Humulus lupulus) in Azerbaijan. Possibly this was A. nasturtii.

Aphis luzuriagae López Ciruelos, Brown & Nieto Nafría Colour in life unknown, BL of aptera 1.4-2.0 mm. Alatae have secondary rhinaria distributed ANT III 6-10, IV 1-4. On Luzuriaga radicans and Luzuriaga sp. in southern Chile (Osorno, Chiloé). Sexual morphs and life cycle are unknown (López Ciruelos et al. 2018).

Aphis lycopicola (Shinji) Apterae are yellowish to yellowish green with dark siphuculi and cauda; BL c. 0.9 mm. Both apterae and alatae have c.5 rhinaria on ANT III. On Lycopus spp. in Japan.

Aphis lysimachiae Bozhko Apterae are grass green; BL c. 1.9 mm. On Lysimachia vulgaris in Ukraine, and also in west Kazakhstan (Smailova 1980).

Aphis maculatae Oestlund Host-alternating between Cornus stolonifera and Populus spp., where it sometimes forms large colonies on twigs and leaves, throughout most of North America. Apterae on Populus in summer are brownish black, with black siphunculi and cauda and paler head, legs and antennae, the body marked with conspicuous patches of white wax powder (Palmer 1952 and see influentialpoints.com/Gallery). Wilson & Moore (1986) studied the differential susceptibility of various Populus hybrids to this species, and discussed insecticidal control measures. [Records from Tunisia (Ben Halima-Karmel 2012) and Turkey (Kök & Özdemir 2021) need further confirmation.]

Aphis madderae Robinson Colour of apterae in life is unrecorded; BL 0.9-1.3 mm. On root crown of Taraxacum lacerum (= ceratophorum) in northern Manitoba, Canada.

Aphis madronae (Essig) Apterae are reddish-brown throughout; BL c.2.1-2.2 mm. In small or large dense colonies on trunk and lower leaves of Arbutus menziesi in California. The aphids match the colour of the bark of the tree. Not recorded on Arbutus since the original collection, but specimens from Arctostaphylos sp. (BMNH collection, leg. R.C. Dickson) have been identified as this species. The life cycle is unknown.

Aphis magellanica Nieto Nafría, Mier Durante & Ortego Apterae are emerald green with dark green siphunculi and long pale cauda; BL 1.9-2.4 mm. Living on Lathyrus magellanicus in the Antarctic region (Magellanes) of southern Chile. Oviparae and alate males were collected together with apterous viviparae in February (Nieto Nafría et al. 2020a).

Aphis magnopilosa Nevsky Apterae are blackish-brown or black, BL c.2.5 mm. A long-haired member of the Aphis fabae group (q.v.) described from Centaurea sp. in Uzbekistan, and also recorded from other countries in Central Asia. Morphological variation in subsequent collections of A. fabae-like aphids on Centaurea spp. in Uzbekistan was discussed by Holman (1987).

Aphis malhalhuina Mier Durante, Nieto Nafría & Ortego Apterae are dark brown to lack, rather shiny; BL 1.3-2.1 mm. Alatae have secondary rhinaria distributed III 4-12, IV 0-4. In dense colonies on stems of Senecio subumbellatus, sometimes near the root. In Argentina. Monoecious holocyclic with alate males (Mier Durante et al. 2003).

Aphis mammulata Gimingham & Hille Ris Lambers Apterae are 1.3-1.9 mm, greenish-grey, BL 1.3-1.9 mm. Alatae have secondary rhinaria distributed III 7-12, IV 0-3, V 0-1 (Heie 1986). Found in autumn in dense, ant-attended colonies on petioles of berries and undersides of leaves of Rhamnus cathartica. Monoecious holocyclic; the colonies include oviparae and orange-brown apterous males. Appearance of the colonies earlier in the season is unrecorded. Recorded from UK, Sweden, Finland, Germany, Poland, Czech Republic, northern Spain, Russia (as A. rhamnicola Mamontova) and Kazakhstan (Kadyrbekov 2004b, 2012a). Börner (1952) and Holman (in Remaudière & Remaudière 1997, p. 42) regarded the shorter haired A. versicolor as a synonym.

Aphis mamonthovae Davletshina (= Aphis verbenae Nevsky, 1929, nec. Macchiati, 1883) Apterae are brown with rather dark brown antennae and legs, dark brown siphunculi and pale brown cauda; BL 1.2-1.8 mm. Alatae have secondary rhinaria distributed III 6-10, IV (0-)1-2. On Verbena spp., especially V. officinalis. In central and southern Europe, north Africa (Ethiopia), Middle East (Lebanon, Turkey), Central Asia and northern India (BMNH collection, leg. S.S. Sekhon). Closely related to the A. frangulae group, and European records of this group from Verbena are probably referable to this species. Anholocyclic in Sicily (Barbagallo & Stroyan 1982), sexual morphs unknown.

Aphis (Bursaphis) manitobensis Robinson & Rojanavongse Apterae are pale green with pale siphunculi; BL 1.6-2.2 mm. On Ribes sp. in Canada (Manitoba), and on R. nigrum in Mongolia (BMNH collection., leg. H. Szelegiewicz). Monoecious holocyclic with apterous males (BMNH collection). Its distinction from A varians needs verification (Rakauskas et al. 2011).

Aphis marthae Essig Apterae are shining black; BL 1.5-2.5 mm. Alatae have secondary rhinaria distributed III 11-21, IV 0-5. Described from an unknown shrub or tree, and subsequently collected from Quillaja saponaria (Remaudière 1994). In Chile. There is a very similar species in the BMNH collection, also from Chile, leg. I. Rosembaum, collected on Cryptocarya alba.

Aphis martinezi Nieto Nafría, Ortego & Mier Durante Apterae are black, BL 0.85-1.5 mm. Alatae have 5-7 secondary rhinaria on ANT III. In dense colonies on stems and leaves of Mulinum spinosum in Mendoza Province, Argentina (Nieto Nafría et al. 1999b).

Aphis masoni Richards Apterae are blue-green, wax-powdered; BL 1.9-2.4 mm. Alatae have secondary rhinaria distributed III 8-15, IV 1-4. On Oxytropis spp. in Canada (Baffin Island, Victoria Island, and Churchill, Manitoba (original description and BMNH collection, leg. W.R. Richards). Also on Astragalus alpinus; a slide in the BMNH collection labelled as a paratype (coll. W.R.M. Mason, leg. W.R. Richards and D. Hille Ris Lambers) has an aptera collected from this host on Baffin Island, 11.viii.1959, although this data is not listed in the original description. Apart from the shorter siphunculi it is very similar to A. astragali in northern Europe.

Aphis mastichinae Pérez Hidalgo & Nieto Nafría Apterae are green, greenish blue or yellowish green; BL 0.9-1.2 mm. Alatae have secondary rhinaria distributed III 1-6, IV 0-1. On roots of Thymus mastichina in northern Spain. Monoecious holocyclic with apterous males (Pérez Hidalgo & Nieto Nafría 2004).

Aphis matilei Nieto Nafría, Ortego & Mier Durante Apterae are pale green, BL 0.95-1.25 mm. Alatae not recorded. In compact colonies on the flower-stems of Verbena glauca (= Junellia spathulata var. glauca ) in Argentina (Nieto Nafría et al. 2000). Monoecious holocyclic, with oviparae and apterous males in March.

Aphis matricariae Barjadze & Özdemir Colour of apterae in life is unknown; BL 1.2-1.6 mm. Alatae (2 specimens) have secondary rhinaria distributed ANT III 13-16, IV 0-1, V 0. On a species of Matricaria in northern Turkey (Barjadze et al. 2014). Biology unknown.

Aphis maulensis Mier Durante & Garcia Tejero Apterae are yellowish-green, often with a variably developed dark brown-black dorsal abdominal patch, and with brownish appendages and black siphunculi; BL 1.6-2.2 mm. Alatae with 4-9 secondary sensoria on ANT III. On Euphorbia sp. in Chile at 2430 m (López Ciruelos et al. 2016). Life cycle unknown, but presumably monoecious holocyclic.

Aphis medicaginis (Koch) Apterae are grey-green to black, wax-dusted; BL 1.7-2.0 mm. On basal parts and roots of Medicago and Melilotus spp. Specimens on Ononis spinosa from France (BMNH collection, leg. F. Leclant) are also possibly this species. Widely distributed in Europe (but not recorded from the Iberian peninsula), and also recorded from Kazakhstan (Kadyrbekov 2017a). Many records prior to 1950, including all those from North America, are referable to A. craccivora. A record from Ononis natrix ssp. angustissima in Algeria (Laamari et al. 2013) requires additional confirmation. Hoffmann (1972) provided a redescription (as meliloti Börner). Monoecious holocyclic.

Aphis melosae Mier Durante & Ortego Apterae in spring and autumn shining dark brown to blackish brown with BL 1.5-2.1 mm, in summer light brown to dark green with BL 0.9-1.5 mm. Apterae have 0-6 secondary rhinaria on ANT III, and alatae have them distributed III 7-13, IV 0-4, V 0-2. In dense colonies on stems, leaf axils and undersides of leaves of Grindelia spp. in Argentina (Mier Durante & Ortego 1999), and also found on Haplopappus sp. (BMNH, leg. C.I.B.C.).

Aphis mendocina Mier Durante, Ortego & Nieto Nafría Apterae are shining black (large individuals) or dark green to dark brown (small individuals), with parts of antennae and legs yellow; BL 1.1-2.3 mm. Monoecious holocyclic on Urtica mollis in Argentina (Mendoza), forming dense colonies on stems and leaf petioles. Small (BL less than 1.5 mm) aestivating apterae occur in summer (January-February), and oviparae and apterous males were collected in March (Mier Durante et al. 2006). [Collections from two species of Adesmia (Fabaceae) at many localities in Argentina could not be distinguished from A. mendocina either by multivariate morphometrics (principal components) or by DNA sequence analysis, and were therefore provisionally identified as this species (Nieto Nafría et al. 2016c). However, apterae of the Adesmia aphid are brownish or reddish brown, males are alate. and the disparity between host plants leads to the conclusion that further verification is needed.]

Aphis mimuli Oestlund Apterae are pale to mid yellowish green or apple-green with pale siphunculi and cauda; BL 1.9-2.2 mm. Spring colonies curl young leaves of Ribes spp. (aureum, nigrum) in North America. Heteroecious holocyclic, migrating in May-June to plants in wet situations (Mimulus, Rorippa, Veronica, Bidens cernua), paralleling A. triglochinis in Europe (Hille Ris Lambers 1974b, as A. ribiensis). Cook (1984a) gave a redescription.

Aphis minima Tissot Apterae are yellowish-brown to dark reddish-brown with darker siphunculi and cauda; BL c.1.0-1.1 mm. On young twigs of Prunus americana, and also feeding along main veins of leaves, which may become tightly curled. Only recorded from Florida, USA. Monoecious holocyclic, with apterous males (Tissot 1933).

Aphis (Zyxaphis) minutissima (Gillette & Palmer) Apterae are dark brown to black, covered with a white powdery reticulation; BL 0.9-1.25 mm. On leaves and stems of Artemisia filifolia and Seriphidium tridentatum in western USA. Oviparae and alate males were collected in October (Palmer 1952).

Aphis mirifica (Börner) Apterae are yellowish, green or mottled bluish green with black siphunculi; BL 1.2-2.2 mm. Alatae have secondary rhinaria distributed III 6-9, IV 0(-1), V 0. On Epilobium angustifolium, forming ant-attended colonies on flowerheads, stems and undersides of leaves (which may be curled), or in ant shelters on stem at or below ground level (where the aphids may be yellowish). Mainly recorded from northern Europe, but it has also been found in Italy (Barbagallo & Patti 1998), in Russia (west Siberia, Altai) and in Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic, with apterous males.

Aphis mizutakarashi Shinji Apterae are green with slight bluish-white waxy pulverulence, with mainly yellowish antennae, and dark siphunculi and cauda (Moritsu 1983); BL not recorded. Immatures are very pale yellow-green. On Brassicaceae (Barbarea, Cardamine) in Japan.

Aphis mizzou Lagos & Puttler Apterae are black dusted with white wax, with black femora and mainly yellowish tibiae; BL 1.4-2.2 mm. Alatae have secondary rhinaria distributed ANT III 7-14, IV 0-3. On new terminal growth and stems of Hypericum kalmianum in Missouri, USA (Lagos et al. 2012). Life cycle and sexual morphs are unknown.

Aphis mohelnensis Holman Apterae are bluish green with dark head, siphunculi and cauda; BL 1.2-1.9 mm. On Hieracium spp., living on terminal parts, mainly in the inflorescences. In Czech Republic, Bulgaria, west Siberia, Uzbekistan (Holman 1998) and Kazakhstan (Kadyrbekov 2013d).

Aphis molluginis (Börner) Apterae are green, BL 1.2-1.7 mm. Alatae have c.2 secondary rhinaria on distal part of ANT III). At stem bases or on subterranean parts of Galium spp. in north and north-west Europe, west Siberia, Kazakhstan and Mongolia. A record from Turkey (Görür et al. 2011b) perhaps requires further confirmation. Oviparae are known from Denmark (Heie 1986).

Aphis monardae Oestlund Apterae are pale yellow to pale green, waxy, with siphunculi dusky to dark on distal part and pale cauda; BL 1.1-1.8 mm. Alatae have secondary rhinaria distributed ANT III 4-9, IV 1-3, V 0. On undersides of leaves of Monarda fistulosa, causing leaf-curl, in mid-western USA. Monoecious holocyclic on Monarda, with oviparae and apterous males in early October (Lagos-Kutz et al. 2014). This species is very similar in morphology to both A. nasturtii and A. gossypii; Lagos-Kutz et al. (2014) provide some distinguishing features.

Aphis mongolica Szelegiewicz Apterae are greenish yellow with dark siphunculi and cauda; BL 0.8-2.0 mm. Alatae (Holman 1988) have 7-14 secondary rhinaria on ANT III (= III + IV fused). On ground-level stems of Potentilla tanacetifolia var. filipendula and Potentilla sp. in Mongolia and east Siberia. Monoecious holocyclic (original description).

Aphis montanicola Hille Ris Lambers (= pulsatillae Ossiannilsson 1959) Apterae are shiny bronze-black (original description) to dull dark green or greenish black with mainly pale antennae and tibiae; BL 1.5-2.0 mm. Alatae have 3-7 secondary rhinaria on ANT III. On Pulsatilla spp. and Anemone spp. in Europe (Switzerland, Sweden) and also recorded from west Siberia, Kazakhstan, Mongolia and east Siberia (Holman 1987; as A. pulsatillae). In Kazakhstan it is also recorded from Adonis vernalis (Kadyrbekov 2017a) Colonising stem bases early in season, and later on leaves and flowers; attended by ants. Monoecious holocyclic; sexual morphs are unknown, but fundatrices have been described from Pulsatilla (= Anemone) montana (original description) and from P. vulgaris (= Anemone pulsatilla) (Ossiannilsson 1959).

Aphis mulguraeae Nieto Nafría & Mier Durante Apterae are pale yellow or green with yellow or reddish markings, siphunculi almost black and cauda pale; BL 1.0-1.3 mm. On twigs of Mulguraea aspera in Argentina (Mendoza province). Alatae have 4-6 rather large secondary rhinaria on ANT III only (González Rodríguez et al. 2018). Other morphs and life cycle are unknown.

Aphis mulini Hille Ris Lambers Colour of apterae in life unknown, probably blackish; BL 1.7-2.0 mm. Alatae are undescribed. On Mulinum sp. (probably spinosum) in Patagonia, Argentina. [A. mulini is one of a group of South American Aphis with thick siphunculi and dorsal abdominal sclerotisation that is often fragmented. A. melosae is also closely related, and has very different-looking spring and summer generations, suggesting a need for more detailed studies on the Mulinum feeders.]

Aphis mulinicola Hille Ris Lambers Colour of apterae in life unknown, probably blackish; BL c. 1.8-1.9 mm. Alatae are undescribed. On Mulinum sp. (probably spinosum) in Patagonia, Argentina.

Aphis multiflorae Barbagallo & Stroyan Apterae are green with brown head, siphunculi and cauda; BL 0.9-1.4 mm. Alatae have secondary rhinaria distributed III 3-5, IV 0-2. On apical parts of young shoots of Erica spp. and Daboecia cantabrica. In Italy (Sicily, Tuscany), south of France and Spain. Probably anholocyclic (Barbagallo & Stroyan 1982).

Aphis mutini Pashchenko Colour of apterae in life unknown, probably with shining dark brown or black dorsal shield; BL 1.8-2.2 mm. Alatae have 7-9 secondary rhinaria on ANT III. On Spiraea media in east Siberia, and since recorded from Spiraea sp. in western Siberia (Altai Republic) by Stekolshchikov & Novgorodova (2013), who noted some differences from the type material. Fundatrices were found in early June (original description).

Aphis myopori Macchiati Apterae are predominantly dark green, BL not recorded. Described from a plant named as “Myoporum pictum” in Sardinia, and subsequently found on the same plant in mainland Italy (Calabria; Macchiati 1882). Possibly it was A. gossypii.

Aphis myrsinitidis Petrović & Leclant Apterae are dark brown to very dark brown-red with thick white pleural wax spots; BL 1.7-2.1 mm. Alatae have secondary rhinaria distributed III 9-11, IV 3-5, V 1-3. On Euphorbia myrsinites in the Balkans (Petrović & Leclant 1998), and also found on Euphorbia spp. in Jordan, Lebanon, Iran, Turkey and Morocco (BMNH collection, leg. T. Mustafa, D. Hille Ris Lambers, E.S. Brown, N. Tuatay and J.H. Martin respectively). Monoecious holocyclic; oviparae and alate males were collected in Iran in late October (BMNH collection).

Aphis narzikulovi Szelegiewicz Apterae are bright yellow with dark head, distal sections of antennae and legs, siphunculi and cauda; BL 0.8-1.7 mm (Holman 1988). Alatae have secondary rhinaria distributed III 7-10, IV 0-4, V 0-3 (fused III+IV with 7-10 in specimens with 5-segmented antennae). Monoecious holocyclic; the original description was of fundatrices which are dark green to greenish black, with black siphunculi and cauda; BL 1.8-2.2 mm. Collected in Mongolia from a plant originally identified as Salvia sp., but later identified as Schizonepeta (= Nepeta) multifida (Holman 1988). One of the many aphids of the A. frangulae group living on Lamiaceae.

Aphis nasturtii Kaltenbach Plate 6g Apterae are yellow to yellowish green with dark apices to antennae and leg segments and siphunculi (see influentialpoints.com/Gallery); BL 1.3-2.0 mm. Alatae have secondary rhinaria distributed III 4-18, IV 0-7, V 0-2. On a wide range of herbaceous plants in summer, including Nasturtium officinale, Solanum tuberosum, Veronica beccabunga, Drosera rotundifolia (Müller, 1978) and Rumex spp.; Stroyan (1984) lists host genera in 28 different families. Now almost world-wide (but not yet in Australasia). Heteroecious holocyclic, with a sexual phase on Rhamnus spp., usually R. cathartica in Europe, but many other Rhamnus spp. can be colonised (see Gleiss 1959, 1966). Apterae on Rhamnus in spring are rather bright green to greenish-yellow, BL 1.6-2.4 mm, and cause distortion of young leaves. 2n=8.

Aphis neilliae Oestlund Apterae are dark olive-green to black or reddish brown (see influentialpoints.com/Gallery); BL 1.3-1.8 mm. Alatae have secondary rhinaria distributed III 25-37, IV 14-23, V 7-15). On leaves and stems of Physocarpus spp. in North America. Monoecious holocyclic with apterous males (Palmer 1952).

Aphis nelsonensis Cottier Colour of apterae in life is unrecorded; BL c. 1.3 mm. Alatae have 2-4 secondary rhinaria on ANT III (BMNH collection, leg. A.D. Lowe). On leaves and stems of an Epilobium sp. in New Zealand. Not found since 1965, and possibly extinct (Teulon et al. 2013).

Aphis neoartemisiphila Pashchenko Apterae are green, BL c.1.4 mm. Alata (1 specimen) has 0-1 rhinaria on ANT III. On stems of Artemisia mandshurica in east Siberia.

Aphis neogillettei Palmer Apterae are 1.0-1.4 mm, olive-green with some red-brown mottling, dusted with grey wax, with blackish siphunculi, cauda and apices of appendages (see aphidtrek.org and influentialpoints.com/Gallery); BL 1.0-1.4 mm. In dense colonies curling leaves of Cornus stolonifera. Monoecious holocyclic, with olive-brown apterous males (Palmer 1952). Widely distributed in North America. A non-host-alternating member of the asclepiadis group (Lagos et al. 2014). 2n=8.

Aphis neomonardae Rojanavongse & Robinson Apterae are are brownish to bluish green, BL 1.2-2.0 mm, on Agastache foeniculum and Monarda fistulosa, in Manitoba, Canada. Alatae are light green to bluish green with brown marginal and postsiphuncular sclerites.

Aphis neonewtoni Pashchenko Apterae are shining dark brown, with black siphunculi and pale cauda; BL c. 1.8 mm. Alatae are undescribed. On upper sides of leaves of Iris uniflora in east Siberia.

Aphis neopolygoni Shinji Aptereae are salmon red; BL c. 1.25 mm. Alatae have secondary rhinaria distributed III 28-32, IV 12-15, V 2-5. On Barbarea spp. and Fagopyrum esculentum in Japan. The short antennal terminal process and alatae with numerous rhinaria suggest that this may be A. triglochinis.

Aphis neospiraeae Takahashi Apterae are reddish brown to dark brown with black dorsal sclerotisation, wax-dusted, middle parts of tibiae whitish; BL 1.3-2.3 mm. Alatae have secondary rhinaria distributed III 6-20, IV 0-7, V 0-3. On shoot apices of Spiraea salicifolia and Rosa maximowicziana in east Asia (east Siberia, Korea, Japan). Monoecious holocyclic in Japan on S. salicifolia; Sugimoto & Seito (2004) studied morphological variation in apterae and alatae and described oviparae and males, collected in September-October, and fundatrices collected in late May.

Aphis neothalictri Pashchenko Apterae are dark green to black, wax-dusted; BL c.2.4 mm. Alatae have secondary rhinaria distributed III 18-27, IV 6-10, V 2-5. On flower stems, pedicels and undersides of leaves of Thalictrum spp. in Siberia, east (Kamchatka) and west (Stekolshchikov & Novgorodova 2013, Novgorodova & Stekolshchikov 2013) and north-west Russia (Stekolshchikov 2017). Described as a subspecies of A. thalictri.

Aphis neothesii Pashchenko Apterae are brown, wax-dusted; BL c.1.8 mm. Alatae have 38-46 secondaru rhinaria on ANT III, and also some on IV and V (numbers not specified). In dense, ant-attended colonies on stems of Thesium chinense in east Siberia.

Aphis nepetae Kaltenbach Apterae are yellowish with head, siphunculi and distal parts of antennae and legs black; BL 1.3-1.7 mm. Alatae have secondary rhinaria distributed III 3-17, IV 0-6, V 0-3. On Nepeta spp. throughout most of Europe (not UK nor Scandinavia, but there is one record from southern Finland, Albrecht 2010), and also reported to occur in Iran (Rezwani et al. 1994) and Kazakhstan (Kadyrbekov 2009b). Also recorded from USA (Smith & Parron 1978), but a record from Minnesota is unsubstantiated (Foottit et al. 2006), and alatae on slides labelled as nepetae in the BMNH collection from Utah and Idaho (leg. G.F. Knowlton) have more numerous rhinaria distributed over ANT III, IV and V, indicating that they must be another species in the frangulae group. Monoecious holocyclic in Germany according to Börner (1952). Mier Durante & Nieto Nafría (1978) redescribed this species on the basis of Spanish material. Biological studies are needed to verify the taxonomic relationships of this and other frangulae-like species on Lamiaceae.

Aphis nerii Boyer de Fonscolombe Plate 6h Apterae are bright lemon yellow with mainly dark antennae and legs, and black siphunculi and cauda (see aphids of Karnataka website); BL 1.5-2.6 mm. Alatae have dark wing veins and secondary rhinaria distributed III 3-19, IV 0-8. In large dense colonies on growing shoots and along leaf midribs of Nerium oleander and other Apocynaceae. Although a specialised feeder on Apocynaceae, it is also found on many plants in other families. Distributed world-wide especially in tropical and subtropical regions including many Pacific islands, and sometimes forming large colonies in summer on ornamental Apocynaceae such as Dregea sinensis in gardens in temperate regions. Anholocyclic almost everywhere; sexual morphs are produced in Japan, but the life cycle is uncertain (Takada & Miyazaki 1993). There have been detailed studies of its ecology (Groeters 1989, 1993) and (lack of) genetic variation (Harrison & Mondor 2011) in USA. The aposematic coloration may be linked with sequestration of cardiac glycosides from its host plants (Malcolm 1990). For an illustrated online account see http://entnemdept.ufl.edu/creatures/orn/shrubs/oleander_aphid.htm. 2n=8.

