SYSTEMATIC TREATMENT OF APHID GENERA

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

Five palaearctic species with a broadly rounded cauda living without host alternation on Centaurea. Holman (1991) provided a revision and a key to species. They are adapted for ant attendance.

Acaudinum 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 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 roumanicum Holman Apterae are matt or slightly shiny jet black, BL 1.6-2.7 mm. At stem bases of Centaurea 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, 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) analysed mitochondrial (COI) and nuclear (EF-1α) sequences of the species in Lithuania, 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) appled 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, Cuba and USA (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), 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 (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). 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 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 Pycnantherum 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), 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) 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 in the complex (e.g. balloticola, 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 and R. alnifolia are accepted as primary hosts (Voegtlin et al. 2004, 2005; Yoo et al. 2005). 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.). Tax