Aphis nevskyana Kadyrbekov ( = A. taraxacicola (Nevsky)) Apterae have pale yellow body and appendages, and short siphunculi (only about 0.1 × BL, just a little longer than cauda); BL c.1.7-1.8 mm. Alatae have a pale yellow abdomen with brown pigment medially on tergites 1-6, and only c.2 secondary rhinaria on ANT III. This species is described both from undersides of leaves of Aesculus hippocastanum in June (presumably vagrants), and from inside leaf rosettes of Taraxacum spp. in June and August, in central Asia (Nevsky 1951, as Cerosipha taraxacicola), and probably also in South Korea (see below). The biology and life cycle are unknown. [Nevsky’s name is a homonym of A. taraxacicola (Börner) and was synonymised with that species by Remaudière & Remaudière (1997). However, Börner’s species differs from Nevsky’s description in its mottled dark green colour and longer siphunculi, and alatae have many more secondary rhinaria on ANT III than Nevsky’s species. It was therefore given the replacement name A. nevskyana by Kadyrbekov (2019b). An aphid from Taraxacum officinale in South Korea is described under the name taraxacicola Börner by S. Lee et al. (2002c), but is apparently Aphis nevskyana.]

Aphis nevskyi (Ivanoskaya) Colour of apterae in life unknown, with black markings laterally and on posterior abdominal segments, and red eyes; BL c.1.4 mm. On Euphorbia sp. in Azerbaijan.

Aphis newtoni Theobald Apterae are dark green to greenish brown or black, with black siphunculi and cauda, immatures often marked with white wax (see influentialpoints.com/Gallery); BL 1.7-2.4 mm. Alatae have secondary rhinaria distributed III 9-20, IV 3-10, V 0-3. Low down on leaf blades, and later on flower stalks and in inflorescences, of Iris spp, often on plants growing in damp places or in water. In Europe (not Scandinavia, but in southern Finland), and also recorded from Kazakhstan, Korea and Mongolia. Accessible colonies are usually ant-attended. Monoecious holocyclic with apterous males. 2n=8.

Aphis nigra Wilson Apterae are greyish-black with yellowish antennal flagellum and tibiae; BL c.0.8 mm. Collected on Cyrilla racemiflora at Batesburg, South Carolina, USA in 1910, and not since recorded (Wilson 1911).

Aphis nigratibialis Robinson Apterae are blackish in colour with tibiae also wholly black; BL 1.3-2.1 mm. On Cornus stolonifera in Idaho, Oregon, Utah, Washington and Wyoming, USA. Life cycle is unknown (Robinson & Chen 1969). A member of the asclepiadis group (Lagos et al. 2014).

Aphis nivalis (Hille Ris Lambers) Apterae are of unknown colour; BL c.1.8-2.0 mm. On Epilobium sp. in Greenland.

Aphis nonveilleri Petrovic & Remaudière Apterae are very pale, finely wax-powdered; BL 0.9-1.3 mm. On roots of Asperula cynanchica, in ant shelters. Described from Montenegro, and subsequently reported from Galium verum and Galium spp. in Kazakhstan (Kadyrbekov 2017a), and from Cruciata taurica in Turkey (Şenol et al. 2017). Monoecious holocyclic with oviparae in mid-late August (Petrovic & Remaudière 2002).

Aphis nudicauda Danielsson Apterae are pale green; BL 1.1-1.8 mm. On roots of Filipendula vulgaris in Sweden, and subsequently recorded from F. ulmaria in Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic; the fundatrix is described (original description), but alatae vivip. and sexuales are unknown.

Aphis ochropus Koch Apterae are pale yellow, yellowish green or greenish yellow (see influentialpoints.com/Gallery); BL 1.2-2.0 mm. Alatae have secondary rhinaria distributed III 10-11, IV 2-3. In ant-attended colonies on basal and subterranean parts of Dipsacus sylvestris (= fullonum) in central and eastern Europe, and recently found in Wales (Baker 2009a). This species was frequently regarded as a synonym of A. confusa Walker until Müller (1987a) showed that it could not be transferred to Knautia arvensis, and listed morphological differences for most morphs. Monoecious holocyclic, with oviparae and apterous males in September.

Aphis odinae (van der Goot) Plate 8a Apterae are usually grey-brown to reddish-brown (see influentialpoints.com/Gallery), but in east Asia a dark green form also occurs; BL 1.3-2.4 mm. They feed on undersides of leaves of host plants along main veins and in dense colonies on young shoots, attended by ants. This aphid is rather polyphagous, occurring on numerous shrubs and some trees in east and south-east Asia, especially in the families Anacardiaceae (Anacardium, Mangifera, Rhus), Araliaceae (Aralia, Polyscias, Kalopanax), Caprifoliaceae (Viburnum), Ericaceae (Rhododendron), Pittosporaceae (Pittosporum), Rubiaceae (Coffea, Mussaenda), and Rutaceae (Citrus). It has become widespread in Africa south of the Sahara (Barbagallo & Alcantara Santos 1989, Martin 1989, as Toxoptera; see also Blackman & Eastop 2000, pp. 355-6, as Toxoptera), and is also in Greece (Margaritopoulos et al, 2013), Turkey (Akyürek et al. 2019) and Hawaii. Apparently anholocyclic almost everywhere. In Japan, however, there is at least a partial sexual phase, with alate males and dark purple-brown oviparae produced on various plants (Kalopanax, Rhus, Viburnum) in October-November, and very dark brown to bluish-black fundatrices collected on Rhus and Juglans in April (Blackman et al. 2011). Togashi (1987) studied relationships with parasitoids, predators and ants. Maruthadurai (2019) studied the predators of A. odinae on Anacardium in India. Pergandeida kalopanacis Hori 1927 is a synonym.

Aphis odorikonis Matsumura Apterae are dark green; BL 1.8-2.2 mm. Alatae have 6-13 secondary rhinaria on ANT III. On Lamium album var. barbatum and Scutellaria indica in Japan (Takahashi 1966, Moritsu 1983), and also recorded from Ixeridum dentatum in South Korea (Lee et al. 2002c), so the host relationship requires clarification..

Aphis (Bursaphis) oenotherae Oestlund (= ribigillettei Allen & Knowlton; ?= neomexicana Cockerell & Cockerell) Apterae are pale yellowish green to dark green with pale siphunculi; BL 1.5-2.0 mm. Alatae have secondary rhinaria distributed III 9-16, IV 3-8, V 0-3. Apparently host-alternating in North America between Ribes spp. (aureum, alpinum, nigrum), and various Onagraceae (Epilobium, Oenothera), although this needs to be confirmed experimentally. It is also reported from Korea (Kim et al. 2006b) and Japan (Lagos et al. 2014), and is now established in all mainland states of Australia, possibly to the detriment of indigenous Epilobium-feeding species (Hales et al. 2014). In New Zealand there is a bright green aphid with very pale siphunculi curling the tips of shoots of Epilobium ciliatum, similar to A. oenotherae but distinguished by the characters given in the key. Introduced anholocyclic populations identified as A. oenotherae also occur on Onagraceae in Europe (Müller 1974). In Spain, large colonies have been found on the introduced composite plant Baccharis halimifolia (Nieto Nafría et al. 2005a). Populations in northern Europe, however, have now been distinguished as another species, monoecious holocyclic on Oenothera (A. holoenotherae, q.v.), and the identity of anholocyclic populations in the south and west of Europe (i.e., how many different forms have been introduced from North America) is therefore uncertain. A. oenotherae is closely related to the European species A. grossulariae, which has a very similar biology. 2n=8.

Aphis oestlundi Gillette Apterae are pale green with blackish apices to antennae, legs and siphunculi; BL c.1.8-2 mm. On leaves and stems of Oenothera biennis in North America. Monoecious holocyclic, with apterous males (Hottes & Frison 1931). 2n=8.

Aphis ogilviei Theobald Apterae are yellowish brown or tan, with black head, eyes, siphunculi and cauda; BL 1.8-2.3 mm. Alatae have secondary rhinaria distributed III 6-10, IV 0-2. On Lilium longiflorum (var. eximium) in Bermuda.

Aphis (Anthemidaphis) oligommata Tashev (Fig.5k) Apterae are wax-dusted, colour unknown; BL c.1.5 mm. Alatae are undescribed. On roots of Anthemis tinctoria in southern Bulgaria, and it is also recorded from France, Spain (Nieto Nafría et al. 2005a), Slovakia, Russia (Ural region) and Kazakhstan (Kadyrbekov 2017a). Apterae and oviparae assigned to this species have also been collected on subterranean parts of Achillea spp. in Poland and Sicily. Barbagallo & Stroyan (1982) provided a morphometric comparison with A. ligusticae.

Aphis ononidis Schouteden (= kaltenbachi Hille Ris Lambers) Apterae are grey due to heavy coat of wax meal; BL 1.2-1.7 mm. Alatae have 3-8 secondary rhinaria on ANT III. On Genista anglica, in small compact colonies on young shoots, not visited by ants, and also on Ononis spp. (Hille Ris Lambers 1956a, as A. schoutedeni). In continental Europe (Netherlands, Belgium, France, Germany, Spain). Monoecious holocyclic, with oviparae and apterous males in early October.

Aphis orchidis Bozhko Apterae are dull black; BL c.2.5 mm. On Orchis purpurea in Ukraine.

Aphis (Zyxaphis) oregonensis Wilson Apterae are dark greyish brown tinged with wine red; BL c.1.7 mm. On leaves and young twigs of Artemisia ludoviciana (var. gnaphalodes) and Seriphidium tridentatum in western USA. There is some confusion in the literature with A. canae and A. hermistonii, but the three species seem separable by the key characters given.

Aphis origani Passerini Apterae are dark mottled green (see influentialpoints.com/Gallery); BL 1.0-1.8 mm. Alatae have 2-8 secondary rhinaria on ANT III. On Origanum spp., forming ant-attended colonies on stems and in young leaves and inflorescences, and causing strong leaf-curl. There are records from other Lamiaceae, but these could be due to confusion with related species. In Europe (not Scandinavia), North Africa (Morocco), Central Asia and Pakistan. Recorded as monoecious holocyclic, although the sexual morphs do not seem to have been described.

Aphis ornata (Gillette & Palmer) (Fig.21l) Apterae are pale pinkish, with darker head and pronotum, and with variably-developed dark markings or bands on dorsal abdomen; BL 1.4-1.8 mm. Alatae have secondary rhinaria distributed III 7-10, IV 1-2. On leaves and stems of Chrysothamnus spp. and Ratibida columnaris in Colorado, USA (eastern slopes of Rockies) The preferred host may be Ch. nauseosus var. graveolens). Oviparae and apterous males occur in October (Palmer 1952). Closely related to A. gregalis, and there seem to be no reliable distinguishing characters for slide-mounted specimens.

Aphis orobanches Passerini Apterae are dark green; BL unknown. On root and at base of stem of Orobanche spp. in Italy. Described from O. ramosa near Bologna, and recorded on O. mutelii in Calabria by Macchiati (1883). Wilson & Vickery (1918) also list records from O. lutea and Lathraea squamaria (Orobanchaceae), but the species is regarded as a nomen dubium.

Aphis orocantabrica García Prieto & Nieto Nafría Apterae are yellow, greenish yellow or dark green with black siphunculi; BL 1.4-2.6 mm. Alatae have secondary rhinaria distributed III 7-14, IV 1-6, V 0(-1). In dense, ant-attended colonies at base of stem and on basal leaves of Gentiana lutea, at 1,300-1,900m in the Cantabrian Mountains (northern Spain). Monoecious holocyclic; fundatrix, oviparae and apterous males were included in the original description (García Prieto & Nieto Nafría 2005). Apterae and oviparae found on the same host in the French Alps (BMNH collection, leg. G. Remaudière) may also be this species but have a longer antennal terminal process (ANT PT/BASE of apterae 1.9-2.2 versus 1.1-1.8 for the Spanish population).

Aphis ortegoi Mier Durante, García-Gómez & Nieto Nafría Apterae are emerad to dark green with dark pleural spots, a reddish posterior abdominal cross-band and partially dark appendages; BL 1.5-2.3 mm. Alatae have 13-21 secondary rhinaria on III and (0-)1-3 on IV. On Glyzyrrhiza astragalina, causing deformation of stems and leaves. In Argentina (Chubut, Mendoza, Neuquén and San Juan provinces; Mier Durante et al. 2021). The life cycle is unstudied, but presumably monoecious and holocyclic.

Aphis oxytropiradicis Pashchenko Apterae are green; BL c. 1.0 mm. Alatae are undescribed. On an Oxytropis sp., living on underground part of stem near root. In east Siberia.

Aphis oxytropis Pashchenko Apterae are brown, wax-dusted; BL c.2.1-2.2 mm. Alatae are undescribed. In ant-attended colonies on flower stems and lower parts of flowers of Oxytropis oxyphylla. In east Siberia. It appears closely related to A. astragali.

Aphis paludicola Hille Ris Lambers (= Pergandeida palustris Börner) Colour of apterae in life unknown, probably dark and wax-dusted; BL 2.3-2.7 mm. Alatae have secondary rhinaria distributed III 9-15, IV 0-7, V 0-4. On Euphorbia palustris in Germany and Ukraine. Tashev (1966) provided a redescription based on some of Börner’s specimens.

Aphis panzeriae Holman Apterae are dark green, bluish green or dull grass-green; BL 1.5-2.1 mm. On root collars of Panzeria (= Panzerina) lanata in Mongolia.

Aphis papillosa Mier Durante, Nieto Nafría & Ortego Apteraea are dark brown to black, rather shiny (imm. matt); BL 1.0-1.9 mm. Alatae have secondary rhinaria distributed III (1-) 4-14, IV 0-3(-5). On Senecio spp., living in dense colonies on stems, sometimes close to the root. Argentina and Chile. Monoecious holocyclic with apterous males (Mier Durante et al. 2003).

Aphis paradoxa (Mier Durante, Ortego & Nieto Nafría) Apterae are pale green to yellowish green powdered with white wax; BL 1.1-2.0 mm. On Senecio subumbellatus, and perhaps on another Andean Senecio sp., forming small dense colonies on aerial parts, especially flower stems. In Argentina. Monoecious holocyclic, but males are undescribed.

Aphis paravanoi Nieto Nafría, Ortego & Mier Durante Apterae are orange or dark green with yellowish siphunculi and cauda; BL 0.95-1.5 mm. Alatae have secondary rhinaria distributed III 2-6, IV 0-1. On stems and leaves of Mulinum spinosum in Argentina. Monoecious holocyclic, with oviparae and apterous males in March (Nieto Nafría et al. 1999b).

Aphis parietariae Theobald Apterae vary from pale to dark bluish green, sometimes with a waxy bloom (see influentialpoints.com/Gallery), or as pale yellowish summer dwarfs; BL 0.9-1.7 mm. Alatae have secondary rhinaria distributed III 4-8, IV 0-2, V 0. In dense colonies on stems, under leaves and on inflorescences of Parietaria spp. Widely distributed in Europe, and in Azores, Canaries, Madeira, north Africa, Mediterranean and Middle East. Monoecious holocyclic, with apterous males (with secondary rhinaria distributed III 2-14, IV 7-20, V 1-12). 2n=8.

Aphis pashtshenkoae Remaudière Apterae are yellow-green, wax-dusted; BL c. 2 mm. Alatae are undescribed. In dense colonies on undersides of apical leaves of Lysimachia davurica (Pashchenko 1993, as A. lysimachiae). In East Siberia (Maritime Territory). Records in Holman (2009) from Ukraine and Kazakhstan should be referred to A. lysimachiae, and a record from Turkey (Akyürek et al. 2011) requires further confirmation.

Aphis passeriniana (Del Guercio) Apterae are mottled dull pale green to dark green, often with a waxy bloom (see influentialpoints.com/Gallery); BL 1.2-1.8 mm. Alatae have secondary rhinaria distributed III 3-12, IV 0-3, V 0-1. Up growing shoots, in curled leaves and among inflorescences of Salvia spp. Ant-attended. In Europe (UK, Germany, Poland, Italy, Spain, Italy, Switzerland, Bulgaria, Croatia, Serbia, Hungary, Greece), Turkey (Bardjadze 2017) and Israel. Presumably monoecious holocyclic, although sexual morphs have apparently not been described.

Aphis patagonica Blanchard Apterae are brick red, BL 1.5-2.3 mm. Alatae have 2-6 secondary rhinaria on ANT III. On Berberis spp. in Chile and Argentina. Ortego & Mier Durante (1997) redescribed the species, including oviparae collected in January. Apterous males occur in January (BMNH collection, and Nieto Nafría et al. 2004).

Aphis patriniae Takahashi Apterae are yellowish green to green; BL 1.0-1.4 mm. On Patrinia spp., colonising axils of stem leaves (Holman 1987). In east Siberia, Japan and Korea. There are records from Valeriana spp. in Korea (Lee et al. 2002c).

Aphis patrinicola Holman Apterae are pale green, BL 1.05-1.45 mm. On Patrinia scabiosifolia, colonising root collar and basal part of stem, sheltered by ants. In east Siberia, and also recorded from Kazakhstan on P. intermedia (Kadyrbekov 2014e).

Aphis patriniphila Holman Apterae are green or yellow; BL 1.25-1.63 mm. On Patrinia rupestris, probably living on terminal parts of plant, in Mongolia, and also recorded from Kazakhstan on P. intermedia (Kadyrbekov 2012a). This species seems close to A. spiraecola.

Aphis patvaliphaga Pashchenko Apterae are green or yellow-green, sometimes with the end of the abdomen dark brown; BL c. 1.4-1.5 mm. On leaves and stem of Patrinia spp. and Valeriana coreana (= fauriei). Described from east Siberia, and subsequently recorded from Kazakhstan (Kadyrbekov 2017a).

Aphis pavlovski Narzikulov Apterae are brown with black siphunculi; BL 1.7-1.9 mm. On roots of Hypericum scabrum in Tajikistan.

Aphis pawneepae Hottes Apterae are brown to reddish-brown with antennae and legs wholly dusky brown; BL 1.5-1.7 mm. Found by Hottes (1934) in shelters constructed by the ant Crematogaster lineolata at the bases of trunks of young Cercis canadensis trees. Parker (1935), however, found colonies arranged along undersides of twigs and branches (always on old wood). Fundatrices and oviparae have also been collected, so this species is monoecious holocyclic on Cercis. Recorded from Illinois, Kansas, Montana, North Carolina and Pennsylvania, USA.

Aphis pediculariphaga Pashchenko Apterae are dark dingy green, almost black; BL c.1.6-1.7 mm. On Pedicularis resupinata in east Siberia, colonising basal or upper part of stem, and undersides of leaves, attended by ants.

Aphis pentstemonicola Gillette & Palmer Apterae are pale green, with brownish head and dark green dorsal markings (see aphidtrek.org); BL 1.8-2.0 mm. On leaves and stems of Penstemon spp. in western USA (Utah, Idaho, Washington, Wyoming, Colorado), and also collected on P. virens in Alberta, Canada (BMNH collection, leg. A.M. Harper). Monoecious holocyclic, with oviparae and apterous males in October (Palmer 1952). This seems to be a very variable species; see footnote in Palmer (1952:161).

Aphis periplocophila Zhang Apterae are dark green, with black siphunculi: BL c. 2.0 mm. On Periploca sepium in China. A member of the frangulae/gossypii group.

Aphis pernilleae Heie Apterae are green or bluish green, siphunculi dark or pale with dark apices; BL 0.7-1.2 mm. In dense, ant-attended colonies on roots of Hypochaeris sp. (probably glabra), down to 21 cm below surface. In Denmark, and aphids on Andryala spp. in Spain and Italy have now been assigned to this species (Nieto Nafría et al. 2005a; Barbagallo & Massimino Cocuzza 2014). Monoecious holocyclic; oviparae occur in September (males are undescribed).

Aphis peucedani Szelegiewicz Apterae are dark green; BL 1.6-2.2 mm. Alatae have secondary rhinaria distributed III 5-6, IV 0-2. On Peucedanum officinale in Hungary, colonising young shoots and later in summer within leaf sheaths (original description).

Aphis (Debilisiphon) peucedanicarvifoliae (Bozhko) Apterae are dirty green, BL c. 2.0 mm. On Peucedanum carvifolia in Ukraine.

Aphis phaceliae Gillette & Palmer Apterae are yellowish green to canary-yellow mottled with bluish green, with rusty yellow head, black siphunculi and pale cauda; BL c. 1.2-1.7 mm. Alatae have secondary rhinaria distributed III 7-12, IV 1-4. On leaves and stems of Phacelia sp. in Colorado, USA (Palmer 1952), with subsequent records from Idaho, Washington and Oregon, (including collections from Ph. nemoralis and Ph. heterophylla; aphidtrek.org).

Aphis philadelphicola Pashchenko Apterae are green, with blackish siphunculi and cauda; BL 1.6-2.1 mm. Alatae have secondary rhinaria distributed III 5-9, IV 0-2. On Philadelphus spp. in east Siberia, and also recorded from Kazakhstan (Kadyrbekov & Karimova 2011).

Aphis phlojodicarpi Pashchenko Apterae are dingy blackish green, wax-powdered; BL c. 1.9 mm. On Phlojodicarpus spp. in ant-attended colonies on stems, in inflorescences and in upper leaf sheaths. In east Siberia. Monoecious holocyclic (according to original description).

Aphis (Pseudoprotaphis) picridicola Holman (= Protaphis striata Hille Ris Lambers, = Protaphis funicularis Müller) Apterae are bluish grey, bluish green, pale green or dark grey-green, with distinctive black dorsal markings, often powdered with bluish grey wax (see influentialpoints.com/Gallery); BL 1.4-2.1 mm. Apterae have secondary rhinaria distributed III 0-11, IV 1-8, V 0-5, and alatae have them distributed III 9-15, IV 1-6, V1-4. On Hypochaeris spp., Leontodon spp. and Picris hieracioides, living basally on root collar and lower leaves, attended by ants. In Europe and west Siberia. Monoecious holocyclic, with apterous males (original description and Müller 1968a, as Protaphis funicularis).

Aphis picridis (Börner) (= stroyani Szelegiewicz) Apterae are bright yellow with black siphunculi (see influentialpoints.com/Gallery); BL 1.1-1.8 mm. Alatae have secondary rhinaria distributed III 3-6, IV 0(-1). On rosette leaves and root collar of Picris spp. and Reichardia picroides. In south, central and eastern Europe, Iran (Rezwani et al. 1994), and there is now also a record from Algeria (Laamari et al. 2013). Monoecious holocyclic (Börner 1950, as Doralina picridis). Patti & Barbagallo (1997) provided a detailed redescription of apterae and alatae.

Aphis pilosellae (Börner) Apterae are pale green to dark green, mottled, with black siphunculi and cauda (see influentialpoints.com/Gallery); BL 0.9-1.7 mm. Alatae have 7-14 secondary rhinaria on ANT III. On Hieracium spp., mostly of the Pilosella group, colonising undersides of basal leaves and subterranean runners. Ant-attended. In Europe, eastward to Russia and Ukraine (Holman 1998).

Aphis pilosicauda Gillette & Palmer Apterae are olive green, lightly wax-powdered; BL 1.5-1.9 mm. On stems of Cirsium sp. near ground, or on roots, in Colorado, USA. Oviparae and apterous males occur in October (Palmer 1952).

Aphis plantaginis Goeze Apterae are dark green, mottled with paler green, immatures paler, sometimes with a covering of particulate wax (see influentialpoints.com/Gallery); BL 1.2-2.2 mm. Alatae have secondary rhinaria distributed III 3-10, IV 0(-1). Under leaves and on leaf bases and subterranean stems of Plantago spp. In Europe, Central Asia, west and east Siberia, Mongolia, and also recorded from USA (New York), but that record is unsubstantiated (Foottit et al. 2006). Monoecious holocyclic with apterous males (Heie, 1986). [A. plantaginis ssp. asiatica Daniyarova, described from Taraxacum in Tajikistan, is a synonym of A. taraxacicola (Börner) (Stekolshchikov 2014a).]

Aphis platylobii Carver & White Apterae are orangey-brown appearing greyish due to dusting of white wax; BL 1.1-1.6 mm. Alatae have only 0-3 secondary rhinaria on ANT III. On young shoots and leaves of Platylobium formosum. Not attended by ants. In New South Wales, Australia. 2n=8*.

Aphis pleurospermi Pashchenko Apterae are dark green to dark brownish green; BL c.2.2 mm. On stems or inflorescences of Pleurospermum uralense in east Siberia. Monoecious holocyclic (original description; fundatrices in early June).

Aphis poacyni Zhang, Chen, Zhong & Li Colour of apterae in life unrecorded; BL c.1.4 mm. On Apocynum venetum in Xinjiang, China (Zhang, 1999). These specimens could be hot-weather dwarfs of A. apocynicola.

Aphis podagrariae Schrank Apterae are black or blackish-green in life (see influentialpoints.com/Gallery), BL 1.4-2.5 mm. Immatures often have pleural spots of wax as in A. fabae. Alatae have secondary rhinaria distributed III 12-18, IV 1-7, V 0-2. In ant-attended colonies causing tightly bunched leaf-curl on young foliage of Aegopodium podagraria. Later it may occur on other parts of the plant, but does not live preferentially in the inflorescences, as do summer generations of A. fabae (s. lat.) on this host. Also recorded from Seseli libanotis (Ossinnilsson, 1959) and S. (Libanotis) condensata (Kadyrbekov 2013d). Monoecious holocyclic on Aegopodium, with alate or occasionally brachypterous males. Throughout Europe (except Iberian peninsula), and eastward to Kazakhstan. Difficulties of distinguishing preserved specimens of this species from members of the fabae complex that colonise Aegopodium are discussed by Stroyan (1984).

Aphis polaris Stekolshchikov & Khruleva Apterae are bluish-black; BL 1.6-2.1 mm. Alatae have secondary rhinaria distributed III 8-10, IV 1-3, V 0(-1). On Astragalus alpinus and Oxytropis sp. on Wrangel Island, in arctic east Siberia, and only known from this location, where it is common and widespread (Stekolshchikov & Khuleva 2014). The life cycle is abbreviated, with fundatrices throughout July and apterae, alatae and an ovipara all collected on July 30^th.

Aphis polemoniradicis Pashchenko Apterae are dark green, slightly pruinose; BL c.1.5 mm. Apterae have 0-2 secondary rhinaria on ANT III, alatae have them distributed III 4-5, IV 0-1, V 0-1. On roots or leaf bases of Polemonium racemosum. Ant-attended. In east Siberia.

Aphis polii Barjadze, Blackman & Özdemir Apterae are yellowish to dark green with dark siphunculi and cauda; BL 0.7-1.4 mm. In ant-attended colonies on apical parts of shoots and sometimes on undersides of leaves of Teucrium polium. Recorded from France, Corsica, Italy, Czech Republic, Lebanon, Iran and Ukraine. Monoecious holocyclic with oviparae collected in France in early December, and apterous males (Barjadze et al. 2015).

Aphis pollinaria (Börner) Apterae are reddish brown with a dorsal wax pattern; BL 1.3-1.8 mm. Alatae have secondary rhinaria distributed III 6-14, IV 0-4, V 0(-2). On apical parts of Epilobium spp. in central and eastern Europe (Germany, Czech Republic, Slovakia). It often forms mixed colonies with A. epilobii, and sometimes also with A. spiraephaga (Holman 1990). No key characters could be found to distinguish Aphis onagraphaga Pashchenko, described from Epilobium spp. in east Siberia, from A. pollinaria, and it seems likely to be a synonym.

Aphis polygonacea Matsumura Apterae are dark green; BL 1.7-2.0 mm. Alatae have secondary rhinaria distributed III 7-9 (-16?), IV 0-2 (-7?). On Fallopia sachalinense, Polygonum sp. and probably other Polygonaceae, in Japan and Kuril Islands (Pashchenko 1988a). Colonies are ant-attended and distort leaves (BMNH collection data). Some alatae found on Polygonum hydropiper in Queensland, Australia (Eastop 1966, p.483; as Aphis sp.) could possibly be this species. A record from Rumex acetosella in India (Chowduri et al. 1969) needs confirmation.

Aphis polygonata Nevsky (= avicularis Hille Ris Lambers) Apterae are brown to black with short pale siphunculi (see influentialpoints.com/Gallery); BL 1.6-2.2 mm. Alatae have secondary rhinaria distributed III (4-)12-17, IV (0-)1-8, V 0-1. On creeping mats of Polygonum spp. and Persicaria maculosa, attended by ants. In Europe, North Africa, Middle East, Central Asia, Pakistan (Naumann-Etienne & Remaudière 1995), and also in USA (California, Illinois). Monoecious holocyclic with alate males (which are described by Stekolshchikov et al. 2008c). 2n=8*.

Aphis pomi De Geer Apterae are bright apple-green with black siphunculi and dark cauda (see influentialpoints.com/Gallery); BL 1.3-2.2 mm. Alatae have secondary rhinaria distributed III 6-12, IV 0-7. In dense colonies on young growth of several genera of woody Rosaceae including Chaenomeles, Cydonia, Malus and Pyracantha; ant-attended, and causing slight leaf-curl (Blackman & Eastop 2000, p.236). Later generations occur on the undersides of leaves, and are yellow-green. Monoecious holocyclic, with apterous males. In Europe, north Africa, Asia eastwards to India and Pakistan, and North America (where there is some confusion in the literature with A. spiraecola on the same hosts). Alatae have darker wing veins than A. spiraecola; Halbert & Voegtlin (1992) gave some other discriminants based on North American populations of the two species. Hogmire et al. (1992) compared population development and insecticidal susceptibility of A. pomi and A. spiraecola, Foottit et al. (2009b) and Rakauskas et al. (2015a) respectively compared morphological and DNA differences between the two species in North American and Eurasian populations. Naaum et al. (2012) developed a technique for DNA barcoding them under field conditions. Records of A. pomi from east Asia all seem to apply to other species, although it is surprising that it has not spread world-wide. Cottier (1953) in New Zealand had A. spiraecola. Lushai et al. (2004) studied genetic variation in A. pomi populations. Jaśkiewicz & Kot (2007) studied its population dynamics on Cotoneaster divaricatus in Poland. The male genitalia were described and illustrated by Wieczorek et al. (2011). 2n=8.

Aphis ponomarenkoi Holman Apterae are green with brown head and appendages; BL 1.3-1.6 mm. On Eleutherococcus senticosus in east Siberia.

Aphis popovi Mordvilko Apterae (fundatrices) are green, BL 1.7-2.0 mm. On Ribes sp. in east Russia (Yakutia). A member of the A. grossulariae group. Rakauskas (1996) re-examined Mordvilko’s specimens, which are fundatrices and their immature progeny, designated lectotypes, and provided a key to fundatrices of palaearctic Aphis on Ribes.

Aphis potentillae Nevsky Apterae are very dark green or black; BL c.1.7 mm. On aerial parts of Potentilla spp. in central and eastern Asia (Pashchenko 1988a, Rezwani et al. 1994), Lee et al. 2002c). A record from Algeria (Laamari et al. 2013) requires additional confirmation.

Aphis praeterita Walker (= diphaga Walker; epilobiina Walker) Apterae are yellow or greenish yellow, sometimes mottled with green, with siphunculi usually dark and cauda pale (see influentialpoints.com/Gallery); BL 1.2-1.9 mm. Alatae have secondary rhinaria distributed III 7-12, IV 0-5, V 0-1. On undersides of leaves and at shoot apices of Epilobium hirsutum, and sometimes other Epilobium spp., but apparently usually not on E. angustifolium; Walker’s original description from this host was apparently in error (Stroyan 1984). Monoecious holocyclic, with alate males. Not usually ant-attended. Rakauskas (2008) found an Oenothera sp. colonised by A. praeterita including sexual morphs and eggs in autumn in the Czech Republic, but fundatrices hatching the following spring did not complete their development when reared on Oe. biennis. Widely distributed in Europe, also reported from west Siberia (Ivanoskaya 1977), Iran (Rezwani et al. 1994), Kazakhstan (Kadyrbekov & Aoitzhanova 2005), Pakistan (Naumann-Etienne & Remaudière 1995) and China (Jiang et al. 2011b), and introduced to North America (Foottit et al. 2006).

Aphis proffti (Börner) Apterae are pale light green or yellowish green with pale legs and antennae; BL 0.7-1.5 mm. Alatae have secondary rhinaria distributed III 12-18, IV 4-9, V 3-5. On undersides of leaves and in influorescences of Agrimonia spp. (eupatoria, odorata) in Europe (Denmark, Sweden, France, Spain, Italy, Germany, Austria, Poland, Hungary, Czech Republic, former Yugoslavia), and also recorded from northern Turkey (Akyürek et al. 2011). Biology is unstudied, but an oviparae was collected on A. eupatoria in Sweden in October (BMNH , coll. F. Ossiannilsson). Closely related to the east Asian species A. argrimoniae.

Aphis propinqua Holman Colour of apterae in life unknown, probably blackish dusted with whitish wax powder; BL 1.9-2.3 mm. Alatae have secondary rhinaria distributed III 12-19, IV 0-4. On Euphorbia sp., collected from terminal parts and inflorescences, in Iran (original description).

Aphis psammophila Szelegiewicz Apterae are green to dark green with brownish head, dark siphunculi (sometimes pale at base) and a pale/dusky cauda; BL 1.0-1.5 mm. Alatae have secondary rhinaria distributed III 3-8, IV 0-2. On Campanula rotundifolia and Jasione montana, living on subterranean parts of the plant in ants’ nests, in dunes and other sandy areas (Heie 1986). In Europe (Netherlands, Denmark, France, Spain, Germany, Poland, Lithuania, Czech Republic, Bulgaria, Serbia), and also reported to occur in Kazakhstan on Codonopsis clematidea (Kadyrbekov 2014d). Monoecious holocyclic, with apterous males.

Aphis pseudeuphorbiae Hille Ris Lambers Appearance of apterae in life unknown, probably wax-dusted: BL 1.4-1.8 mm. On Euphorbia lanata in Israel, and also recorded from E. macroclada in Jordan. Closely related to A. euphorbiae and A. tirucallis, and possibly an unpigmented, hot-weather form of the latter species.

Aphis pseudocomosa Stroyan Apterae are dark warm brown with shiny black dorsum, imm. reddish brown to magenta with waxy bloom; BL 1.5-2.3 mm. Alatae have secondary rhinaria distributed III 4-11, IV 0-2. In small ant-attended colonies up shoots and in inflorescences of Lathyrus pratensis (Stroyan 1984), and also since found on L. tuberosus Onobrychis viciifolia and . In Europe (UK, Denmark, Sweden, Finland, Germany, Austria, Switzerland, Italy, Poland, Czech Republic, Spain, Turkey), west Siberia and Kazakhstan. Monoecious holocyclic with apterous males.

Aphis pseudocytisorum Hille Ris Lambers Apterae are black, densely powdered with pearly grey wax; BL 1.2-1.6 mm. Alatae are not recorded. Described from colonies on young shoots and petioles of Cytisus nigricans in Italy, and aphids identified as this species have subsequently been collected in Hungary and France on other Cytisus (and Chamaecytisus) spp., and in Italy on Spartium junceum. However, the features distinguishing pseudocytisorum from cytisorum (shorter siphunculi and less dorsal sclerotisation) vary greatly in this group and should be treated warily. Of the specimens in the BMNH collection, only those from Italy have siphunculi as short as the type material.

Aphis pseudopaludicola Tashev Colour of apterae in life unrecorded, probably dark and wax-dusted; BL 1.8-2.2 mm. Alatae have secondary rhinaria distributed III 12-17, IV 2-3. On stem, petioles and undersides of leaves of Euphorbia cyparissias in Bulgaria.

Aphis pseudopulchella Blanchard Colour of apterae in life unknown; BL c. 1.8 mm. Alatae have not been described. On Euphorbia portulacoides in Argentina. A record from Iran (Rezwani 2010) needs further confirmation.

Aphis pseudovalerianae Gillette & Palmer Apterae are pale olive-green with brown head, darker marginal areas, dark antennae and legs, and black siphunculi and cauda; BL 1.4-1.7 mm. Alatae have secondary rhinaria distributed III 14-20, IV 3-7, V 1-5. On roots of a Valeriana sp. in Colorado, USA. Monoecious holocyclic, with oviparae in September (Palmer 1952).

Aphis pulchella Hottes & Frison Apterae are bluish green and lightly wax-dusted in life; BL c.1.6-1.7 mm. Alatae have 5-7 secondary rhinaria on ANT III. On Euphorbia sp. in Illinois, USA, and there are also records from Montana, Iowa (on Eu. corollata) and North Carolina. Monoecious holocyclic; oviparae and alate males were collected in late September (Hottes & Frison 1931).

Aphis pulegii Del Guercio Apterae are pale yellow to dark green; BL 0.9-1.4 mm. Alatae have secondary rhinaria distributed III 4-14, IV 0-5, V 0-2. On Mentha pulegium, feeding on young shoots and undersides of leaves in spring, moving in summer to base of stem. Ant-attended. In Spain, Portugal, Sicily and Morocco. Ilharco (1987) redescribed this species and compared it with the closely related A. affinis.

Aphis pulsatillaephaga Pashchenko Apterae are green to dark green, BL of holotype 1.55 mm. On Pulsatilla spp. in east Siberia, living at base of stem. Alatae are unknown.

Aphis pulsatillicola Holman Apterae are dark green to blackish, in some specimens the dorsal abdomen is bright green; BL 1.4-2.2 mm. Alatae have 4-6 large secondary rhinaria on ANT III. Described from Pulsatilla pratensis ssp. nigricans, living on the root collar and stem base early in season, later up the stem and on leaves; ant-attended. In Czech Republic, and also recorded from P. grandis in Hungary (Basky 2014), and from P. patens in Kazakhstan (Kadyrbkov 2017a). Monoecious holocyclic, with apterous males (original description).

Aphis punicae Passerini Apterae are yellowish‑green, green or grey-green (see aphids of Karnataka website) with siphunculi pale green to brown with darker apices (but wholly dark in spring); BL 1.0-1.7 mm. Alatae have secondary rhinaria distributed III 4-11, IV 0-5. On upper sides of mature leaves of Punica granatum, concentrated along the mid‑ribs and around the leaf margins. Also recorded from Duranta plumieri (= erecta) and Lantana camara (Verbenaceae), Bignonia sp. and Campsis radicans (Bignoniaceae), and Plumbago capensis (Plumbaginaceae). In the Mediterranean region, Middle East, Ethiopia, India, Pakistan, and now established to occur in Japan, where A. punicae Shinji is a junior synonym (Sugimoto 2011a), and also in Korea (Lee et al. 2015). Monoecious holocyclic on Punica, with alate males, but in Israel there is also anholocyclic overwintering on Duranta (Swirski 1954b). Sugimoto (2011a) provided a redescription of apterae, alate, oviparae and males of Japanese populations, and compared it with A. gossypii, and Lee et al. (2015) provided morphometric and molecular (COI) comparisons with other Aphis species on pomegranate. See also Blackman & Eastop (2000). 2n = 8.

Aphis raji (Kumar & Burkhardt) Apterae are creamy white, BL c. 1.4-1.7 mm. Alatae have 6-10 secondary rhinaria on distal part of ANT III. Mostly on Lamiaceae (Callicarpa, Colebrookea, Salvia; BMNH collection, leg. S.P. Maity, A.N. Chowdhuri, R. Kumar), and there are also records from Cyathula tomentosa (Amaranthaceae; David et al. 1970, as A. leptorhyncha) and Buddleia paniculata (BMNH collection, leg. A.N. Chowdhuri). In northern India (Himachal Pradesh, Uttar Pradesh). A. paraverbasci Chakrabarti, described from an unidentified labiate in Himachal Pradesh and subsequently recorded from Buddleia sp. and Debregeasia sp. (A.K. Ghosh 2006), is probably a synonym. S.K. Das et al. (1981) described an ovipara under that name, collected in October, from an unidentified host. [Note: Information about chaetotaxy and relative length of R IV+V given in the original description of A. raji does not agree with paratypes in the BMNH collection; see also A.K. Ghosh (2006).]

Aphis ramona Swain Apterae are rich dark green, with head dusky brown; BL 1.1-1.2 mm. On Salvia stachyoides in California, and also in Utah, USA (BMNH collection and aphidtrek.org).

Aphis renjifoanae Ortego & Nieto Nafría Apterae are green with an orange transverse band at the level of the siphunculi; BL 1.2-1.4 mm. In rather compact colonies on fruits of Adesmia renjifoana in Mendoza province, Argentina (Nieto Nafría et al. 2016c). Other morphs and life cycle are unknown.

Aphis reticulata Wilson Apterae are brown, with a reticulate pattern over the entire body; BL c.1.7 mm. Alatae have c.40 protruberant secondary rhinaria on ANT III, but none on IV and V. Described from Seriphidium tridentatum in Oregon, USA, and apparently not found since 1915.

Aphis rhamnicola Lee, Lee & Kim (nec. Mamontova 1953) Apterae blackish green, green mottled with dark green, or (on secondary hosts) green lightly dusted with wax; BL 1.2-1.9 mm. On Rhamnus davurica in Korea, migrating to Commelina communis and various other secondary hosts (Y. Lee et al. 2014b). Oviparae and males were collected on Rhamnus in September to November. This aphid is primarily distinguished from A. gossypii by its DNA barcode (COI sequence), and by the usual presence of 1 to 4 marginal tubercles on ABD TERG 2-4, but its distribution and host relationships need to be clarified by further study, and more work is particularly necessary to show whether it is indeed distinct from other members of the frangulae group (q.v.).

Aphis rhamnifila David, Narayanan & Rajasingh Apterae are blackish-green, with brown legs and antennae, and darker siphunculi and cauda; BL 1.1-1.8 mm. A photomicrograph of a slide-mounted aptera is provided by García Prieto & Nieto Nafría (2007). Collected on undersides of leaves of Rhamnus virgatus in India (Uttar Pradesh, Himachal Pradesh, Garhwal Himalaya). Possibly this name is being applied to more than one species (David et al. 1971e).

Aphis rheicola Nevsky Aptere are yellowish orange or shining brown; BL c.2.3 mm. On undersides of leaves of Rheum maximoviczii in Ukraine, Kazakhstan and Uzbekistan (Kadyrbekov 2017a). [This species is tentatively keyed to the Aphis fabae group on the basis of Nevsky’s description, but not enough information is given to be certain of this. Hairs on antennae and body are said to be short and sparse.]

Aphis rhoicola Hille Ris Lambers Apterae are brown with pale antennae and legs (except black tarsi), and black siphunculi; BL 1.7-2.1 mm. On leaves and branches of Rhus abyssinica in Eritrea (Hille Ris Lambers 1954). Resembling A. gossypii, but with a consistently shorter antennal terminal process. Life cycle is unknown; however, very similar aphids occur on Rumex in northern India (David et al. 1971e, Raychaudhuri 1984, BMNH collection leg. D. Hille Ris Lambers), so this is a possible secondary host. The Indian specimens in the BMNH collection have a longer antennal terminal process (PT/BASE 1.9-2.2, compared with 1.5-1.9 for Eritrean types), shorter siphunculi (siphunculi 1.5-1.7 × cauda, compared with 1.8-2.1 ×) and a darker cauda, so may be another species in the difficult A. frangulae group.

Aphis ripariae Oestlund Apterae are pale yellowish green; BL not recorded. On undersides of leaves of Vitis riparia in Minnesota, USA.

Aphis roberti Nieto Nafría, Ortego & Mier Durante Apterae are shiny black; BL 0.9-1.7 mm. Alatae are dull black. In dense colonies on flowerheads of Mulinum spinosum in Patagonia (Argentina) and southern Chile (Nieto Nafría et al. 1999b). Oviparae and alate males occur in late March (Gonzáles Rodríguez et al. 2017)

Aphis roepkei Hille Ris Lambers Apterae are pale greenish yellow; BL 1.3-1.7 mm. Alatae have 3-6 secondary rhinaria on ANT III. In small ant-attended colonies on lower parts of stems and leaf petioles, and on roots, of Potentilla spp., especially reptans. In Europe (Sweden, Germany, northern Italy, Poland, Czech Republic, and recently found in Wales – Baker 2011); Iran, Kazakhstan (Kadyrbekov 2017a); and also recorded from P. chinensis in east Siberia (Pashchenko 1988a).

Aphis roripae (Palmer) Apterae are pale to mid green, with pale siphunculi and cauda; BL 1.3-1.8 mm. Apterae have 0-9 secondary rhinaria on ANT III, and alatae have 10-19 (antennae are 5-segmented). On stems and leaves of Rorippa hispida (= palustris) in western USA (Palmer 1952).

Aphis roumanica Holman Apterae are green, BL 1.4-1.9 mm; appearance of colonies in life is unrecorded but probably similar to A. farinosa, to which it is closely related. Monoecious holocyclic on Salix purpurea in Romania, with oviparae collected in July (Holman 1990). It has been found subsequently on several other Salix spp. (Holman 2009). Alate viviparae and males are undescribed.

Aphis rubiae Narzikulov Apterae are yellowish green, with black siphunculi and brown cauda; BL c.1.8 mm. Alatae have secondary rhinaria distributed III 7-9, IV 3-4. In large colonies on stems and undersides of leaves of Rubia tinctorum in Tajikistan.

Aphis rubicola Oestlund Apterae are pale yellowish green, with pale appendages (see influentialpoints.com/Gallery); BL 0.6-1.2 mm. Alatae have 3-10 secondary rhinaria on ANT III. On leaves and shoot apices of Rubus idaeus var. strigosus and R. occidentalis in North America. A record from Turkey (Akyürek et al. 2019) requires additional confirmation. Monoecious holocyclic, with apterous males. The minute long-haired summer dwarfs with 5-segmented antennae may often have been misidentified as A. rubifolii. See also Blackman & Eastop (2000). 2n=8.

Aphis rubicolens (Hori) Recorded from Rubus sp. in Japan (Hori 1929a, as “ichigo-gosetsu-aburamushi [= berry aphid with 5-segmented antenna], Cerosipha rubicolens Hori”), but the original description cannot be found.

Aphis rubifolii (Thomas) Apterae are pale greenish-yellow with pale appendages (see aphidtrek.org and influentialpoints.com/Gallery); BL 0.9-1.4 mm. Alatae have 2-7 secondary rhinaria on ANT III. On wild and cultivated Rubus spp., especially blackberries; the leaves are rolled and crinkled into pseudogalls. Records from R. idaeus var. strigosus may be confusing this species with summer dwarfs of A. rubicola. In North America, and also recorded from north-eastern India and Nepal (A.K. Ghosh & L. K. Ghosh 2006). Monoecious holocyclic with apterous males.

Aphis rubiradicis Robinson Apterae are grey, BL 1.2-1.4 mm. On roots of Rubus chamaemorus in Saskatchewan, Canada. Monoecious holocyclic, with oviparae and apterous males present in early August (original description).

Aphis ruborum (Börner & Schilder) Apterae are blue-green in spring, and blue-green or pale yellow-green in summer (see influentialpoints.com/Gallery); BL 0.8-2.0 mm. Alatae have rather large secondary rhinaria distributed III 4-10, IV 0-3. In dense, ant-attended spring colonies on young shoots of Rubus fruticosus (agg.), and later under leaves, in flowers and on developing fruits. Also sometimes on Fragaria × ananassa, and found in Chile on R. idaeus (BMNH collection, leg. R.H. Converse). In Europe, north Africa, south-west and central Asia eastward to India and Pakistan, introduced to South America (Chile, Argentina) and USA (Washington; Havelka et al. 2012). Its presence on blackberries from Mexico intercepted in Florida indicated a wider distribution in North America (Skvarla et al. 2017), and it is now reported to occur in Mississippi, where colonies were found to be parasitised by two Aphelinus spp. of Eurasian origin imported originally to western USA to control Diuraphis noxia (Riddick et al. 2019). In fact it is now known to be widely distributed in Mexico, where it is parasitised by Aphidius colemani (Peña-Martinez et al. (2020). Monoecious holocyclic with both apterous and alate males. Havelka et al. (2012) reviewed all the parasitoids of this species. Some of the records from India may apply to the longer-haired Aphis longisetosa (q.v.). Populations in the BMNH collection from several countries of former Yugoslavia on both R. fruticosus and R. caesius have a longer antennal terminal process and may be an undescribed species. 2n=8.

Aphis rukavishnikovi Ivanoskaya Apterae are bright green with dark siphunculi and cauda; BL 3.5-4.0 mm. Alatae have secondary rhinaria distributed III 8-10, IV 4-5, V 1-2. In small colonies on undersides of leaves of Caltha silvestris (= C. palustris). In northern Asia.

Aphis rumicis L. Apterae are dull black or very dark brown (see influentialpoints.com/Gallery); BL 1.4-2.8 mm. Alatae have secondary rhinaria distributed III 5-20, IV 0-4. On Rumex spp., feeding on undersides of leaves in spring and rolling them longitudinally into tubes. Later it occurs on stems and inflorescences. It can also occur on Rheum spp. (Stroyan 1984, Lee et al. 2002c), but records from other plants mostly apply to A. fabae. Ant-attended. Widely recorded in the northern hemisphere, although many North American and at least some Japanese records (e.g. Takahashi 1966) apply to different species. Monoecious holocyclic, with apterous males and oviparae with barely swollen hind tibiae. 2n=8.

Aphis rumicivora Heie Apterae are dark green to greenish black; BL 1.5-2.3 mm. On roots of Rumex acetosella in northern Europe (UK, Denmark, Sweden, Finland, Poland), in ants’ nests, especially in sandy soil and dune areas. Alatae have secondary rhinaria distributed III 4-8, IV 0. Monoecious holocyclic on R. acetosella. It was described as a subspecies of A. acetosae by Heie (1986), but given full species status by Stekolshchikov et al. (2007b), who recorded it from European Russia (Chuvashia), and described the ovipara, collected in late August. It is also now recorded from Kazakhstan (as acetosae ssp. rumicivora; Kadyrbekov 2016d).

Aphis salicariae Koch (= corniella Hille Ris Lambers) Apterae are reddish-brown with a light dusting of wax, pale legs (except tarsi), and black siphunculi and cauda (see influentialpoints.com/Gallery); BL 1.6-2.4 mm. Alatae have secondary rhinaria distributed III 12-28, IV 0-2. Causing leaf curl to Cornus spp. (alba, mas, sanguinea, stolonifera) in spring, and also feeding in developing flower umbels. Host-alternating to Epilobium angustifolium, where colonies are formed on undersides of leaves along midribs and are often attended by ants. It is also recorded from E. obscurum (Chumak 2004). Migration back to Cornus occurs in October. Of holarctic distribution; throughout Europe (except southern Spain and Portugal), across Siberia, Iran, Kazakhstan, and also widespread in North America. It is a member of the asclepiadis group (Lagos et al. 2014). 2n=8.

Aphis salsolae (Börner) Apterae are reddish brown, and unusually active when disturbed (J. Holman, pers. comm.); BL 1.4-1.9 mm. On certain genera of Amaranthaceae (Kochia, Salicaria, Salsola) in southern Europe, and the Mediterranean and Black Sea regions (France, Germany, Italy, Israel, Bulgaria, Romania, Turkey, Iran, Crimea, Georgia), and also now recorded from Poland (Osiadacz & Hałaj 2017). When mounted on slides this species is almost indistinguishable from A. craccivora.

Aphis salviae Walker Apterae are brown, shining, BL 1.5-2.1mm. Alatae have secondary rhinaria distributed III 4-10, IV 0-1. On Salvia spp, and also recorded from Lavandula multifida and Teucrium polium. Europe (France, Spain, Portugal, Switzerland, Poland, Hungary, Czech Republic, Bulgaria, Serbia), Algeria (Laamari et al. 2013), Israel, Turkey, Iran and Kazakhstan. Monoecious holocyclic with apterous males (Nieto Nafría et al. 1986).

Aphis sambuci L. Plate 7a Apterae are dark grey-green or brown to dark blue-green powdered with greyish wax (see influentialpoints.com/Gallery); BL 1.4-3.5 mm. In dense colonies around young stems of Sambucus spp. in spring. Alatae produced on Sambucus have secondary rhinaria distributed III 11-32, IV 0(-11), V 0(-4). Host alternation occurs to roots of plants in several genera, particularly Caryophyllaceae (Cerastium, Dianthus, Silene, Melandrium, Moehringia, Spergula), also often on Rumex, Capsella, Oenothera and Saxifraga, where they are attended and sheltered by ants. Alatae produced on secondary hosts have secondary rhinaria distributed III 30-44, IV 10-27, V 3-15. Colonies may persist through summer on Sambucus and may produce oviparae, but males only seem to be produced on secondary hosts. Anholocyclic overwintering on roots occurs regularly in Europe (Jacob 1949, Iglisch 1966), and probably elsewhere. Holarctic (except that long-haired populations in Japan and Korea are regarded as a different species, A. horii), in north Africa (according to Nieto Nafría et al. 2005a), and in North, Central and South America. It is often recorded in North American literature as A. sambucifoliae Fitch. Its population dynamics in elderberry orchards using different production systens was studied by Holb et al. (2010). 2n=8.

Aphis sanguisorbae Schrank Apterae are blackish brown, sometimes with copper and red-brown hues on the posterior abdomen (see influentialpoints.com/Gallery); BL 1.1-2.2 mm. Alatae have secondary rhinaria distributed III 3-9, IV 0-4, V 0-2. In ant shelters on basal parts of Sanguisorba spp. In Europe and the Middle East (Sweden, England, France, Spain, Germany, Finland, Poland, Bulgaria, Czech Republic, former Yugoslavia, Israel (BMNH collection, on S. spinosum), Turkey, Ukraine, Iran) and also now recorded from Kazakhstan (Kadyrbekov 2014e). Blackish green aphids on S. minor are regarded as a subspecies, A. sanguisorbae ssp. poterii (Börner), following Stroyan (1984). They tend to have shorter hairs and more weakly developed marginal tubercles (Holman 1987). Monoecious holocyclic with apterous males.

Aphis sanguisorbicola Takahashi Apterae are shining dark brown to black, with black siphunculi and whitish green cauda; BL 1.0-2.3 mm. On Sanguisorba spp. in Asia (east and west Siberia, Japan, Korea).

Aphis saniculae Williams Apterae are green (original description) or golden yellow with yellowish brown head (Hottes & Frison 1931, as A. luridis); BL c. 1.5 mm. Alatae have secondary rhinaria distributed III 21-50, IV 6-15, V 1-7. In ant-attended colonies on undersides of leaves of Sanicula canadensis, which may be curled downwards to form small “leaf-nests” (Williams 1911, Stroyan 1970); also recorded from leaves and flower stalks of Zizia spp., and from Cicuta maculata (Rojanavongse & Robinson 1977). In USA (Illinois, Iowa, Minnesota, Nebraska, Kansas) and Canada (Manitoba). It is a member of the asclepiadis group (Lagos et al. 2014), and very similar in morphology to A. thaspii inhabiting the same range of umbelliferous hosts, but apparently there are consistent colour differences in life which correlate with the presence/absence of secondary rhinaria in the apterous morph. Experimental work is needed to confirm whether or not these two species are really distinct.

Aphis sassceri Wilson Apterae are greenish-brown with dusky siphunculi, BL c.1 mm; alatae are predominantly orange-yellow. Apterae have distinct marginal abdominal tubercles. Known only from the original collection on Annona “rectilineata” (= ?reticulata) in Florida, USA (Wilson 1911).

Aphis saussurearadicis Pashchenko Apterae are brown, wax-powdered; BL c.1.6 mm. Alatae have 9-12 secondary rhinaria on ANT III. On Saussurea spp., living in small, dense colonies on radical leaves and roots, attended by ants. In east Siberia, Kazakhstan (Kadyrbekov 2017a), and aphids identified as this species are also recorded from Korea (Lee et al. 2002c; although the alata illustrated by Lee et al. has ANT III with at least 20 rhinaria). Monoecious holocyclic with oviparae in mid-August (original description). In Korea it is also recorded from Ligularia stenocephala.

Aphis schilderi (Börner) Apterae are dark green; BL 1.5-2.0 mm. Alatae have secondary rhinaria distributed III 4-7, IV 0-2. On stems, and later in summer on leaf sheaths, of Peucedanum spp. in Germany, Hungary and Poland (Szelegiewicz 1964b, 1966). Only recorded from that genus in Europe, but reported to occur on plants in other genera of Apiaceae (Angelica decurrens, Carum carvi, Cicuta virosa and Libanotis condensata0 in Kazakhstan (Kadyrbekov 2012a, 2017a). Attended by ants. Monoecious holocyclic according to Börner (1952).

Aphis schinifoliae Blanchard Apterae are blackish-brown in life with antennae and legs mainly dark, and cauda dark, but siphunculi pale, rather short and almost flangeless; BL 1.4-1.7 mm. The antenna has a short terminal process (ANT PT/BASE ratio about 1.4), and there are well-developed abdominal marginal tubercles (Blanchard 1939). Monoecious holocyclic on Schinus spp. (dependens, molle) in Argentina, Chile and Bolivia. Remaudière et al. (1992) provided a redescription including alatae, and also oviparae collected on Schinus molle in November. Ortego et al. (2007) recorded oviparae in December–February, provided further morphometric data and described the apterous males (collected in January).

Aphis schinivora Ortego, Nieto Nafría & Mier Durante Apterae are shining black with legs partially white; BL 0.9-1.4 mm. Monecious holocyclic on Schinus johnstonii in Argentina, forming very dense colonies on stems and leaves. The life cycle is apparently abbreviated, as oviparae and apterous males were collected in late January (Ortego et al. 2007).

Aphis (Toxoptera) schlingeri (Tao) Apterae are blackish-brown with banded antennae, almost indistinguishable from A. (T.) aurantii; BL 1.6-1.8 mm. Described from Ficus sp. in Hong Kong (Tao 1961). There are alatae in the BMNH collection from Ficus microcarpa (leg. D.S. Hill) agreeing with Tao’s description; also alatae with host unrecorded from China (Foochow; leg. M.S. Yang) and one alata trapped in Nepal (leg. K.C. Sharma). Its biology and life cycle are unknown. [This species is synonymised with A. (T.) aurantii by Remaudière & Remaudière (1997), and the apterae are very similar, but differences in the antennal sensoriation of alatae (see key to aphids on Ficus) indicate that they must be distinct species.]

Aphis (Bursaphis) schneideri (Börner) Apterae are dark green, dusted with bluish-grey wax (see influentialpoints.com/Gallery); BL 1.2-2.3 mm. Alatae have secondary rhinaria distributed III 9-20, IV 2-11, V 0-5. In spring colonies on young growth of Ribes spp., most typically R. nigrum, causing curling and crumpling of young leaves., and later on undersides of leaves. Ant-attended. Widely distributed in Europe, eastward to Turkey, Kazakhstan and west Siberia. Monoecious holocyclic with apterous males. Rakauskas (2003b) reported evidence of natural hybidisation with A. grossulariae.

Aphis schuhi Robinson Appearance in life is unknown; BL 1.3-1.6 mm. Only found once, on Holodiscus discolor in Oregon, USA (see Jensen 2021). Alatae are unknown.

Aphis sedi Kaltenbach (= Aphis tahosalea Hottes & Wehrle) Apterae are dark green to almost black (see influentialpoints.com/Gallery); BL 1.0-1.7 mm. Alatae have secondary rhinaria distributed III 4-12, IV 0-3. On young stems, in inflorescences and under leaves of Sedum spp. and other Crassulaceae. Often ant-attended. Throughout Europe, Iran, Turkey, Kazakhstan (Kadyrbekov 2009b), North Korea (Lee et al. 2002c), South Africa, and in North and South America, Australia and New Zealand (BMNH collection). Monoecious holocyclic with apterous males (Heie 1986, Kring & Kring 1991). The mitochondrial COI and COII sequence of samples identified as A. sedi in Italy was indistinguishable from that of A. gossypii (Cocuzza et al. 2009), although there are clear biological differences, and Lagos-Kutz et al. (2014) provided some distinguishing morphological features. 2n=8.

Aphis sediradicis Pashchenko Apterae are green or dark green, BL c. 1.2-1.3 mm. In small colonies on root and stem base of Sedum aizoon. In east Siberia.

Aphis selini (Börner) Apterae are pale or dark blue-green; BL 1.2-1.5 mm. Alatae have secondary rhinaria distributed III 7-10, IV 3-4, V 0-1. On prostrate stems/ stolons of Selinum carvifolia in Germany, and also in southern Finland (Albrecht 2010). It seems possible that A. umbelliferarum (Shaposhnikov) is a synonym.

Aphis sempervivae Mamontova-Solukha Apterae are dark green, wax-dusted; BL c.0.7 mm (?). On Sempervivum soboliferum in Ukraine.

Aphis seneciocrepiphaga Pashchenko Apterae are blackish green, slightly wax-dusted; BL c. 1.9 mm. Alatae have secondary rhinaria distributed III 26-33, IV 8-15, V 0-5. On Senecio spp. and Crepis tectorum, living on undersides of upper leaves and flower stems, attended by ants. In east Russia (Sakhalin, Kuril Islands, Yakutia).

Aphis senecionicoides Blanchard Apterae are black, rather shiny; BL 1.1-2.1. Alatae have secondary rhinaria distributed III 5-15 (-23), IV 0-7, V 0-3. On Senecio spp., forming dense, tranquil colonies on stems. In Argentina. Monoecious holocyclic with alate males (Mier Durante et al. 2003).

Aphis senecionis Williams Apterae are dark olive-green, with dusky to dark brown legs and dusky brown siphunculi; BL 1.7-1.9 mm. In large colonies on upper stems, leaves and flower-stalks of Senecio canus. In mid-western USA (Nebraska, New Mexico). Closely related to A. lugentis.

Aphis senecioradicis (Gillette & Palmer) Apterae are dusky blue-green with head brownish, and dark appendages; BL 2.2-2.8 mm. Apterae have secondary rhinaria distributed III 1-5, IV 1-5; alatae have them distributed III 5-8, IV 1-6. On roots of Senecio spartioides and Senecio sp. in western USA (Colorado, Idaho). Monoecious holocyclic with oviparae and apterous males in late October (Palmer 1952).

Aphis sensoriataeuphorbiae Bozhko Apterae are brown, wax-dusted; BL c.1.8-2.0 mm. Alatae have secondary rhinaria distributed (1 antenna only) III 19, IV 11, V 8. On Euphorbia lathyris in Ukraine.

Aphis septentrionalis Pashchenko Apterae are brown or black; BL c.1.5 mm. On Pedicularis resupinata and Pedicularis sp. in east Siberia, colonising inflorescences, and upper and basal parts of stem, often in dense colonies. Monoecious holocyclic, with oviparae and apterous males in late July (original description). Recently it has been reported to occur on a Crepis sp. in western Siberia (Altai Republic; Stekolshchikov & Novgorodova 2013).

Aphis serissae (Shinji) Apterae are pale green, BL 0.8-1.3 mm. Alatae have c.4 secondary rhinaria on ANT III. On Serissa japonica in Japan, and recorded on Artemisia princeps in Korea (Paik 1965).

Aphis serpylli Koch Apterae are dark green with a slight waxy bloom; BL 1.0-1.4 mm. Alatae have secondary rhinaria distributed III 4-9, IV 0-2, V 0-1. On shoots and inflorescences of Thymus spp., and certain other labiates (Micromeria, Thymbra). Attended and sometimes tented over by ants. In Europe, Middle East and Central Asia. Monoecious holocyclic with apterous males. In the BMNH collection there is an undescribed, closely related species collected on Hyssopus officinalis in Switzerland.

Aphis serratularadicis Pashchenko Apterae are pale green; BL c.1.2 mm. In small dense colonies on roots of Serratula wolfii in east Siberia.

Aphis (Debilisiphon) seselii (Bozhko) Apterae are green or dark green; BL c.1.6 mm. On lower part of stem of Seseli tortuosum in Ukraine. [This is probably a synonym of A. umbelliferarum; the main difference from A. umbelliferarum that Bozhko gives in her (1963) key is the lack of marginal tubercles, and we have followed this in our key, yet a specimen in the BMNH collection identified as seselii by Bozhko has well-developed marginal tubercles like those of umbelliferarum.]

Aphis shaposhnikovi Holman Apterae are yellowish brown, covered with a whitish grey powder; BL 1.75-2.0 mm. On Arctium tomentosum, feeding at bases of leaves close to ground, sheltered by ants (original description). Pashchenko (1997b) recorded it from petioles and stem bases of Synurus deltoides. In east Siberia (Maritime Territory).

Aphis sibirica Stekolshchikov & Novgorodova Apterae are grey, brown, brownish grey or brownish grey-green, powdered with wax; BL 1.3-2.3 mm. On species in several genera of Orobanchaceae (Euphrasia, Odontites, Pedicularis, Rhinanthus), in colonies on tops of stems or between flowers, attended by ants (Stekolshchikov & Novgorodova 2019). Smaller colonies were also found on some plants in other families (Elaeagnus, Medicago). Alatae have secondary rhinaria distributed III 9-19, IV 0-5. In northern Russia (western and eastern Siberia; provinces of Novosibirsk and Irkutsk).

Aphis sierra Essig Apterae are deep green, often with a bluish cast, with mainly dark antennae, and dark markings on posterior abdomen which may include a broad dark cross-band on abdominal tergite 6, bridging the large postsiphuncular sclerites; BL 1.5-1.8 mm. On Penstemon newberryi in Sierra Nevada, California, and also collected on P. fruticosus in British Columbia, Canada (BMNH collection, leg. R. Danielsson). Monoecious holocyclic, with oviparae and apterous males in the Sierra Nevada in August (original description). Very closely related to A. penstemicola; it needs to be confirmed whether the differences between these two warrant separate species status.

Aphis silaumi Bozhko Apterae are dirty green, wax-dusted, with dark siphunculi and cauda; BL c. 1.4 mm. On Silaum besseri in Ukraine.

Aphis silenephaga Pashchenko Apterae are dark dingy green, wax-dusted; BL c.1.6 mm. Apterae have secondary rhinaria distributed III-IV 0-2, V 0-1; alatae have them distributed III 15-21, IV 5-9, V 2-5. On upper part of stem and undersides of leaves of Silene repens in east Siberia.

Aphis silenicola Holman & Szelegiewicz Apterae are black, BL 1.6-2.3 mm. Apterae have secondary rhinaria distributed III 0-6, IV 0; alatae have them distributed III 5-8, IV 0-2. On apical parts of Silene spp. in Kazakhstan (Kadyrbekov 2017a), Mongolia and east Siberia.

Aphis smilacisina Zhang Apterae are green with black siphunculi and cauda; BL 1.2-1.5 mm. On Smilax china in Hangzhou, China. [The original description gives the number of caudal hairs as 12-16; nine paratypes were studied and found to have only 6-8 caudal hairs.]

Aphis smirnovi Orlova Apterae are matt dark green; BL 1.2-1.8 mm. Alatae have 2-4 secondary rhinaria on ANT III. At base of stem of Sedum purpureum (= telephium), ant-attended. Russia (Moscow and Volgograd regions).

Aphis soan (Ivanoskaya) Apterae are greenish brown; BL c. 2.6 mm. On Sanguisorba officinalis in Russia (Novosibirsk and Krasnoyarsk regions, and Tuva).

Aphis sogdiana Nevsky Apterae are pale or yellow-green, with pale antennae, legs, siphunculi and cauda; BL 1.6-2.2 mm. Alatae have 8-10 secondary rhinaria on ANT III. On undersides of leaves of Ulmus spp. in central Asia (Azerbaijan, Kazakhstan, Uzbekistan).

Aphis solanella Theobald Apterae are dull black, sometimes with white wax markings; BL 1.2-2.6 mm. Alatae have secondary rhinaria distributed III 9-22, IV 0-7, V 0-1. On a wide range of herbaceous plants, including many colonised by its close relative A. fabae, but additionally and characteristically Solanum nigrum, and also Fallopia convolvulus (Stroyan, 1984, as A. fabae solanella). A. remaudieri Börner, described from Rhododendron ferrugineum in the French Alps is possibly this species. In Europe, and with a more extensive distribution than A. fabae in Asia, Africa and South America. Heteroecious holocyclic in northern Europe with a sexual phase on Euonymus europeaus. Thieme & Dixon (2004) reviewed the taxonomic status of A. solanella. 2n=7 (5 samples from Iran) and 8 (UK).

Aphis solidaginis (Börner) Apterae are presumably dark brown or black; BL not recorded. On Solidago virgaurea in Austria, and also reported to occyr in Germany and Kazakhstan (Kadyrbekov 2017a). Apparently a member of the A. fabae group.

Aphis solidagophila Pashchenko Apterae are yellow-green; BL c.1.5 mm. Alatae have secondary rhinaria distributed III 9-20, IV 0-3 (antennae 5-segmented). Living in small to moderate-sized colonies on inner sides of leaves of Solidago decurrens near roots, attended and sheltered by ants. In east Siberia.

Aphis (Bursaphis) solitaria McVicar Baker Apterae are greenish to brownish, slightly pruinose; BL 1.6-2.2 mm. Alatae have secondary rhinaria distributed III 8-34, IV 0-8, V 0-3. On Ribes spp. in Mexico. Heteroecious holocyclic, migrating to various Compositae (Baccharis, Bidens, Cosmos, Chrysanthemum, Eupatorium, Oxyloba, Simsia, Stevia), and occasionally Penstemon (Plantaginaceae). Anholocyclic populations also persist through the year on secondary hosts. Remaudière (1993b) provided a full account of this species.

Aphis spiraecola Patch (= A. citricola van der Goot of some authors) Plate 7b Apterae are bright greenish yellow to apple green with brown head and black siphunculi and cauda (see influentialpoints.com/Gallery); BL 1.2-2.2 mm. Alatae have secondary rhinaria distributed III 6-11, IV 0-5. On a very wide range of secondary hosts in over 20 families, but especially on Caprifoliaceae, Compositae, Rosaceae (including hosts of the very similar A. pomi, q.v.), Rubiaceae and Rutaceae, and particularly on plants of shrubby habit. It is a major pest of Citrus (Blackman & Eastop 2000), and has more recently attracted interest as a pest of commercially grown yarrow, Achillea collina (Gama et al. 2010). Genetic differentiation between populations on different species of Citrus has been demonstrated in North Africa (Lebbal et al. 2019), and between populations on Rosaceae and those on other plant families in southern China (Q. Li et al. 2020; but could this be due to confusion with A. pomi?). In eastern North America it has displaced A. pomi as the most abundant aphid on apple (Brown et al. 1995), and it has also become a major pest of pears in China (Cao et al. 2012). Distribution is almost world-wide. Anholocyclic through most of the world, but a holocycle involving Spiraea spp. as primary hosts is recorded from North America, Brazil and Japan. In Japan there are also populations with a sexual phase on Citrus (Komazaki et al. 1979), but these may be a separate species or subspecies (see also Blackman & Eastop 2007). 2n=8.

Aphis spiraephaga Müller Apterae are dark greyish brown, with transverse dorsal bands of wax; BL 0.9-2.0 mm. Alatae have secondary rhinaria distributed III 6-20, IV 0-7, V 0-2. Both apterae and alate have a very variably developed pattern of dorsal sclerotisation (Müller 1961b). In dense ant-attended colonies on young shoots of Spiraea spp. However this species seems to be rather polyphagous, with records from Epilobium spp. (see Holman 1990), and other genera in several plant families (Arabis, Carum, Centranthus, Erica, Filipendula, Helipterum, Symphoricarpus, Trinia and Valeriana; Müller 1987b). Barbagallo & Stroyan (1982) reported it as commonly occurring on Centranthus ruber (Valerianaceae) in the region of Etna in Sicily. In Europe (but not UK), west Siberia, Iran, Kazakhstan (Kadyrbekov 2009b) and Mongolia (Heie 1986). Monoecious holocyclic on Spiraea, with alate males; Müller (1961b) fully described all morphs.. 2n=8.

Aphis spiraephila Patch Apterae are hazel to chestnut-brown to almost black with powdery reticulations, dark head, siphunculi and cauda, antennae and legs mainly pale (see influentialpoints.com/Gallery); BL 1.3-1.7 mm. On leaves and twigs of Spiraea spp., widespread in North America. Monoecious holocyclic with oviparae in October (Palmer 1952), males are apparently undescribed. Records from Ukraine (e.g. Coeur d’acier et al. 2010) seem to be erroneous, and records from east Asia are probably all ascribable to A. neospiraeae (see Pashchenko 1997a). 2n=8.

Aphis stachydis Mordvilko Apterae are yellowish green or dark green, with brown siphunculi and a pale cauda; BL 1.3-1.8 mm. Alatae have secondary rhinaria distributed III 4-9, IV 0-1(-3). In rolled leaves or on flower-stems and flowers of Stachys spp., especially S. recta. In central and eastern Europe, and eastward to west Siberia, Iran (Rezwani et al. 1994), Turkey (Akyürek et al. 2019), Transcaucasia (Jörg & Lampel 1988) and Kazakhstan (Kadyrbekov 2012a). Holman (1961b) found aphids with shorter siphunculi on roots of S. cretica in Crimea, that he suggested were midsummer dwarfs of A. stachydis. Monoecious holocyclic; oviparae and apterous males were described by Garcia-Prieto et al. (2004).

Aphis stranvaesiae Takahashi Apterae are green, with appendages almost entirely black; BL c.2.0 mm. Alatae have secondary rhinaria distributed III 7-9, IV 0-2. On Stranvaesia niitakayamensis (= Photinia davidiana) in Taiwan, and also recorded from Rosa sp. (Tao 1999).

Aphis subnitida (Börner) Apterae are shining black; BL 1.5-2.2 mm. Apterae have secondary rhinaria distributed III 9-18, IV 2-9, V 0-3; alatae have them distributed III 14-22, IV 4-8, V 1-3. Ant-attended colonies occur on succulent stems of Pimpinella spp. in Europe (Sweden, Denmark, Netherlands, Finland, Germany, Poland, Austria, Switzerland, Italy, Spain, Portugal, Czech Republic, former Yugoslavia, Ukraine) and west Siberia. 2n=8.

Aphis subviridis (Börner) Apterae are pale green; BL 1.0-1.3 mm. On roots and rosette leaves of Potentilla spp., especially P. argentea, in central and eastern Europe (Germany, Poland, Czech Republic, Latvia, Byelorussia, Ukraine) and Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic; oviparae and apterous males were described by Stekolshchikov (2008c).

Aphis succisae Holman Apterae are brown to velvet black; BL 1.3-2.0 mm. Alatae have secondary rhinaria distributed III 12-16, IV 1-3. On Succisa pratensis, at stem bases and on ground-level leaves in ant shelters. In Czech Republic (southern Bohemia). Monoecious holocyclic with oviparae and apterous males in September-October (original description).

Aphis sumire Moritsu Apterae are matt dark green to black; BL 1.0-1.2 mm. Alatae have 4-6 secondary rhinaria on ANT III only (Takahashi 1966). In ant-attended colonies on Viola spp. in east Asia (Japan, China, Korea, east Siberia).

Aphis symphyti Schrank Apterae are pale yellowish green to dark green, rarely lemon yellow, with dark head and siphunculi (see influentialpoints.com/Gallery); BL 1.3-2.0 mm. Alatae have secondary rhinaria distributed III 8-12, IV 1-4, V 0-2 (cf. gossypii). Living scattered under leaves of Symphytum officinale, or when numerous extending up the stems and in the influorescences (Stroyan 1984). Also recorded from various other Boraginaceae, but many of these records may apply to A. gossypii, with which there is wide overlap in many features of morphology. Recent molecular work suggests a close relationship with A. frangulae (Cocuzza et al. 2009, Cocuzza & Cavalieri 2014; but see also under frangulae). Throughout Europe (except Scandinavia and Iberian peninsula). Monoecious holocyclic, with alate males; the life cycle is only known to be completed on S. officinale.

Aphis sywangi Zhang, Cheng, Zhong & Li Colour of apterae in life is unrecorded; BL c.2.3 mm. On Apocynum venetum in Xinjiang, China (G. Zhang et al. 1999a).

Aphis tacita Huculak Apterae are dull blackish green, finely wax-dusted; BL 1.4-2.5 mm. In large ant-attended colonies on roots, and in ant shelters at bases of stems, of Astragalus arenarius. Described from Poland, and subsequently recorded from Astragalus sp. in Kazakhstan (Kadyrbekov 2017a). Alatae are unknown. Monoecious holocyclic, oviparae and apterous males appearing in late August-September (original description). A. astragalicola in central Russia is probably a close relative.

Aphis takagii Takahashi Colour of apterae in life is unrecorded; BL 1.7-2.2 mm. Alatae have secondary rhinaria distributed III 8-13, IV 0-3. On stem and upper and lower leaf surfaces of Veratrum spp. in Asia (Japan, and east and west Siberia). Monoecious holocyclic, with oviparae and aptedrous males appearing in June (original description, as A. veratri ssp. takagii).

Aphis talcensis Nieto Nafría, Ortego & Mier Durante Apterae are very dark green to black, without wax powder; BL 1.1-1.7 mm. On Adesmia papposa (var. papposa) in Chile (Talca province; Nieto Nafría et al. 2021b). Alatae and other morphs are unknown.

Aphis talgarica Kadyrbekov Apterae are blackish brown with grey tinge; BL 1.6-1.9 mm. In small colonies on leaf petioles of Bupleurum spp. in Kazakhstan (Kadyrbekov 2001a, 2012a), and also recorded from B. aureum in western Siberia (Stekolshchikov & Novgorodova 2013).

Aphis taraxacicola (Börner) Apterae are mottled dark green (see influentialpoints.com/Gallery); BL 1.3-2.3 mm. Alatae have secondary rhinaria distributed III 5-13, IV 0-4, V 0-1. On root collar and under rosette leaves of Taraxacum spp., tented over with soil by ants, most often in dry, open locations. Throughout Europe, and in Iran (Hodjat 1993), Kazakhstan (Kadyrbekov 2017a), Tajikistan (as A. plantaginis ssp. asiatica; see Stekolshchikov 2014a), Pakistan (Naumann-Etienne & Remaudière 1995), Siberia (Pashchenko 1997a, Holman 1987), China (G. Zhang 1999), Japan (Sorin 1992), and an alata has been trapped in Quebec, Canada (Cloutier et al. 1986). Some distribution records may apply to A. nevskyana Kadyrbekov (q.v.). A taraxacicola (Börner) is monoecious holocyclic with apterous males. A. eugenyi (Ivanoskaya), described from T. officinale in west Siberia, is not clearly distinct and may be this species. 2n=8.

Aphis tashevi Szelegiewicz Apterae are probably green; BL 1.8-2.0 mm. Alatae have secondary rhinaria distributed III (7-) 21-25, IV (0-) 9-14, V (0-) 3-5. On Stachys sp. in Bulgaria. The apterae are very similar to A. stachydis, and might be considered as alatiform specimens of that species, but for the correspondingly high rhinarial numbers of four of the five alatae present in the type series (the fifth alata may be stachydis).

Aphis taukogi Shinji Only alatae are described; green, BL c. 1.6 mm, with secondary rhinaria distributed III 28-32, IV 8-15. On leaves, stems and flowers of Bidens tripartita in Japan. Monoecious holocyclic according to Shinji, with eggs laid on stem near ground, but the species has apparently not been more recently collected.

Aphis tehuelchis Nieto Nafría & López Ciruelos Apterae are green with or without variably developed dorsal abdominal brown-black cross bands or a central patch; BL 1.1-1.8 mm. Alatae have 4-8 secondary rhinaria on ANT III. On Euphorbia sp. in Argentina (López Ciruelos et al. 2016). The life cycle is unknown, but presumably monoecious holocyclic. [Morphologically very similar to A. brotericola, but differing in appearance in life and number of secondary rhinaria in alatae.]

Aphis teucrii (Börner) Apterae are bright green to dark green or dark grey-blue; BL 1.0-1.9 mm. Alatae have 4-6 secondary rhinaria on ANT III. On Teucrium spp., especially chamaedrys, feeding on shoot apices and causing curling of leaves (cf. A. alienus). Known from various countries in Europe (England, Channel Islands, France, Austria, Switzerland, Spain, Italy, Czech Republic, Slovakia, Slovenia, Poland, Hungary, Rumania, Ukraine), and from Iran, Iraq and Lebanon. Monoecious holocyclic.

Aphis thalictri Koch Apterae are greyish to yellowish green powdered with grey wax; BL 1.9-2.8 mm. Alatae have secondary rhinaria distributed III 6-11, IV 0-2. Under leaves, up stems and in flowerheads of Thalictrum spp. (Stroyan, 1984). In Europe (England, Denmark, Sweden, Finland, Germany, Switzerland, Russia, Byelorussia), west Siberia, Central Asia, and also recorded from North Korea (Lee et al 2002c). Foottit et al. (2006) reported its occurrence in British Columbia, Canada. Monoecious holocyclic. Bozhko (1976a) described a population from Thalictrum sp. in Ukraine as a subspecies, A. thalictri ssp. orangii. [Another form that was originally described as a subspecies of A. thalictri (neothalictri Pashchenko) is here treated as a separate species.]

Aphis thaspii Oestlund Apterae are dark green with chocolate brown head and thorax, and brown patches at bases of siphunculi (Hottes & Frison 1931, as A. signatis); BL 1.3-1.8 mm. Alatae have secondary rhinaria distributed III 28-56, IV 5-11, V 0-6. On various Umbelliferae/ Apiaceae (Cicuta maculata, Sanicula sp., Thaspium spp., Zizia aurea) in USA and Canada (Manitoba). Very similar in morphology to A. saniculae (q.v.). 2n=8.

Aphis thecomae Davletshina Apterae are dark brown; BL c.1.5 mm. On stem apices, flowerheads and fruit of Campsis radicans in Uzbekistan. Monoecious holocyclic with oviparae and alate males in October (original description).

Aphis thesii Holman Apterae are black; BL 1.4-1.9 mm. Alatae are undescribed. At stem bases and on root collars of Thesium spp., attended by ants. In Czech Republic, Slovakia, and Poland (Osiadacz & Halaj 2010). Monoecious holocyclic with apterous males (original description).

Aphis thomasi (Börner) Apterae are straw yellow; BL 1.1-1.3 mm. Alatae have secondary rhinaria distributed III 4-12, IV 0-4, V 0-1. On basal parts of Scabiosa spp., and also recorded from species of Dipsacus and Knautia, and Pycnocomon rutifolium. In Europe, eastward to Ukraine and Turkey, Kazakhstan (Kadyrbekov 2012a) and also recorded from Algeria (Laamari et al. 2013). Müller (1987a) compared morphology of apterae with those of A. confusa and A. ochropus. Closely related to A. confusa, but with shorter hairs. Kanturski & Bezdĕk (2019) provided a redescription, and a first description of the sexual morphs, collected on Scabiosa in late September.

Aphis tianschanica Kadyrbekov Apterae are blackish brown; BL c.2.0-2.1 mm. On roots of Potentilla bifurca in south-east Kazakhstan, ant-attended (Kadyrbekov 2001a).

Aphis tirucallis Hille Ris Lambers Apterae are dark brown to black, wax-dusted; BL 1.5-2.0 mm. Alatae have secondary rhinaria distributed III 4-11, IV 0, V 0. On Euphorbia spp. in southern Europe, Azores, Canaries, Madeira, Balearics, Yemen (van Harten et al. 1994), widely distributed in Africa (Millar 1994), and this seems to be the species introduced to Australia (Eastop 1966, as A. euphorbiae). There are also specimens in the BMNH collection from Lebanon, Turkey and Pakistan. It was described originally as a subspecies of A. euphorbiae. Nieto Nafría et al. (2005a) provided a redescription. Apparently anholocyclic everywhere, although Nieto Nafría & Mier Durante (1977, cited in Nieto Nafría et al. 2005a) found viviparous females with scent glands on the hind tibiae in the Canaries.

Aphis tomenthosi Bozhko Apterae are brown, with dark siphunculi and cauda; BL c. 1.8 mm. On Arctium tomenthosa in Ukraine, and also recorded from Kazakhstan (Kadyrbekov 2017a) .

Aphis toriliae Rusanova Apterae are brownish yellow or chocolate-coloured, with dark siphunculi and cauda; BL 1.6-2.1 mm. Alatae have secondary rhinaria distributed III 5-20, IV 0-2. On Torilis arvensis in Azerbaijan.

Aphis tormentillae Passerini Apterae are greenish black (see influentialpoints.com/Gallery); BL 1.0-1.7 mm. Alatae have 4-8 secondary rhinaria on ANT III. On Potentilla spp., especially erecta, tending to be dispersed on stems, petioles and undersides of leaves, not usually visited by ants. Widely distributed in Europe, and also now recorded from Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic with alate males.

Aphis torquens Holman Apterae are dark brown to blue-black; BL 1.9-2.9 mm (spring), 1.4-1.8 mm (summer). Alatae have secondary rhinaria distributed III 7-20, IV 0-4, V 0. On undersides of young leaves of Petasites spp., which are strongly curled and crumpled, attended by ants. Recorded from Czech Republic and Hungary (BMNH collection). There are differences in R IV+V length and hair length between these two populations. A record from east Siberia (Holman 1987) is referable to A. fukii (Pashchenko 1997b). Monoecious holocyclic, with oviparae and apterous males in October (original description).

Aphis triglochinis Theobald Apterae are dirty green or brownish, BL 1.5-2.7 mm. Alatae have secondary rhinaria distributed III 25-60, IV 9-25, V 7-17. Migrating from the primary host Ribes to a variety of secondary host plants living in wet places, including species of Anemone, Barbarea, Borago, Cardamine, Drosera, Myosotis, Nasturtium, Portulacea, Rorippa, Triglochin and Veronica. In north, central and eastern Europe, and across Asia to Mongolia and east Siberia. Possibly also in Japan, as either or both of two species described by Shinji, A. cardamine (currently regarded as a nomen dubium) and A. neopolygoni (q.v.), could be this aphid, having similarly short antennal terminal processes and alatae with numerous secondary rhinaria, although described as salmon red/brown in life. A. veroniciphaga (q.v.) described from Korea is also possibly this species. Rakauskas (1998a) reviewed the literature and host records of A. triglochinis, and discussed the possible existence of host races. 2n=8.

Aphis tripolii Laing Apterae are green with dusky head, with appendages dusky to dark distally (see influentialpoints.com/Gallery); BL 1.3-2.5 mm. Alatae have secondary rhinaria distributed III 6-18, IV (0-)2-6, V 0-3. On upper parts of leaves and in inflorescences of Aster tripolium, in coastal salt marshes or on mud flats. In Europe (UK, Franc, Germany, Italy) it seems to be specific to this plant and habitat, but Kadyrbekov (2017a) reports it as occurring on Bidens cernua in Kazakhstan. Monoecious holocyclic with alate males.

Aphis tropaeoli Nieto Nafría & Brown Apterae are brown lightly powdered with wax; BL 1.4-2.4 mm. Alatae have 9-17 secondary rhinaria only on ANT III. On Tropaeolum incisum in Argentina (Mendoza province, at high altitude) and on an unidentified Tropaeolum sp. at lower altitudes in Chile (Nieto Nafría et al. 2019).

Aphis tsujii Shinji Apterae are dark brown, with black siphunculi and cauda; BL 1.9-2.5 mm. Alatae have secondary rhinaria distributed III 13-18, IV 0, V 0-6. On Solidago and Eupatorium spp. in Japan (Takahashi 1966).

Aphis typhae Mamontova Appearance of apterae in life unknown; BL c.1.7 mm. On Typha spp. in Ukraine.

Aphis ucrainensis Zhuravlev (= Aphis turkestanica Kadyrbekov) Apterae are shining black, slightly wax-dusted; BL 1.2-2.1 mm. Alatae have secondary rhinaria distributed III 7-10, IV 0-2. In compact colonies on young shoots of Spiraea spp. in Ukraine and eastward to west Siberia and Kazakhstan (Kadyrbekov 2001a, as A. turkestanica, and 2012a). Monoecious holocyclic with oviparae and apterous males in October (Zhuravlev 1997).

Aphis ulicis Walker Apterae are blackish green with a shiny black dorsal shield, coated with silvery grey wax (see influentialpoints.com/Gallery); BL 1.3-2.6 mm. Alatae have secondary rhinaria distributed III 3-9, IV 0-1. In dense ant-attended colonies on young growth, flowers and seed-pods of Ulex spp., especially U. europaeus. In western Europe (UK, Ireland, Netherlands, France, Spain, Portugal, Madeira). Monoecious holocyclic with alate males. For a fuller account see Jacob (1948b). 2n=8.

Aphis ulmariae Schrank Apterae are green, mottled with dark green, and darker head and legs; BL 1.1-2.4 mm. Alatae have secondary rhinaria distributed III 4-8, IV 0-1. On Filipendula spp., in ant-attended colonies, curling leaves, and also recorded from Spiraea salicifolia. In Europe, west and east Siberia (Holman 1987), Kazakhstan (Kadyrbekov 2009b) and Korea (Lee et al. 2002c). Records from USA are referable to other species (Foottit et al. 2006). Monoecious holocyclic, with alate males.

Aphis (Debilisiphon) umbelliferarum (Shaposhnikov) Apterae are very dark green slightly shiny, with antennal segment III and basal parts of tibiae whitish; BL c. 1.8-2.1 mm. In colonies at stem bases and on radical leaves of Umbelliferae/Apiaceae (Cachrys, Cenolophium, Peucedanum, Seseli, Torilis), usually in ant shelters. In Russia (Ural Mountains) and Kazakhstan (Kadyrbekov 2017a). [A. seselii (Bozhko) is probably a synonym, and it is possible that umbelliferarum is itself a synonym of A. selini (Börner).]

Aphis umbrella (Börner) (= Aphis rostellum; see Stekolshchikov & Buga 2017) Apterae are pale green or yellowish green, usually mottled with darker green, with pale or dark-tipped siphunculi (see influentialpoints.com/Gallery); BL 1.7-2.3 mm. Alatae have secondary rhinaria distributed III 4-13, IV 0-5, V 0-1. On Malva spp. and certain other Malvaceae, causing umbrella-like leaf-curl of terminal leaves. Ant-attended. In Europe, North Africa (Laamari et al. 2013), across Asia to China (where described as Rhopalosiphum rostellum), and introduced to North America (Foottit et al. 2006); records from elsewhere are probably mostly due to confusion with A. nasturtii and A. gossypii. Monoecious holocyclic with alate males in northern Europe, probably anholocyclic in warmer regions. Stekolshchikov & Buga (2017) provided descriptions of the fundatrix and sexual morphs. 2n=6* and 7 in Iran, 2n=8* in Israel, Cyprus, Italy and UK.

Aphis unaweepiensis Hottes Apterae are pale blue-grey to greenish grey; BL 2.0-3.0 mm. On roots of Sarcobatus vermiculatus in Colorado, USA. This species possibly belongs in Protaphis, and may be related to P. middletonii, but differs in its longer R IV+V and less hairy cauda.

Aphis urticata Gmelin Apterae are dark bluish green in spring, with dwarfish pale yellow forms appearing in summer (see influentialpoints.com/Gallery); BL 0.90-2.2 mm. Alatae have secondary rhinaria distributed III 5-11, IV 0-5, V 0-1. In dense ant-attended spring colonies at tips of growing shoots of Urtica dioica, and later as summer dwarfs on undersides of leaves. In Europe, Middle East, Central Asia, Pakistan (Naumann-Etienne & Remaudière 1995) and North America. Monoecious holocyclic with apterous males; life cycle and biology were studied by Müller & Holtfreter (1978).

Aphis (Zyxaphis) utahensis (Knowlton) Colour of apterae in life is unrecorded, BL 1.5-1.6 mm. Described from Seriphidium tridentatum in Utah, USA.

Aphis utilis Zhang Apterae about 1.4 mm, appearance in life unrecorded. Collected on Rhamnus utilis at Beijing, China (Zhang & Zhong 1983). Apparently a member of the A. frangulae group.

Aphis utsugicola Monzen Apterae are yellowish green with brown head and large dark brownish dorsal abdominal spot; BL c.1.9-2.0 mm. Alatae have secondary rhinaria distributed III 8-10, IV 5-6. Colonies roll the terminal leaves of Deutzia crenata and D. scabra var. sieboldiana in Japan.

Aphis uvaeursi Ossiannilsson Apterae are dark brownish green or black; BL 1.3-1.9 mm. Apterae have secondary rhinaria distributed III 0-14, IV 0-2 , and alatae have them distributed III 9-17, IV 0-2, V 0-1. On Arctostaphylos uva-ursi, forming dense colonies on shoot apices of prostrate mats, tented over by ants. In Scotland, Norway, Sweden, Finland, Germany, Poland and north-west Russia. Monoecious holocyclic with apterous males (Szelegiewicz 1976).

Aphis vaccinii (Börner) Apterae are shining greenish black with a fine wax bloom; BL 1.3-2.0 mm. Alatae have secondary rhinaria distributed III 8-17, IV 0-4. On stems of Vaccinium spp. and Andromeda spp., ant-attended. In Europe north of Alps, west Siberia, and introduced to British Columbia, Canada (Foottit et al. 2006). Monoecious holocyclic with alate males (Heie 1986). Closely related to the Aphis fabae group, but usually smaller and with more development of dorsal sclerotisation.

Aphis valerianae Cowen Apterae are blackish green with dusky or brownish antennae and legs, and black siphunculi and cauda; BL 1.8-2.5 mm. Alatae have secondary rhinaria distributed III 27-42, IV 0-16, V 0-14. On leaves and stems of Valeriana spp. in western North America (British Columbia, Oregon, Idaho, Utah, Colorado, New Mexico; BMNH collection). Monoecious holocyclic with oviparae and alate males in September (Palmer 1952).

Aphis vallei Hille Ris Lambers & Stroyan Apterae are black covered in greyish white wax powder; BL 1.4-2.1 mm. Alatae have secondary rhinaria distributed III 9-16, IV 2-5, V 0-3. Described from “a very tall Euphorbia sp.” in Italy, and since found on various Euphorbia spp. Reported to occur in Spain, Portugal, the Balearics, France, Bulgaria, Greece, Ukraine and Turkey (Nieto Nafría et al. 2005a).

Aphis (Toxopterina) vandergooti (Börner) (Fig.5j) Apterae are dark blue-green, or sometimes yellow (see influentialpoints.com/Gallery); BL 1.4-2.0 mm. Alatae have 3-7 secondary rhinaria on ANT III. In ant shelters on roots, stolons and basal leaf petioles of composite plants of tribe Anthemideae, e.g. Achillea, Matricaria, Tanacetum. Widespread in Europe, in Iran (Rezwani et al. 1994), Kazakhstan (Kadyrebekov 2012a), and there is also a record from China (L. Zhang & G. Zhang 2002b). Monoecious holocyclic, with apterous males. A. obiensis (Ivanoskaya), described from Achillea millefolium in Russia are probably small specimens of this species.

Aphis (Bursaphis) varians Patch Apterae are dark blue-green with pale siphunculi (see aphidtrek.org); BL 1.0-2.2 mm. On spring growth of Ribes spp., crumpling leaves and stunting growth. Heteroecious holocyclic, migrating in late spring to Onagraceae, especially Epilobium angustifolium, where its relations with ants have been intensively studied (e.g. Breton & Aldicott, 1992). Patch (1927) gave life cycle details. In North America, and possibly Mongolia (BMNH collection, leg. H. Szelegiewicz) and east Siberia (Pashchenko 1988a), although Pashchenko recorded Borago officinalis and Portulaca oleracea as secondary hosts, suggesting that the east Asian species may be distinct. 2n=8.

Aphis veratri Walker Apterae are dark brown or black, not shining, with slight greyish bloom (see influentialpoints.com/Gallery); BL 1.5-2.3 mm. Alatae have secondary rhinaria distributed III 10-16, IV 0-3, V 0-1. On undersides of leaves of Veratrum spp. in Europe (not UK, Scandinavia or Iberia), eastward to eastern Kazakhstan. Records from east Siberia (Pashchenko 1988a) seem to apply to the North American species A. cercocarpi. Monoecious holocyclic, with apterous males and oviparae in August in Switzerland (Lampel & Meier 2007).

Aphis verbasci Schrank Apterae are bright golden yellow to pale green, slightly pulverulent, with black somewhat curved siphunculi (see influentialpoints.com); BL 1.7-2.5 mm. Alatae have secondary rhinaria distributed III 20-27, IV 2-8. Under basal leaves of Verbascum spp., ant-attended, and also recorded from Buddleja spp., and from Scrophularia . Monoecious holocyclic on these hosts; Kanturski et al. (2014a) described and illustrated the oviparae and apterous males, collected in UK and Poland in October. In Europe (except Scandinavia), Middle East, North Africa, Kazakhstan (Kadyrbekov 2014e), India and Pakistan. A record from west Siberia (Ivanoskaya 1977) is probably a misidentification (Stekolshchikov & Khureva 2020). Some records from north India (Chowdhuri et al. 1969a) are from unlikely hosts (Senecio and Polygonum), and possibly apply to another species. 2n=8.

Aphis vernoniae Thomas (Fig.57d) Apterae are greenish yellow with pale appendages; BL c.1.4-1.6 mm. Alatae have secondary rhinaria distributed III 4-8, IV 0-1. On growing points, flower stems and undersides of leaves of Vernonia spp., and also recorded from Eupatorium and Helenium (Hottes & Frison 1931). In eastern and central USA.

Aphis veronicicola Holman Apterae are dull black; BL 1.6-2.1 mm. Alatae have secondary rhinaria distributed III 4-7, V 0-2. On stem apices and inflorescences of Veronica (= Veronicastrum) sibirica, attended by ants. In east Siberia (Ussurijsk).

Aphis veroniciphaga Kim & Lee Colour of apterae in life unrecorded; BL 1.3-1.5 mm. Alatae are undescribed. In small colonies on shoots, stems and leaves of Veronica nakaiana on Ulleung-do Island, South Korea (Kim et al. 2006a). Possibly this is a synonym of A. triglochinis.

Aphis versicolor (Börner) Apterae have body and siphunculi dark green, BL 1.1-1.6 mm. On leaves of Rhamnus cathartica in Austria and Bulgaria (Börner 1950). This species has similar very large marginal tubercles to A. mammulata on this host from other parts of Europe, but much shorter hairs on antennae and marginal areas of abdomen. It was synonymised with A. mammulata by Börner (1952), and should perhaps be regarded as a variant population of that species. Monoecious holocyclic; males are apterous and orange-brown, as in mammulata.

Aphis verticillatae (Börner) (Fig.48g) Apterae are mottled yellow-green; BL 1.5-1.8 mm. Alatae have 3-5 secondary rhinaria on ANT III. On Salvia verticillata, on undersides of basal leaves and young shoots. In central and eastern Europe (Austria, Germany, Hungary, Poland, Czech Republic). Aphids identified as this species have also now been collected in southern Italy, on S. fruticosa (Cocuzza et al. 2014).

Aphis viburni Scopoli Apterae are brownish green to black, sometimes with paired dorsal wax markings (see influentialpoints.com/Gallery); BL 1.5-3.0 mm. Alatae have secondary rhinaria distributed III 8-24, IV 0-6. On Viburnum spp. (especially V. opulus), in curled leaves, ant-attended. Possibly sometimes on orchids of the genus Epipactis, if A. epipactis is a synonym (Stroyan 1984). Europe, Central Asia, and west Siberia. Records from North America seem to be unsubstantiated (Foottit et al. 2006). Monoecious holocyclic with apterous males. 2n=8.

Aphis viburniphila Patch Apterae are reddish brown mottled with paler shades, with blackish head, and with broad dark areas around and behind the black siphunculi (see influentialpoints.com/Gallery); BL 1.7-2 mm. Alatae have secondary rhinaria distributed III 20-28, IV 5-7, V 0-2. On leaves and stems of Viburnum spp., apparently widely distributed in North America. Monoecious holocyclic, with oviparae in October (males apparently not recorded).

Aphis (Toxoptera) victoriae (Martin) Apterae are shining black, immatures are dull reddish brown; BL 1.6-1.8 mm. In ant-attended colonies on young shoots of Zanthoxylum scandens in Hong Kong (Martin 1991), and subsequently recorded from Guanxi province on Z. scandens and Z. schinifolium (Qiao et al. 2008). There is also a record from Schima crenata in Guanxi, but this is unlikely to be a true host. It may occur on other Zanthoxylum that are of tree habit, and will key to Aphis odinae in the polyphagous aphid key. For distinguishing features see Martin (1991).

Aphis vineti Hoffmann Apterae are blackish, without wax (?); BL 1.3-1.9 mm. Alatae have secondary rhinaria distributed III 3-10, IV 0-1. On ground stems and roots of Coronilla (= Securigera) varia in Germany. Monoecious holocyclic with oviparae and apterous males in October (Hoffmann 1972).

Aphis violae Schouteden Apterae are dark green mottled with pale green (see influentialpoints.com/Gallery); BL 1.1-2.0 mm. Alatae have secondary rhinaria distributed III 5-17, IV 0-3. On basal and subterranean parts of Viola spp., tented over by ants. In Europe (UK, Belgium, Denmark, Netherlands, Finland, Germany, Austria, Czech Republic, Hungary, Poland, and one record from Spain). Monoecious holocyclic with apterous males. 2n=8.

Aphis violaeradicis Pashchenko Apterae are brown, immatures green; BL c.1.5 mm. Alatae have secondary rhinaria distributed III 3-5, IV 0, V 0-1. On Viola spp., living at base of stem or on root or leaf rosette, often in an ant shelter. In east Siberia. Monoecious holocyclic (original description). Although it is clearly distinct in its biology, the description of this species does not differentiate it morphologically from A. gossypii. Chu (1949) recorded A. gossypii with eggs on roots of Viola spp. in China which could be this species.

Aphis viridissima Mamontova Apterae are bright green, BL c.1.6-1.7 mm, on terminal shoots of Tamarix sp. in Ukraine (Mamontova 1955), and since reported from Tamarix ramosissima in Kazakhstan (Kadyrbekov 2017a). Others morphs and life cycle are unknown.

Aphis vitalbae Ferrari Apterae varying from very dark green to pale green or yellowish in summer, without wax (cf. A. clematidis), with considerable variation in pigmentation of body and appendages, the siphunculi being pale, dark-tipped or wholly dark; BL 1.2-1.9 mm. Alatae have secondary rhinaria distributed III 5-8, IV (0-)1-5, V 0-1. On shoot tips, petioles and undersides of leaves of Clematis spp. (vitalba, flammula) in Europe (Poland, Czech Republic, Bulgaria, France, Greece, Italy, Rumania, Spain, UK) and Georgia (Barjadze et al. 2009). Monoecious holocyclic with oviparae and apterous males in October-November (Tashev 1964b).

Aphis vitexicola Kim & Lee Colour in life is unrecorded; BL 1.5-1.6 mm. Alata (one specimen) has 6 secondary rhinaria on ANT III. On shoots, stems and undersides of leaves on Vitex negundo in South Korea (Kim et al. 2006a). This species is not included in the key to apterae on Vitex because there are no clear characters that distinguish it from Aphis viticis.

Aphis viticis Ferrari Apterae are grass-green with brown siphunculi in spring to pale straw-coloured with pale siphunculi in midsummer dwarfs; BL 0.8-1.7 mm. Alatae have secondary rhinaria distributed ANT III 4-9, IV 0-2. On Vitex agnus-castus in southern Europe and the Middle East (Spain, Mallorca, France, Italy, former Yugoslavia, Israel, Iran, Turkey). Monoecious holocyclic, with alate males. Barbagallo & Stroyan (1982) redescribed apterous and alate viviparae, and Nieto Nafría et al. (1986) provided brief descriptions of the sexual morphs.

Aphis vladimirovae Orlova Apterae are matt brownish black; BL 1.7-2.1 mm. Alata (1 specimen) has 4-5 secondary rhinaria on ANT III. On stems of Corispermum marschalii, attended by ants. In Russia (Volgograd province).

Aphis vurilocensis Nieto Nafría, Brown & López Ciruelos Colour in life is unknown; BL 1.1-1.2 mm. On Mulinum spinosum in Rio Negro province, Argentina (Gonzáles Rodríguez et al. 2017).

Aphis wahena Hottes & Wehrle Apterae are probably black; BL c.1.1-1.5 mm. Alatae have secondary rhinaria distributed III 12-15, IV 6-9, V 3-4. On Viguiera cordifolia in Arizona, USA.

Aphis wartenbergi (Börner) Colour of apterae in life is unrecorded; BL c.1.5-1.7 mm. On Carduus spp., rolling ground-level leaves. Recorded from Austria, Slovakia, Romania, Turkey and Georgia. Monoecious holocyclic according to Börner.

Aphis wellensteini (Börner) Apterae are dark green or olive; BL 1.9-2.1 mm. On Verbascum spp. in central and eastern Europe (Austria, Czech Republic, Ukraine).

Aphis whiteshellensis Rojanavongse & Robinson Apterae are reddish-brown, BL 1.5-1.8 mm. In tightly curled leaves at ends of small branches of Amelanchier alnifolia in Manitoba, Canada; also collected from Amelanchier sp. in Montana, USA (BMNH collection, leg. D.J. Voegtlin). Monoecious holocyclic, with apterous males (Rojanavongse & Robinson 1977).

Aphis xylostei (Börner) Apterae are probably dark greenish brown; BL unrecorded. On Lonicera xylosteum in Austrian Alps.

Aphis yangbajaingana Zhang Apterae are dark brown, wax-dusted; BL 1.7-1.8 mm. On Medicago sativa in Tibet. Oviparae on a paratype slide were collected in late September. Very similar to A. coronillae (q.v.).

Aphis zamorana Garcia Prieto & Nieto Nafría Apterae are brownish green, with black siphunculi and cauda; BL 1.1-1.3 mm. Living all-year-round on Rhamnus frangula (= Frangula alnus) in Zamora, Spain. Oviparae and apterous males were collected along with apterous viviparae in late October (Garcia Prieto & Nieto Nafría 2005). Alate viviparae are unknown.

Aphis zapalina Mier Durante & Ortego Apterae are rather plump-bodied, dark brown with black siphunculi and cauda; BL 2.2-2.6 mm. In rather compact groups on thin stems of Adesmia boronioides in Neuquen province, Argentina (Nieto Nafría et al. 2016c). Other morphs and life cycle are unknown.

Aphis zonassa Knowlton Apterae black, with wholly black appendages; BL c.1.5-1.7 mm. Alatae have secondary rhinaria distributed III 15-21, IV 9-11. On Chrysothamnus parryi and Chrysothamnus sp., in Utah and Colorado, USA.

Aphis zweigelti Börner Colour of apterae in life is unrecorded, probably blackish and densely covered with grey wax; BL 1.6-2.0 mm. Described (briefly) from Chamaecytisus supinus (=Cytisus capitatus) in Austria, and subsequently recorded from C. supinus in Austria (BMNH collection, leg. Franz) and Bulgaria (Szelegiewicz 1962b). The specific distinction from A. cytisorum, however, remains unclear.

Aphthargelia Hottes

Aphidinae: Macrosiphini

Two North American species on Symphoricarpos somewhat resembling Brevicoryne but with dark, very distinctly reticulate dorsal sclerotisation, and dark rounded marginal tubercles on ABD TERG 1-5. Jensen (2013) reviewed the genus, providing a key to distinguish it from morphologically similar genera in North America.

Aphthargelia rumbleboredomia Jensen Apterae are distinctive in life, with head and anterior part of dorsum black, broken on first abdominal segment by a broad white wax cross-band with a gap in the middle; legs, basal parts of antennae, posterior dorsal abdomen and cauda pale; siphunculi black with red internal pigment surrounding their bases (Jensen 2013, and aphidtrek.org); BL 1.9-2.3 mm. Alatae have secondary rhinaria distributed III 55-77, IV 25-61, V 0-15. Heteroecious holocyclic, with Aconogonon spp. as secondary hosts, and a sexual phase on Symphoricarpos, on which oviparae were found in September (Jensen 2013). In mountainous regions of north-western USA. Colonies on the secondary host may be attended by ants, but the actions of Formica fusca were reported to be predatory by Andersen (1991, as A. symphoricarpi).

Aphthargelia symphoricarpi (Thomas) Plate 12h Apterae are dorsally black, dark blue or dusky green with variably developed powdery white cross-bands on abdominal tergites 1, 4, 5 and 8; siphunculi black, but antennae and legs mostly pale, as are the end of abdomen and cauda (see aphidtrek.org); BL 1.3-2.8 mm. Alatae have secondary rhinaria distributed III 38-64, IV 20-37, V 3-14. Jensen (2013) provided a redescription of all morphs. On twigs of Symphoricarpos spp. in western and central USA and Canada. Monoecious holocyclic, with oviparae and apterous males in September-October in the Rocky Mountain region (Palmer 1952), but Jensen (2013) collected small alate males in summer as well as autumn, and oviparae with many more scent plaques on the hind tibiae than those described by Palmer. He has also made successful transfers of an aphid identified as A. symphoricarpi from Symphoricarpos to Bistorta (= Polygonum) bistortoides (see aphidtrek.org), suggesting that there may be more than one species under this name. 2n=14.

Aploneura Passerini

Eriosomatinae: Fordini

A small palaearctic genus with alternation between galls on Pistacia and the roots of Poaceae and vines, but also a strong tendency to anholocycly on the roots of secondary host plants. Apterae have very short 4‑ or 5‑segmented antennae; alatae have 6‑segmented but unusually short antennae which are distinctive because ANT III and IV both bear a single, large secondary sensorium on the distal half, resembling the primary sensoria on V and VI. Apterae on roots produce copious white wax and are not attended by ants. Alatae hold their wings flat in repose, unlike Baizongia and Slavum (q.v.), in both of which the wings are held roof‑like. However in the case of Slavum, no satisfactory structural characters have been found to support its generic separation from Aploneura, which is therefore in doubt.

Aploneura ampelina Mokrzecky (= Slavum lentiscoides Mordvilko?) Apterae are pale, in white wax; BL c.1.6 mm. On roots of Vitis vinifera in southern Europe, Crimea and Central Asia; also in South Africa and probably more widespread. Also reported from roots of woody Rosaceae (Cydonia, Malus, Pyrus; Mamontova 1955). Occurrence on grass roots requires confirmation. Alatae of A. ampelina may be distinguished from those of A. lentisci by the shape of the sensorium on ANT VI, which has its long axis across the segment in ampelina, along the segment in lentisci. Populations seem to be mostly anholocyclic, but whether a sexual phase occurs anywhere on Pistacia is still unclear; it may be synonymous with Slavum lentiscoides which forms galls on Pistacia vera, but experimental work is needed to confirm this. See Mordvilko (1935) for further information.

Aploneura lentisci (Passerini) Plate 1f, fig. 81A Apterae on grass roots are long-bodied, pale yellow with darker head, covered with fine white wax which is flocculent at posterior end (see influentialpoints.com/Gallery); BL 1.1‑3.0 mm. They colonise the roots of numerous species of Poaceae, and occasionally of dicots (e.g. Ranunculus, Veronica). In western, central and southern Europe, Middle East, Central Asia, Africa (Morocco, Kenya, Nigeria, Zimbabwe), Australia, New Zealand, Argentina, Chile, and North America. Contrary to Blackman & Eastop (1994, 2000), it has been in North America since at least 1879 (Foottit et al. 2006). Heteroecious holocyclic in Mediterranean region, galling the leaves of Pistacia lentiscus, with the two-year life cycle that is typical of Fordini. The galls are pocket-like, often kidney-shaped, formed by invagination of the leaf alongside the mid-rib (fig. 126A). Their ultrastructure was studied by Álvarez et al. (2016). They appear in late March-April (in Israel), and grow quickly to attain their maximum size by May, although the fundatrix inside does not start to reproduce until early June. Galls open in August-November and emigrant alatae (1.3-2.3 mm) leave as they become mature, over an extended period. Wool & Manheim (1986, 1988) studied the population ecology of gall generations in Israel. Elsewhere it is anholocyclic on grass roots, but large flights in late summer may consist almost entirely of sexuparae, even where primary host is unavailable. Wool & Kurzfeld-Zexer (2008) studied colonies on cereal roots in culture, noting successful colonisation of wheat and barley, but not of oats. Nahum et al. (2010) found little genetic differentiation between populations on P. lentiscus in the Mediterranean basin. See also Blackman & Eastop (2000). 2n = 16.

Aploneura werthi Börner Apterae are wax-covered; BL 1.4-1.8 mm. On roots of Carex ?curvula in the French Alps (Börner 1952, p. 476).

Appendiseta Richards

Calaphidinae: Therioaphidini

A genus with one North American species similar to Pterocallis, but with two pairs of anterior prothoracic marginal hairs and a small hair arising from the siphunculus, on the ventral side near its base (Richards 1965).

Appendiseta robiniae (Gillette) Plate 9a Viviparae all alate, pale yellow-green with spinopleural and marginal longitudinal rows of pale powdery spots (see aphidtrek.org); BL 1.6-1.9 mm. On undersides of leaves of Robinia pseudacacia and R. neomexicana, and also now recorded from Sophora japonica (Forbes & Chan 1989). Sexuales (alate males and apterous oviparae) occur in September-November (Palmer 1952). Widespread in North America, and introduced into Chile and Argentina (Pagnone et al. 1993), Europe (England, Netherlands, Germany, Poland, Belarus, western Russia, Hungary, Italy, Greece, Spain, Switzerland) and the Middle East (Iran, Jordan: BMNH collection). Ecological studies have been conducted in Poland (Borowiak-Sobkowiak & Durak 2012). The male genitalia were described and illustrated by Wieczorek et al. (2011). 2n=10*.

Apulicallis Barbagallo & Patti

Calaphidinae: Myzocallidini

A genus for one species in Italy related to Myzocallis, but with one pair of spinal hairs on each abdominal tergite in all morphs and other differences in larval chaetotaxy.

Apulicallis trojanae Barbagallo & Patti Alatae are lemon yellow with head and thorax very slightly brownish; BL 1.3-2.3 mm. Living all-year-round on Quercus trojana in south-east Italy. Oviparae and alate males occur in November. Barbagallo & Patti (1991) described all morphs.

Artemisaphis Knowlton & Roberts

Aphidinae: Macrosiphini

A genus erected for a North American Artemisia-feeding aphid with low antennal tubercles, a ventral protruberance on the anal plate and a short, distinctively-shaped cauda. Possibly it belongs in Obtusicauda, as the distinction is blurred by a species described from China (longicauda Zhang) which has a similar anal plate but a longer cauda more typical of Obtusicauda (see under Obtusicauda moldavica). Metopeurum also has a similar development of the anal plate, a well as low antennal tubercles, but has small spiracles like Macrosiphoniella, whereas Obtusicauda and Artemisaphis have large rounded spiracles.

Artemisaphis artemisicola (Williams) (Fig.13b,c) Apterae are shiny dark reddish wine-coloured, BL 1.7-2.0 mm. Forming large colonies on leaves and stems of Artemisia (and Seriphidium) spp. in western USA (Palmer 1952). There are single records also, possibly of vagrants, on Chrysothamnus sp. and Cirsium vulgare.

Asiphonaphis Wilson & Davis

Aphidinae: Aphidini

A genus for two North American and one east Asian species with different biologies and host associations, related to Rhopalosiphum and/or Aphis but completely lacking siphunculi. Both alatae and apterae have prominent marginal tubercles on abdominal tergites 1-7 (Robinson 1964). The genus is unlikely to be monophyletic.

Asiphonaphis japonica Torikura Apterae are dark brown; BL 1.4-1.8 mm. In small colonies on basal parts of a Carex sp. in Japan. Affinity with North American species placed in this genus is uncertain.

Asiphonaphis pruni Wilson & Davis Apterae are pale yellow to whitish green, sometimes banded with darker green, with darker head and pronotum, in colonies on leaves and distorting the new growth of Prunus spp. (armeniaca, serotina, virginiana) in Canada and USA. Monoecious holocyclic, with apterous males. Robinson (1964) gives an account of this species on P. virginiana in Manitoba.

Asiphonaphis utahensis Knowlton Only alatae are described, collected from Salsola pestifer in Utah, USA . This is possibly not the true host.

Asiphonella Theobald

Eriosomatinae: Fordini

Two species with host alternation between Pistacia, on which they form large, cock’s comb‑like galls, and the roots of Poaceae, especially Cynodon dactylon. The life cycle takes two years. Apterae on grass roots have 3‑faceted eyes which are noticeably protuberant, and well‑developed segmental wax glands secreting a coating of white flocculent wax. The two species are difficult to distinguish on their secondary host plants, although the migrants from Pistacia are readily distinguishable. Both apterae and alatae have 6‑segmented antennae. Accounts are given by Remaudiere & Tao (1957) and A.K. Ghosh (1984).

Asiphonella cynodonti (Das) Host-alternating between Pistacia khinjuk and roots or ground level parts of Cynodon dactylon in Iran, Pakistan and India, and also found galling P. palaestina in Israel (Wool 1984). Galls are formed by the invagination of the leaf alongside the mid-rib to form a large, cock’s comb-like structure (fig. 126B and Davatchi 1958). Their ultrastructure was studied by Álvarez et al. (2016). The emigrant alatae (BL about 1.8-1.9 mm) (fig. 81D) leave through secondary openings in the wall of the gall in September to November. Apterae on grass roots are light green to dull green with a dark head, whole body covered with white wax powder; BL 1.6-2.0 mm. They are found on roots of Cynodon dactylon and probably certain other grasses. The original description (Das, 1918) included biological observations.

Asiphonella dactylonii Theobald Plate 1g Host-alternating in China between Pistacia sinensis and roots and ground level parts of Cynodon dactylon and certain other grasses, with anholocyclic populations on grass roots in other parts of the world (Blackman & Eastop 2000, p.242). Aptera (fundatrix?) on Pistacia is yellow, secreting long wax filaments; BL c. 1.7 mm (W. Zhang et al. 1995, as Chaetogeoica utriculata). Galls are similar to those of A. cynodonti, but mature earlier in the year, with alate emigrants (BL 1.5-1.9 mm) collected in May-June (Remaudiere & Tao 1957; W. Zhang et al. 1995). Apterae on grass roots are pale yellow, yellowish green to green, head darker, whole body covered in bluish‑white wax; BL 1.6-2.0 mm. They are found on roots of Cynodon dactylon and certain other grasses (Oplismenus compositus, Chrysopogon ciliata, and Digitaria decumbens). Anholocyclic populations occur in Egypt, Sudan, Zimbabwe, Philippines, Bermuda, New Mexico, Guyana, Brazil, and Argentina. Blanchard (1944) gave an account of this aphid on Cynodon in Argentina, under the name Paraprociphilus graminis, and Silva (1985) provided a more recent account from Brazil.

Asiphonipponaphis Chen, Sorin & Qiao

Hormaphidinae: Nipponaphidini

A genus for one species in China forming galls on the leaves of Distylium and lacking siphunculi in all known morphs.

Asiphonipponaphis vasigalla Chen, Sorin & Qiao Galls arising on the upper sides of the leaves of Distylium racemosum var. chinense in China (Hunan Province) are green, vasiform, c.3.2 cm long and 1.1 cm wide when mature, splitting at the tip into a flower-like opening. Alatae have 5-segmented antennae with secondary rhinaria distributed ANT III 37-44, IV 16-21,V 6-9, and media of forewing unbranched; they migrate in late April to an unknown secondary host (Chen et al. 2011a).

Aspidaphis Gillette

Aphidinae: Macrosiphini

Two species with 5-segmented antennae, reduced siphunculi (in both apterae and alatae), very short hairs, and abdominal tergite 8 produced into a large cowl-like supracaudal process. Immatures have spinulose hind tibiae.

Aspidaphis adjuvans (Walker) Plate 14i (Fig. 42o,p) Apterae are yellowish, brownish yellow or pale bluish green (see influentialpoints.com/Gallery); BL 1.3-2.0 mm. Living cryptically on Polygonum aviculare, often at roadsides or on paths. In Europe, south-west and central Asia, Pakistan (Naumann-Etienne & Remaudière 1995), and North America. Monoecious holocyclic with apterous males (Tuatay & Remaudière 1964). 2n=12 (Canada, Robinson & Chen 1969), 14 and 16 (Israel, Iran, Cyprus; significance of karyotype variaton is unknown). Polygonaphis avicularisucta Zhang, Chen, Zhong & Li is a synonym (G. Zhang & Qiao 2002; Eastop & Blackman 2005).

Aspidaphis porosiphon Börner Apterae are dull, pale green; BL c.1.7-1.9 mm. Living singly on leaf blades of Festuca rubra growing in lush herbage. Europe (UK, Sweden, Germany, Czech Republic). Monoecious holocyclic with oviparae and apterous males in October (Stroyan 1966).

Aspidophorodon Verma

Aphidinae: Macrosiphini

A genus with four or five species on Salix in east Asia (subgenus Aspidophorodon), and four species associated with Rosaceae, three in Asia and one in North America (subgenus Eoessigia). Life cycles of most species are unknown; one species in north-west India host-alternates between Cotoneaster and Potentilla, and one in China possibly migrates from Salix to Polygonum. The antennae are 4- or 5-segmented with terminal process 0.6-1.6 times base of last segment, and the siphunculi have clavate ends with the aperture small and subapical. The head of the aptera bears anteriorly-directed medial and lateral frontal projections, and in subgenus Eoessigia there are variably developed spinal processes. Stekolshchikov & Novgorodova (2010) provided a key to apterae. J. Chen et al. (2015) reviewed the genus, described three new species (one with host unknown) and established some synonymies.

Aspidophorodon cornuatus Qiao Apterae are yellow; BL 1.2-1.5 mm. On Salix cupularis in Tibet, mostly feeding on undersides of leaves (J. Chen et al. 2015). Alatae and life cycle are unknown.

Aspidophorodon harvensis Verma (figs 101A,C) Apterae mainly pale yellowish with pale brown head; BL 1.5-1.9 mm. Alatae have black head and thorax and brownish transverse bars on dorsal abdomen. On undersides of leaves of an unidentified Salix species, sitting tightly alongside the veins, in Kashmir, India (Verma 1965). The life cycle is unknown.

Aspidophorodon (Eoessigia) indica (David, Rajasingh & Narayanan) (Fig.24c-e) Apterae are narrow-bodied, pale green or pale brown with a dark green streak; BL 1.2-2.1 mm. In spring on Cotoneaster obtusus (= bacillaris), feeding along the main veins on upper sides of young leaves; heteroecious holocyclic, migrating in April-May to Potentilla sp. (Chakrabarti & Banerjee 1993a). In north-west India (Himachal Pradesh, Uttar Pradesh).

Aspidophorodon (Eoessigia) longicauda (Richards) Apterae are milky white, lightly pulverulent; BL 1.7-2.0 mm. Alatae have a dusky/dark head and thorax and milky white abdomen with two longitudinal pigmented stripes. On undersides of leaves of Spiraea spp. in British Columbia, Canada, and also now recorded from Washington, USA (aphidtrek.org). Life cycle is unknown. 2n=20.

Aspidophorodon (Eoessigia) longituberculatus (Zhang, Zhong & Zhang) (Fig.42a) Apterae are unknown, alatae have a black head and thorax and whitish abdomen with a dark central patch; BL c. 2.1 mm. On Polygonum sp. in China. This species was originally described in a separate genus, Margituberculatus, and transferred to Eoessigia by J. Chen et al. (2015), who provided a redescription.

Aspidophorodon obtusus Qiao Apterae are green; BL c.1.8-1.9 mm. On undersides of leaves of an unidentified Salix sp. in Sichuan Province, China. Fundatrices were collected along with apterae in May (J. Chen et al. 2015). The life cycle is unknown.

Aspidophorodon salicis Miyazaki (fig. 101B) Apterae are whitish, pale yellow or pale yellowish-green, BL 1.4-1.7mm in spring; dwarf summer apterae have BL 0.9-1.4 mm and 4-segmented antennae. Alatae have a black head and thorax and a large dark central abdominal patch. On undersides of leaves of Salix udensis in Hokkaido, Japan (Miyazaki 1971), and also recorded from Sakhalin and the Kuril islands (Pashchenko 1988a), and on S. viminalis in Kazakhstan (Kadyrbekov 2012a). [Also in China, as two nominal species described from China, Aspidophorodon sinisalicis Zhang and Trichosiphonaphis lijiangensis Zhang, Zhong & Zhang – the latter collected on Polygonum, which might be the secondary host – have been synonymized with A. salicis by J. Chen et al. (2015).]

Aspidophorodon (Eoessigia) sorbi (Chakrabarti & Maity) Apterae are pale brown (?); BL 2.5-2.7 mm. On undersides of young leaves of Sorbus foliolosa in Uttar Pradesh, India (Chakrabarti & Maity 1984, as Indotuberaphis sorbi). Biology and other morphs are unknown. Later generations may have smaller spinal processes. Indotuberaphis was recently synonymised with Aspidophorodon (Eoessigia) by Stekolshchikov & Novgorodova (2010).

Aspidophorodon (Eoessigia) vera Stekolshchikov & Novgorodova Apterae are pale green, without any waxy pulverulence, with pale siphunculi and cauda; BL 1.5-2.0 mm. Alatae are unknown. On Potentilla (= Pentaphylloides) fructicosa, feeding on undersides of leaves along the margins. Only known from south-east of Altai Republic, Russia (Stekolshchikov & Novgorodova 2010).

Astegopteryx Karsh

Hormaphidinae: Cerataphidini

An oriental genus of about 24 species with host alternation between Styrax and monocotyledonous plants, chiefly palms and bamboos. The galls on Styrax vary greatly in morphology between species and are sometimes very elaborate (fig. 124). The morphs from these galls differ greatly from those on secondary host plants, which were described in different genera (Trichoregma, Oregma) until Hille Ris Lambers (1953) suggested that Astegopteryx styracophila on Styrax benzoin might be the same species as Oregma pallida van der Goot on bamboos (but see below under pallida). In more recent years studies of gall structure and symbiont type (Fukatsu et al. 1994), analysis of molecular data (Stern et al. 1997) and experimental host transfers (Kurosu et al. 1998) have led to the elucidation of the life cycles of more species, and as a consequence of these studies some species treated under Astegopteryx in Blackman & Eastop (1994) have been synonymised, and some have been moved to other genera. Apterae of Astegopteryx on bamboos and palms are small aphids with well-developed frontal horns and segmentally arranged wax glands, forming large colonies on the leaves. Most species probably have the ability to maintain themselves continuously by parthenogenetic reproduction on their secondary host plants. Noordam (1991) reviewed the Javanese species, describing the available morphs in considerable detail. Qiao et al. (2018) reviewed the species in China. Aoki & Kurosu (2010) reviewed life cycles and gall formation.

Astegopteryx bambusae (Buckton) (= Oregma bambucifoliae Takahashi; synonymy by Li et al. 2019) Host-alternation occurs in Taiwan between Styrax suberifolium and various bamboos. The generations galling Styrax were described as Astegopteryx sasakii (Takahashi 1936), but were linked with the bamboo-feeding generations by Aoki & Kurosu (1989, as Astegopteryx bambucifoliae). The galls comprise about 20 elongate subgalls radiating from one point on the stem (fig. 124A). Kurosu & Aoki (1990a) reported “chimeral” galls formed initially by A. bambusae, but transformed by invading Ceratoglyphina bambusae. Defensive behaviour patterns involving use of the frontal horns for butting are described by Aoki & Kurosu (1985, 1989, as A. bambucifoliae).

Alatae (BL about 1.7 mm) emerge from distal openings in the subgalls over an extended period from August-December to fly to bamboos (Kurosu & Aoki 1991b, Aoki & Kurosu 2010, as A. bambucifoliae). Apterae on bamboos are broadly pear-shaped aphids, BL 1.2-2.1 mm, variably coloured and patterned in life, with rounded apices to their frontal horns, generally colonising the undersides of the leaves of Bambusa and other bamboos. They form spaced-out colonies on undersides of leaves of Bambusa spp., Dendrocalamus latiflorus and Phyllostachys lithophia. It is also recorded from Phyllostachys edulis in China (Guizhou; Huang et al. 2012, Qiao et al. 2018, as A. bambucifoliae).

A. bambusae occurs in the Indian subcontinent and throughout east and south-east Asia. Populations in different localities may show consistent differences in colour and wax gland development, but these characters may be influenced by climate and other environmental factors. The name A. bambucifoliae has been widely used both for the Styrax-galling generations in Taiwan and for populations of yellowish-green apterae on bamboos with two dark green longitudinal stripes and wax tufts on the sides of the abdomen (see aphids of Karnataka website). However, Li et al. (2019) analysed molecular data from mitochondrial and nuclear genes an extensive collection of specimens to show that this name should be treated as a synonym of A. bambusae.

C. bambusae was originally described from Uttar Pradesh, India, colonising the upper sides of leaves of Bambusa arundinacea (Buckton 1893, as Oregma). Buckton recorded the colour as greenish brown, mottled with black, but he was presumably looking at dead specimens. Doncaster (1953) redescribed the Buckton material and made Oregma lutescens van der Goot from Java a synonym. According to van der Goot’s original description, lutescens is yellow-brown with indistinct dorsal transverse bands on abdominal tergite 1 and between the siphunculi. Noordam (1991) provided a detailed description and illustrations of A. bambusae populations in Java and placed similis van der Goot 1917 and striata van der Goot 1917 as additional synonyms of bambusae, regarding them, with lutescens, as colour varieties (see Noordam 1991 plates 1-5). 2n=12 (Kar et al. 1990).

Astegopteryx basalis van der Goot The name applied to bamboo-feeding populations in south-east Asia, which are similar to A. bambusae, but have broadly conical pointed frontal horns. The primary host of this species has now been shown to be Styrax benzoin (Stern et al. 1997), the galls resembling “gall no. 1204” of Docters van Leeuwen-Reijnvaan & Docters van Leeuwen (1926). A. basalis was first described from unidentified Bambusa species in Java, as a yellowish-white aphid, BL 1.1-1.7 mm, with paired brownish red areas on the dorsum and with wax secreted especially on the sides of the abdomen (van der Goot 1917). Noordam (1991) described other colour forms in Java, and placed three Takahashi species in synonymy: vandermeermohri (Takahashi 1935, nec. Hille Ris Lambers 1931a) from Sumatra; flava (Takahashi 1950), apterae of which are entirely yellow with paired sclerotic patches on abdominal tergites 1-4 and transverse bars on 5-7; and malaccensis Takahashi 1950). A. malaccensis has however subsequently been re-established as a distinct taxon (see below).

Astegopteryx chinensis Tao Forms globular galls similar to those of Ceratoglyphina bambusae on an unidentified species of Styrax in Szechuan, China (Tao 1966). Alatae (BL about 1.8 mm) are mature in the galls in July, and fly to an unknown secondary host.

Astegopteryx formosana (Takahashi) Apterae very broad-bodied, pale yellow or greenish yellow, with a very extensive dark bluish green dorsal patch, broadest anteriorly, and a small marginal fringe of white wax (see aphids of Karnataka website); BL 1.0-1.9 mm (Noordam 1991, as A. unimaculata). On lower sides of leaves of bamboos (Bambusa, Dendrocalamus, Gigantochloa in India, China, Taiwan, Indonesia and Malaysia. In Taiwan this species seems almost to be restricted to Dendrocalamus latiflorus (Liao 1976; as A. insularis), although Aoki & Kurosu (1995) did find a few colonies in Taiwan on Bambusa oldhami. Mounted specimens are difficult to distinguish from A. bambusae, but in life the broad abdomen and dark green patch are distinctive. Aoki & Kurosu (1995) clarified the name of this species, although there remains some doubt about whether the Taiwanese and Javanese populations are conspecific. 2n=12 (Chen & Zhang 1985; as A. insularis).

Astegopteryx glandulosa Noordam Apterae are yellow, with orange markings or marbling, wholly covered with wax wool which extends as a wax powder onto the leaf; BL 1.2-1.5 mm. Immatures are bright yellow, developing orange spots. On undersides of leaves of unidentified bamboos in Java, at altitudes above 1,100 m (Noordam 1991).

Astegopteryx lambersi Takahashi Galls are about 2.7 cm long, approximately kidney-shaped, dark brown and hard when dry, with a large circular distal opening (fig. 124C); on twigs of Styrax benzoin in Sumatra (Takahashi 1936). Alatae, BL about 1.75 mm, are adult in February. The secondary host is unknown.

Astegopteryx liukueinensis Liao Apterae are yellow in life with a pair of longitudinal green markings, BL 1.4-1.6 mm, and occur on Bambusa multiplex in Taiwan (Liao 1976).

Astegopteryx malaccensis (Takahashi) Apterae on bamboos in peninsular Malaysia are dark purplish brown form with very well-developed wax glands. The gall on Styrax benzoin is similar to that illustrated as “gall no. 1203” in Docters van Leeuwen-Reijnvaan & Docters van Leeuwen (1926). The life cycle was established by analysis of molecular data (Stern et al. 1997) and a transfer experiment (Aoki & Kurosu, unpubl. data).

Astegopteryx minuta (van der Goot) is the name most commonly applied to populations of bambusae-like aphids with marginal wax pore plates reduced (so that they are small, circular and discontiguous) or totally absent. In life such apterae are generally yellowish white to pale green with two irregular dark green longitudinal markings and scanty wax; BL 1.1-1.8 mm (Noordam 1991). On undersides of leaves of bamboos (Bambusa, Dendrocalamus) in India, China (Qiao et al. 2018) and south-east Asia. Populations in which the apterae have no visible marginal wax pore plates and are not clearly distinguishable from A. minuta have been described from India as new species: A. neelagiriensis David 1959, on Bambusa sp. in Madras, and Pseudastegopteryx himalayensis M.R. Ghosh, Pal & Raychaudhuri 1977, on unidentified bamboo in West Bengal. Alatae occur (in Java) from May to mid-October (Noordam 1991). 2n=12 (in India; Kar et al. 1990).

Astegopteryx nipae (van der Goot) Apterae on palm leaves are small, with yellow-grey head and thorax, bright brownish red abdomen and a broad white wax fringe; BL 0.9-1.6 mm. They can occur in very large, invariably ant-attended colonies on various palms (Calamus, Cocos, Daemonorops, Elaeis, Nypa, Salacca), especially those with fan-like or climbing growth habits. In Malaysia, Indonesia, Solomon Islands, New Hebrides, Fiji, and Tonga. The primary host is reported to be Styrax benzoin (Kurusu & Aoki unpubl., in Aoki et al. 2002).

Astegopteryx pallida van der Goot Apterae greenish white or yellow with variable dark green dorsal markings and white wax, especially on margins; BL 1.1-2.2 mm (Noordam 1991). On undersides of leaves of bamboos (Bambusa, Gigantochloa) in Indonesia. Oregma salatigensis van der Goot (1917) is a synonym, and possibly also Trichoregma bambusae Takahashi (1935). It has also now been shown that A. setigera Noordam, which produces flower-bud galls on Styrax benzoin in Java (Noordam 1991) and Sumatra, is the primary host form of A. pallida (Kurosu et al. 1998).

Astegopteryx pandani (Takahashi) Apterae pale brown to brown with whitish antennae and legs and long marginal tufts of white wax; BL c. 1.4 mm. On Pandanus tectorius in Sumatra (Takahashi 1935), and redescribed from Freycinetia javanica in Java (Noordam 1991). On undersides of leaves, attended by ants. Other morphs and life cycle unknown.

Astegopteryx rappardi Hille Ris Lambers Apterae are small, with pale grey-brown head and prothorax and appendages, and greyish brown to brownish red or dark violet abdomen, fringed with plates of wax; BL 1.2-1.5 mm. In large numbers on undersides of palm leaves (Cocos nucifera, Elaeis guineensis). Infested leaves become spotted with yellow. esides the key characters, Hille Ris Lambers (1953b) and Noordam (1991) gave other features distinguishing this species from A. nipae. Alatae of A. rappardi are black, whereas those of A. nipae have a greenish abdomen. Recorded from Malaysia, Indonesia and the Solomon Islands.

Astegopteryx rhapidis (van der Goot) Apterae are yellowish brown to dark brown, with a wax fringe; BL 1.2-2.0 mm. Forming numerous ant-attended colonies on undersides of leaves of palms (Cocos, Elaeis, Rhapis javanica, Calamus). In Taiwan, Philippines, Palau Islands, Malaysia, Indonesia, and the Solomon Islands. Evans et al. (1995) recorded a parasitoid, Encarsia noordami.

Astegopteryx singaporensis (van der Goot) Apterae are yellowish white or pale brownish yellow, with 1-3 pairs of small pale greenish spots, a somewhat shiny, bright yellow to brownish area on and anterior to the siphunculi, and variably developed marginal plumes of wax; BL 1.2-1.7 mm. On undersides of old leaves of bamboos, which often show yellow or brown patches. Originally described from unidentified bamboos in Singapore (Van der Goot 1918), and redescribed from Java, where it is recorded from species of Bambusa, Dendrocalamus, Melocanna and Schizostachyum (Noordam 1991). Alatae are unknown.

Astegopteryx spinocephala Kurosu, Buranapanichpan & Aoki Galls on Styrax benzoides in northern Thailand are complex multi-chamber structures shaped like bunches of elongate bananas (Kurosu et al. 2006a). They take about a year to mature and the alatae produced in March to April are sexuparae, so this species completes its life cycle without migrating to secondary hosts. Sexual morphs and eggs were found in subgalls guarded by soldiers that plugged the openings of the galls with their sclerotised spiny heads (Kurosu et al. 2006b).

Astegopteryx styracophila Karsh Galls on Styrax benzoin in Java and Sumatra are elaborate multi-chamber structures formed from flower-buds and are similar to those of A. bambucifoliae in Taiwan. Alatae (BL 1.3-1.7 mm, with secondary rhinaria distributed ANT III 10-22, IV 6-14, V 4-9) emerge through a circular opening in Feb-June. Host alternation occurs to gingers (Zingiberaceae) the secondary host forms being previously known as A. muiri (van der Goot). In transfer experiments, Kurosu et al. (1998) found that progeny of the emigrant alatae colonised both a palm (Areca catechu) and Etlingera elatior (Zingiberaceae), and developed into adult apterae with the morphology of A. muiri, the only species of the genus recorded from Zingiberaceae. A. muiri has not been recorded from palms, which may be due either to confusion with the similar species A. nipae, or to palms not being colonised under field conditions. Apterae on secondary hosts are green or brown, sometimes with a reddish or violet tinge, with wax secretion; BL 0.9-1.3 mm. They live in ant-attended colonies on undersides of leaves of Zingiberaceae (Alpinia, Amomum, Languas, Zingiber). Populations on Zingiber sp. in Vietnam and on a species of Zingiberaceae in China (Hong Kong) have also been identified as A. styracophila (Yoshitomi et al. 2015, Qiao et al. 2018). Records of an Astegopteryx sp. from Musa sapientum are also likely to be this aphid, but the close similarity to the palm-feeding species A. nipae (van der Goot), makes definite identification difficult.

Atarsos Gillette

Aphidinae: Macrosiphini

One nearctic species on Compositae/Asteraceae that completely lacks tarsi. (Note: RIV has 10-20 accessory hairs, more than illustrated by Palmer 1952).

Atarsos grindeliae Gillette Apterae are shining pale yellow to pale green with pale to dusky appendages; BL 1.3-1.8 mm. On leaves of Grindelia squarrosa and plants in related composite genera (Aster, Dieteria, Machaeranthera). Described from Coloradoa, USA, and also found in Canada (Manitoba). Oviparae and apterous males occur in September-October (Palmer 1952). 2n=12.

Atheroides Haliday

Chaitophorinae: Siphini

Six palaearctic and one nearctic species of very elongate-bodied grass-feeding aphids, with short antennae and legs, and a sclerotic tergum with fused head and pronotum and often fused abdominal tergites 2-7. Cauda and anal plate are broadly rounded, with cauda usually obscured in dorsal view by abdominal tergite 8, the hairs on which are very long and thick. Alatae are rare and sexual morphs of several species are unknown. Wieczorek (2009) fully reviewed the genus, with a key and illustrations of apterae of all six species, including a new one (A. persianus Wieczorek) from an unknown host in south-west Asia, collected by sweeping Cyperaceae. Wieczorek (2010) provided detailed distribution maps, redescriptions and keys to all known morphs.

Atheroides brevicornis Laing Apterae are brownish to black, very narrow-bodied (see influentialpoints.com/Gallery); BL 1.5-2.4 mm. In rows on upper sides of leaves of grasses (Deschampsia, Puccinellia, Festuca), mainly in coastal habitats around northern Europe and the Black Sea (UK, Sweden, Norway, Netherlands, northern France, Germany, Slovakia, Czech Republic, Moldova, Ukraine), and also found in a salty pond-side habitat in Hungary (Pintera 1965b, as A. aplangi). Records from Carex and Juncus perhaps need confirmation as these aphids are often collected by sweeping from mixed vegetation. Oviparae were recorded as occurring in early October by Pintera (1965, as aplangi), and were redescribed by Wieczorek (2011). Males are undescribed. See Wieczorek (2010) for other information.

Atheroides doncasteri Ossiannilsson Apterae are dirty yellow to brownish; BL 1.7-2.6 mm. Only known from Deschampsia caespitosa. In Europe (Netherlands, Sweden, Slovakia, Hungary, Poland, west Siberia). An ovipara and males collected in Sweden in September-October were described by Wieczorek (2010).

Atheroides hirtellus Haliday Apterae are dark brown to black with ventral side yellowish; BL 2.0-2.8 mm. In small colonies in grooves of leaf blades of grasses such as Deschampsia caespitosa and Arrhenatherum elatius, not attended by ants. In Europe (UK, Ireland, France, Netherlands, Sweden, Finland, Germany, Poland, Latvia, Russia, Switzerland, Spain), and also recorded from Kazakhstan (Kadyrbekov 2017a), and from Jilin province, China (Tao 1999). Juncus and Scirpus spp. are also recorded hosts, but these specimens may have been swept from mixed vegetation. Monoecious holocyclic, with oviparae in September (in Scotland). Males are apterous, dark brown to black. For fuller information see Wieczorek (2010). 2n=8.

Atheroides karakumi Mordvilko Apterae are yellowish brown; BL c. 2.3 mm. In grooves of leaf blades of Stipa (= Lasiagrostis) splendens and Festuca ovina. In Caucasus and Central Asia. Apterae and alatae were redescribed by Wieczorek (2010). Sexual morphs are unknown.

Atheroides serrulatus Haliday Plate 5b Apterae are yellow or yellowish brown (see influentialpoints.com/Gallery); BL 1.7-2.4 mm. On leaves of various grasses (Agrostis, Alopecurus, Brachypodium, Bromus, Calamagrostis, Cynosurus, Dactylis, Deschampsia, Festuca, Holcus, Hordeum, Nardus, Phalaris, Poa). Also recorded quite frequently from Cyperaceae (Carex, Juncus), although this needs confirmation because the specimens may have been collected by sweeping or beating mixed vegetation. Widespread in Europe; also recorded from west Siberia, Iran, Turkey, Kazakhstan (Kadyrbekov 2017a), Korea (BMNH collection, leg. W.H. Paik), and Canada (Quebec, Richards 1972; British Columbia, Forbes & Chan 1989). Monoecious holocyclic, with oviparae and pale olive to yellowish brown apterous males in late September-November. See Wieczorek (2010) for further information. 2n=8.

Atheroides vallescaldera Miller & Jensen Appearance in life unknown; BL c.2.0-2.2 mm. On an unknown species of Poaceae in southern USA (Valles Caldera National Preserve, New Mexico). Apterae and oviparae were collected in September (G.L. Miller et al. 2014). Other morphs are unknown.

Aulacophoroides Eastop & Hille Ris Lambers

Aphidinae: Macrosiphini

A genus for four species in east Asia, on woody Leguminosae/Fabaceae mainly of climbing habit, with a spinulose, Aulacorthum-like head but also with characters similar to Megoura. (The generic classification of the oriental legume-feeding Macrosiphini needs to be clarified). Qiao et al. (2006b) reviewed and keyed the species.

Aulacophoroides formosana (Takahashi) Apterae are green with yellowish-brown head, black antennae and siphunculi, yellowish-brown legs except for black tarsi, and a yellow cauda; BL c. 2.7 mm. Recorded only from Millettia reticulata in Taiwan (Takahashi 1923) and Chekiang, China (Tao 1963).

Aulacophoroides hoffmanni (Takahashi) Apterae are dark red with black antennae and siphunculi; BL c.2.5-2.7 mm. On Wisteria chinensis (= sinensis) in China. The alata was described by Tao (1963, as Subacyrthosiphon). Monoecious holocyclic; oviparae were collected in October (Takahashi, 1937, as Acyrthosiphon). 2n=14*.

Aulacophoroides millettiae Qiao, Jiang & Martin Apterae are dark brown to black; BL 2.2-2.9 mm. On Millettia sp. (not a tree but a woody vine) in Hong Kong, China (Qiao et al. 2006b).

Aulacophoroides virgatae Szelegiewicz Apterae are shining green with black antennae and siphunculi; BL 1.9-2.4 mm. On terminal shoots of Lespedeza virgata in Korea.

Aulacorthum Mordvilko

Aphidinae: Macrosiphini

About 40 species of Aulacorthum are known, the majority from the eastern palaearctic and oriental regions. The remaining species are western palaearctic; the few described from North America seem to belong to other genera, e.g. Ericaphis. Members of the genus are recognisable by the parallel‑sided inner faces of the spinulose antennal tubercles. W. Lee & S. Lee (2013) studied the distinction between Aulacorthum and Acyrthosiphon using molecular and morphometric methods. Neomyzus has hitherto been treated as a subgenus of Aulacorthum, but DNA studies indicate that they are not so closely related. None of the species have host alternation. A. solani is one of the most polyphagous aphids. There is a similar lack of pattern in the hosts of the plant‑specific species. Eight species of Aulacorthum are described from Compositae, 4 from Liliaceae, 3 each from Ericaceae and Lamiaceae, and 2 from Dipsacaceae, Lauraceae, Oleaceae, Rosaceae, and Rutaceae. Accounts are available for western Europe (Hille Ris Lambers 1947b, 1949, Heie 1994, Blackman 2010), Japan (Miyazaki 1971, in Acyrthosiphon), Korea (W. Lee et al. 2009b, W. Lee et al. 2012a), China (Su & Qiao 2011) and lndia (Raychaudhuri 1980), Java (Noordam 2004), and Canada (Richards 1972b).

Aulacorthum aegopodii Börner Apterae are opaque creamy white with yellow spots at bases of siphunculi; BL 2.0-2.4 mm. Alatae have 4-8 secondary rhinaria on ANT III (cf. A. solani). On Aegopodium podagraria in Europe (Netherlands, Germany; as A. solani ssp. aegopodii). Monoecious holocyclic, with alate males; it did not hybridise readily with A. solani s.str. in laboratory experiments (Müller 1976).

Aulacorthum albimagnoliae Lee Apterae are milky white with only tips of siphunculi dark; BL 1.8-2.4 mm. On undersides of leaves, young shoots and young flowers of Magnolia spp. in Korea (S. Lee & Kwon 2006, W. Lee et al. 2009b). Alatae are much darker with black head and antennae, dark brown to black thorax and extensive dark dorsal abdominal sclerotisation. The species is believed to live all year on Magnolia. Sexuales have not yet been observed.

Aulacorthum artemisiphaga Lee, Havelka & Lee Apterae are pale green with dark tips to antennae, legs and siphunculi; BL 2.2-3.0 mm. Alatae are much darker with black head, antennae and thorax and extensive dark dorsal abdominal sclerotisation including dark intersegmental muscle plates. On Artemisia spp. (princeps, stolonifera) in Korea (W. Lee et al. 2011b). Other morphs and life cycle are unknown.

Aulacorthum asteriphagum Lee, Kim & Lee Colour of apterae in life unrecorded; BL 2.1-2.6 mm. On undersides of leaves of Aster scaber (= Doellingeria scabra) in Korea (W. Lee et al. 2009a). Other morphs and life cycle unknown.

Aulacorthum asteris Takahashi Apterae are white or pale yellowish, with dusky sensoriated part of ANT III, dark apices to antennal segments and femora, and siphunculi pale in middle, dark at base and apex; BL 2.5-3.5 mm. On Aster (Kalimeris) yomena in Japan, and an unidentified Aster sp. in Korea (W. Lee et al. 2002c).

Aulacorthum cercidiphylli (Matsumura) Apterae are yellow to pale green with BL 1.8-2.1 mm in spring, but down to 1.1-1.2 mm in summer. Feeding singly on leaves of Cercidiphyllum japonicum in Japan (Honshu, Hokkaido). Presumably monoecious, holocyclic. The alata was described by Shinji (1930), and the fundatrix by Miyazaki (1971). 2n=12.

Aulacorthum cirsicola (Takahashi) Apterae are large, broadly spindle-shaped pale lemon-yellow, pink or green aphids with antennae and legs banded with black, and black-tipped siphunculi; BL 2.5-4.1 mm. Monoecious holocyclic, feeding on upper leaves and stems of Cirsium spp., and also recorded from certain other Compositae/Asteraceae (Arctium, Petasites, Tussilago). In Japan, China, Korea, Taiwan and east Siberia. 2n=10.

Aulacorthum corydalicola Lee, Kim & Lee Apterae are pale green with antennae and distal parts of femora, tibiae and siphunculi dusky/dark; BL 2.2-2.6 mm. Alatae have dark antennae, head and thorax, dark cross-bands on dorsal abdomen and 1-7 secondary rhinaria on ANT III. On young leaves, flowers and seed pods of Corydalis spp. in Korea (W. Lee et al. 2009a). The life cycle is unknown.

Aulacorthum cylactis Börner Apterae are shining green; BL. 1.4-1.9 mm. Alatae have dark head and thorax and rather pale brown dorsal abdominal cross-bands. On undersides of rolled leaves of Rubus saxatilis in Europe (Denmark, Sweden, Finland, Germany), west Siberia (Stekolshchikov et al. 2008a) and Kazakhstan (Kadyrbekov 2013d), and recorded from R. nessensis in Belarus (Buga & Stekolshchikov 2012). Monoecious holocyclic with apterous males (Ossiannilsson 1959). Sometimes treated as a subspecies of A. solani.

Aulacorthum dasi A.K. Ghosh, R.C. Basu & Raychaudhuri Apterae are chocolate brown, with blackish brown head, basal parts of siphunculi, and cauda: BL c.2-7-2.9 mm. Alatae have dark bands across the dorsal abdomen. Described from an unidentified composite plant in India, with a later record from Artemisia sp. (Raychaudhuri, 1984).

Aulacorthum dorsatum Richards Apterae are yellowish green or lemon yellow with dark antennae and variably developed black markings, maximally consisting of broad cross-bands on pro- and mesonotum and a broad, brown/black median stripe extending from metanotum to abdominal tergite VI, joined to a broad band between the black siphunculi (C.-k. Chan, pers. comm.); BL 1.9-2.4 mm. Alatae have a black dorsal abdominal patch. On viscous racemes of Gaultheria shallon, sometimes on the terminal shoot and occasionally producing immatures on lower surface of mature leaves. Monoecious holocyclic (C.-k. Chan, pers. comm.).In western North America (British Columbia, Washington, Oregon, California). DNA barcoding (Foottit et al. 2008) indicates that this species might be more correctly placed in Ericaphis. 2n=12*.

Aulacorthum esakii (Takahashi) Apterae are shining black; BL c. 2.0-2.2 mm. Alatae are undescribed. On Paederia foetida in Japan.

Aulacorthum euphorbophagum Zhang, Chen, Zhong & Li Apterae are yellow; BL c. 1.9 mm. Alatae have dark bars joined between abdominal tergites 3-6 in midline. On undersides of leaves of Euphorbia pekinensis in north-west China (Zhang 1999). Closely related to A. solani, but with longer HT II.

Aulacorthum flavum Müller Apterae are shining yellow or greenish yellow, with yellowish brown spots at bases of siphunculi; BL 1.5-2.1 mm. On leaves and shoot apices of Vaccinium spp., especially V. uliginosum. Rupais (1989) recorded it from Andromeda polifolia. In northern and central Europe. Monoecious holocyclic, with oviparae and alate males on V. uliginosum in late August-September in Finland (Heikinheimo 1990). 2n=12*.

Aulacorthum glechomae Takahashi Apterae are white; BL c. 1.5 mm. Alatae have extensive dark dorsal markings. On Glechoma hederacea (var. grandis) in Japan and Korea, feeding solitarily on undersides of leaves (Miyazaki 1971, as Acyrthosiphon). Also recorded in Korea from Mukdenia (Aceriphyllum) rossii (S. Lee et al. 2002c). Immatures have spinulose tibiae suggesting that this species may really belong in Neomyzus.

Aulacorthum ibotum Essig & Kuwana Apterae are yellow to pale green with dark antennae, yellowish legs except for black tarsi, black siphunculi and a pale cauda (Essig & Kuwana 1918); BL 1.8-2.3 mm. On undersides of leaves of Ligustrum spp. in Japan (Hokkaido, Honshu) and Korea (S. Lee et al. 2002c). The life cycle has not been investigated. 2n=14.

Aulacorthum ixeridis Lee, Havelka & Lee Colour of apterae in life is unrecorded, presumably rather dark; BL 2.2-2.9 mm. Alatae have dark head and thorax and dark dorsal abdominal cross-bands. On Ixeris chinensis var. graminifolia (= Ixeridium graminifolium) and Lactuca indica in North Korea (W. Lee et al 2009b). The life cycle is unstudied.

Aulacorthum kerriae (Shinji) Apterae are black except for yellow central area on dorsal abdomen, basal part of ANT III, basal parts of femora, tibiae and cauda, and middle parts of siphunculi (Shinji 1930); BL c. 2.0-2.3 mm. On Kerria japonica in Japan, curling young leaves at growing points in late spring. Also recorded from Petasites sp. (Takahashi 1965a). Monoecious holocyclic, with oviparae and alate males in late September-October (original description). [The aphid described and illustrated by Moritsu (1983) as Acyrthosiphon kerriae (Shinji) seems to be Aulacorthum solani.]

Aulacorthum knautiae Heie Apterae are yellowish or pale greenish, often shiny, without pigment spots at bases of siphunculi; BL 2.3-2.9 mm. On undersides of leaves of Knautia arvensis in Europe. Monoecious holocyclic, with oviparae and apterous males in September-October (original description).

Aulacorthum kuwanai (Takahashi) Apterae are shining brownish yellow, with blackish apices to segments of antennae and legs, and to siphunculi; BL 1.5-1.8 mm. On Vitis sp. in Taiwan.

Aulacorthum langei (Börner) Apterae are rather evenly pale green, without any darker spots at bases of siphunculi (cf. A. solani); BL 1.9-2.3 mm. Alatae usually have dark dorsal abdominal cross-bands. On Pulmonaria officinalis, feeding on leaves, which become irregularly curled, and petioles. Europe (Germany, Czech Republic, Sweden). Monoecious holocyclic with apterous males (Müller 1976a, as A. solani ssp. langei).

Aulacorthum ligularicola Lee Apterae have a dark reddish brown head, thorax and abdomen milky white or pale yellow with large red patches at bases of dusky siphunculi; BL 2.7-3.4 mm. On undersides of leaves of Ligularia fischeri, causing severe curling. In mountainous regions of South Korea (Lee 2002).

Aulacorthum linderae (Shinji) Apterae are pale green to white or creamy with brownish coloration around bases of siphunculi, antennae and legs banded with black, siphunculi pale with black tips, cauda pale; BL 2.0-2.5 mm. In Japan, on undersides of leaves of Lindera sericea, causing loose leaf-curl and reddish-brown blotches (Miyazaki 1971). Presumably monoecious, holocyclic. [The host was given as Aucuba japonica in the original description (Shinji 1922b), presumably in error.]

Aulacorthum magnoliae see Pseudomegoura magnoliae

Aulacorthum majanthemi Müller Apterae are shining brown; BL c.2.5-2.7 mm. On Maianthemum bifolium, yellowing leaves. Europe (Denmark, Germany, Poland, Russia, Switzerland). Monoecious holocyclic with oviparae and alate (brachypterous) males found in late September in northern Germany (original description).

Aulacorthum muradachi (Shinji) Apterae are milky white, pale yellow to pale green with dark antennal flagellum, bicoloured legs, and conspicuous dorsal pigmentation consisting of paired submarginal rows of red to reddish-brown patches from mesonotum to abdominal tergite 6, converging and broadening out at the bases of the black siphunculi (see Moritsu 1983, as Acyrthosiphon); BL 1.6-2.6 mm. On leaves of Lindera spp. (erythrocarpa, praecox, triloba, umbellata) in Japan (Honshu), and redescribed from L. erythrocarpa in Korea (Lee et al. 2008). Presumably monoecious holocyclic; fundatrices are described from L. triloba (Takahashi 1965, Miyazaki 1971 as Acyrthosiphon). 2n=10.

Aulacorthum myriopteroni Zhang Colour of apterae in life is not recorded; BL c. 2.4 mm. On Myriopteron extensum in Yunnan province, China. Su & Qiao (2011) record an earlier collection from Taiwan, on Paederia. 2n=10 (Chen & Zhang 1985b)

Aulacorthum nepetifolii Miyazaki Apterae are blackish brown; BL 2.5-2.9 mm. On Nepeta subsessilis (incl. var. yesoensis = N. yesoensis). In Japan and South Korea.

Aulacorthum nipponicum see Pseudomegoura nipponica

Aulacorthum palustre Hille Ris Lambers Apterae are pink or pale green, frequently with darker rust-coloured or green spots at bases of siphunculi; BL 2.4-3.2. Immatures have cross-bands of wax powder. On undersides of leaves of liguliflorous Compositae/Asteraceae such as Hypochaeris, Leontodon, Picris and Taraxacum. In Europe, east to Poland and Czech Republic, and south to Italy and Greece. It is also reported Iran (Rezwani 2010) and Saudi Arabia (Hussain et al. 2015b). Males were trapped in UK in October, but possibly it is mainly anholocyclic, as found on underside of rosette leaves of Taraxacum in UK in February (BMNH collection, leg. J.H. Martin). 2n=34*.

Aulacorthum (Perillaphis) perillae (Shinji) Apterae are yellowish brown, rather shining, with black antennae and siphunculi and a pale cauda; BL 1.6-2.0 mm. Alatae have a shiny black thorax and a dark brown dorsal abdominal pattern of sclerotisation. On undersides of leaves of Perilla spp. (frutescens, ocymoides). In east Asia; West Bengal (BMNH collection, leg. A.N. Basu), Japan, Szechuan, Taiwan and Korea. Takahashi (1965a) provided a redescription. Aulacorthum (Perillaphis) siniperillae (Zhang), described from Perilla frutescens in China (as Cryptaphis) is a synonym (Su & Qiao 2011). [Holman (2009) lists numerous other host plants for A. perillae, but these are in error and should all be referred to Aulacorthum (now Pseudomegoura) magnoliae.]

Aulacorthum phytolaccae Miyazaki Apterae have a black sclerotic dorsum, immatures are green; BL 2.0-2.7 mm. Described from Phytolacca esculenta, and specimens of apparently the same species were also found on undersides of leaves of Comanthosphace sp., attended by ants (BMNH collection, R.L. Blackman). Alatae are undescribed. In Japan. 2n=10* (specimens from Comanthosphace).

Aulacorthum pirolacearum Szelegiewicz Apterae are shining orange or dark green, immatures are white or yellowish; BL 1.7-2.1 mm. Alatae have large brown marginal and intersegmental sclerites. On undersides of leaves of various Ericaceae-Pyrolaceae (Moneses, Pyrola, Ramischia) in Poland, and also reported from P. rotundifolia in north-west Russia by Stekolshchikov (2017), who confirmed the constancy of the characters by which it differs from A. rufum. However, specimens subsequently collected from P. grandiflora in west Siberia apparently had more affinity to A. rufum, indicating the need for further work (Stekolshchikov & Khureva 2020).

Aulacorthum pterinigrum Richards Appearance of apterae in life is not recorded, but presumably greenish or yellowish with a mainly brown-black abdomen, laterally as well as dorsally, with pale submarginal longitudinal stripes; BL c.2.2 mm. Host of type material was originally recorded in error as “Pteris”, but was actually Pieris (Ericaceae), and there are subsequent records from Vaccinium spp. and Gaultheria shallon (C.-k. Chan, pers. comm. viii.2005), and also from Akebia quinata (Lardizabalaceae) (Robinson & Chen 1988). Described from British Columbia (Canada), and subsequently found in California and Oregon (aphidtrek.org). Alate males have been reared in the laboratory (C.-k. Chan, pers. comm. x.2005). [This species is listed as a Sitobion in Remaudière & Remaudière (1997), but is transferred back to Aulacorthum here, adopting the narrower concept of Sitobion advocated by Jensen (1997). Its generic position is however uncertain and DNA barcoding (Foottit et al. 2008) indicates that it might be more correctly placed in Ericaphis.]

Aulacorthum rhamni M.R. Ghosh, A.K. Ghosh & Raychaudhuri Apterae are yellowish green with darker siphunculi and cauda; BL c.1.8-2 mm. Described from colonies on apical leaves of Rhamnus nepalensis in West Bengal, India (M.R. Ghosh et al. 1971). . Subsequently recorded from Eurya sp. and other unidentified plants (Raychaudhuri et al 1980). The host plant relationships need clarification. Biology is unknown.

Aulacorthum rufum Hille Ris Lambers Apterae are green or dirty reddish brown, slightly wax-powdered laterally and ventrally; BL 2.0-2.7 mm. On young shoots and undersides of leaves of Vaccinium spp., usually V. myrtillus. In north-west, north and central Europe. Monoecious holocyclic with apterous males.

Aulacorthum scirpi A.K. Ghosh & Raychaudhuri 1972, nec. van der Goot Apterae are dark with black siphunculi; BL 1.7-1.8 mm. On Scleria terrestris (= elata) in Assam, India. Described under the name Aulacorthum scirpi van der Goot, which is now thought to be a species of Carolinaia (Juncomyzus); see Noordam (2004).

Aulacorthum sclerodorsi Kumar & Burkhardt Apterae are pale brown with darker markings on dorsal thorax and abdomen, and black tips to siphunculi; BL c.2.8-2.9 mm. Alatae have a rather extensive black dorsal abdominal patch. On “Nepata” (presumably Nepeta) sp. in India (Himachal Pradesh).

Aulacorthum sedens Müller Apterae in life are whitish, slightly greenish, shiny, with tips of siphunculi rather dark; BL 1.4-2.4 mm. On undersides of lower leaves of Knautia arvensis in Germany and Poland, and also recorded from Italy (van Harten & Coceano 1981). Monoecious holocyclic, with oviparae and alate males in late September (original description).

Aulacorthum sensoriatum David, Narayanan & Rajasingh Apterae are yellow shading to brown on posterior tergum, with contrastingly dark front of head, ANT I and distal parts of femora; BL 2.7-3.1 mm. Alatae have 32-39 small secondary rhinaria on ANT III. Described from an “undetermined creeper”, feeding on underside of leaves; the host was subsequently identified as Vinca major (Dutta & Gautam 1993, and BMNH collection). In Himachal Pradesh, India. Life cycle is unknown.

Aulacorthum smilacis Takahashi Apterae are yellow with variably developed dark markings, antennae and legs banded with black, and black-tipped siphunculi; BL 1.8-1.9 mm. Alatae are undescribed. On Smilax spp. in Japan. 2n=10.

Aulacorthum solani (Kaltenbach) Plate 22i Apterae are shining pale whitish or yellowish green with darker green, orange or rust-coloured spots at bases of siphunculi, to dull green or greenish brown (see influentialpoints.com/Gallery); BL 1.8-3.0 mm. Alatae have dark head and thorax and a variably-developed pattern of dark dorsal abdominal cross-bands, and 8-18 (mostly 9-15) secondary rhinaria on ANT III. Extremely polyphagous, colonising plants in may different families of both dicots and monocots. Piron (2017) provided many new host records from the Netherlands. Probably of European origin, now almost world-wide. Monoecious holocyclic with apterous and (more rarely) alate males, and with the unusual ability to go through the sexual phase on many different plant species. Commonly anholocyclic in mild climates and glasshouses. Populations occur with particular host associations, some of which are persistent. Outbreaks have occurred on soybean in Japan (Nagano et al. 2001, Saito et al. 2001) although it is not reported elsewhere as a pest on this crop. A population with a dark dorsal shield living on maritime Convolvulus near Auckland, New Zealand has been known since 1939 (Cottier 1953); it transferred readily to sprouting potato tubers in 1983 and maintained its dark pigmentation in culture for several generations (V.F. Eastop, unpublished data). A. aegopodii, A. cylactis and A. langei are morphologically similar but biologically distinct taxa in Europe closely related to A. solani. There are also some recognisably distinct permanently anholocyclic forms that are not reliably distinguishable on morphological grounds alone from A. solani, such as A. prasinum Börner, studied in Germany by Müller (1970). Tiwari et al. (2021) have published a preliminary analysis of the draft genome sequence to uncover virulence genes. Miller et al. (2009) found little variation in mitochondrial DNA sequence from different parts of the world. See also Blackman & Eastop (2000). 2n=10.

Aulacorthum speyeri Börner Apterae are shiny black except for pale yellow areas on mesonotum and abdominal tergites 1 and 5, with mainly pale antennae and legs, siphunculi dark at base and apex, and dark cauda; BL 1.9-2.9 mm. On undersides of leaves of Convallaria majalis, Polygonatum multiflorum, and some other Asparagaceae formerly placed in Convallariaceae. Also occasionally found in small numbers in concealed situtations on certain other plants (Anthericum, Lycopus, Potentilla). Infested leaves of Convallaria develop large yellow spots and become desiccated. In north-west Europe, Italy, Hungary, Greece, eastward to Iran, and also now recorded from Argentina (Ortego 1998a). Müller (1979) gave an account of this aphid. Oviparae and alate males occur in October in northern Europe. 2n=10.

Aulacorthum spinacaudatum (Kumar & Burkhardt) Apterae are pale yellow; BL c. 3.0 mm. Alatae are light brown with black head and thorax. Described from Cucumis sativus in India (Himachal Pradesh). Specimens from Cucurbita pepo in West Bengal (BMNH collection, leg. A.N. Basu) and from Sechium edule previously identified as A. magnoliae seem to be this species, and there are other Indian records of A. magnoliae on Cucurbitaceae that should be referred to A. spinacaudatum. A small sample collected in mid-September from Eleusine (BMNH collection, CIE A15689) included an alate male. 2n=12* (Khuda-Bukhsh & Basu 1987, as A. magnoliae).

Aulacorthum syringae (Matsumura) Apterae are pale yellow to yellowish-green, often with reddish-brown coloration around bases of siphunculi; BL c.2 mm. Antennae and legs mainly dark, siphunculi black, cauda pale (Takahashi 1965a). On Syringa spp. (including introductions) in Japan and east Siberia. Presumably monoecious, holocyclic; fundatrices are described from S. reticulata (Miyazaki 1971).

Aulacorthum takahashii (Mason) Apterae are white to pale yellowish, with black tips to antennal segments, tibiae and siphunculi; BL c.2.1 mm.On leaves of Pollia japonica in Taiwan.

Aulacorthum vaccinii Hille Ris Lambers Apterae are shining black, with appendages pale with dark apices, siphunculi sometimes dark; BL 1.3-2.1 mm. On Vaccinium uliginosum in northern Europe and Czech Republic and also recorded from V. japonicum and Leucothoe in Japan (Miyazaki 1971). Monoecious holocyclic on Vaccinium, with oviparae and alate males in August-September in northern Europe (Heikinheimo 1990). One small aptera with very short frontal hairs was given subspecific status in the original description as A. vaccinii ssp. parvulum.

Aulacorthum vandenboschi Hille Ris Lambers Apterae are large, broadly spindle-shaped, orange aphids with black appendages; BL c.3.5 mm. Alatae have brown marginal sclerites and thin dorsal cross-bands. On Cirsium japonicum in Japan and Korea (W. Lee et al. 2012).

Aulacorthum watanabei (Miyazaki) Apterae are yellow to orange-yellow in spring, creamy white in summer, with shining black head and sclerotised parts of thorax and abdomen, mainly pale antennae and legs, dark siphunculi and pale yellow cauda; BL c.3.0 mm. On undersides of leaves of Convallaria keiskei in Japan (Miyazaki 1971).

Avicennina Narzikulov

Aphidinae: Macrosiphini

A genus with three rather large Asian species on Lonicera with long clavate siphunculi and a short hairy cauda. Kadyrbekov (2013b) reviewed the genus and keyed the species, including a new one from Heracleum, so Apiaceae are possible secondary hosts. [The Prunus-feeding species described in this genus (A. indica) belongs in Tumoranuraphis (Eastop & Blackman 2005).]

Avicennina almatina Kadyrbekov Colour of apterae in life is unrecorded; BL 3.7-5.6 mm. Alatae have c.44-53 secondary rhinaria on ANT III. On Heracleum dissectum in south-east Kazakhstan (Kadyrbekov 2013b).

Avicennina sogdiana Narzikulov Apterae (fundatrices?) are green or pale green with darker green markings; BL 2.7-3.4 mm. On Lonicera spp. in Central Asia (Kazakhstan, Kirghisia, Tajikistan, Uzbekistan). Probably host-alternating, but with the secondary host unknown. Oviparae occur on Lonicera in October (Mukhamediev & Akhmedov 1982). [A. spiraecola Akhmedov, described from Spiraea hypericifolia in Kirghisia, was synonymised with A. sogdiana by Kadyrbekov (2013b).]

Avicennina turkestanica Akhmedov Apterae (fundatrices?) are green without darker markings; BL 3.8-5.5 mm. Dispersed on undersides of leaves of Lonicera spp. in Central Asia (Uzbekistan, Kazakhstan; Akhmedov 1994, Kadyrbekov 2013b). The life cycle is unknown; alatae produced in late June presumably migrate to an unknown secondary host..