The Aphids

Two or (probably) more palaearctic species associated with Rhamnus and Polygonum, and probably with at least a partial host alternation between these two plant genera. The genus is well-defined, being characterised by a very long antennal terminal process, spiracles of abdominal segments 1 and 2 well separated, and the cauda reduced to such an extent that it is almost indiscernible. Apterae have secondary rhinaria on ANT III, and alatae have them on III-IV (-V).The antennal tubercles are smooth and divergent, and the siphunculi are tapering and rather short. The species-level taxonomy is difficult and in need of further work.

Macchiatiella itadori (Shinji) Plate 11g Apterae are yellow, brownish yellow or green with shiny black dorsal abdominal patch and black siphunculi; BL 1.7-2.2 mm. They are found on Polygonum and related genera (Aconogonon, Fallopia, Fagopyrum, Reynoutria). in Japan, Korea and east Siberia. Populations thought to be the primary host generations of the same species occur on Rhamnus spp. (japonica, purshiana) in Japan and Korea (Miyazaki 1971). Apterae on Rhamnus are shining yellow-brown or yellow-green with a brown head, an extensive black dorsal abdominal patch, banded antennae and legs, and black siphunculi (for illustration see Moritsu 1983); BL 1.8-2.1 mm. Large colonies are formed on the leaves, young shoots and woody stems, causing leaf-curl (Hori 1927, as Acaudus rhamni). However, host transfers have not been made and the life cycle is uncertain, being based on synonymy with Hori’s species, which may not be correct, because Hori (1927) stated that his aphid lived all year on Rhamnus and produced apterous as well as alate males. Alatae produced on Rh. japonica were shining green according to Hori (1927), whereas those produced on Rh. purshiana were yellow (Miyazaki 1971), so possibly two or more species are involved. Oviparae and males have been collected or Rh. japonica in September-October (Hori 1927, and BMNH collection). 2n=12.

Macchiatiella rhamni (Boyer de Fonscolombe) Apterae on Rhamnus are yellow-brown or yellow-green, and have antennae ringed with black, pale legs with black tarsi, pale siphunculi and variably developed black markings on abdominal tergites 3-5, which are sometimes completely absent; BL 1.9-2.3 mm. Populations on Rhamnus occur in southern France, Italy, Portugal, Spain, and east to Turkey (BMNH collection). Remaudière (1959) found a population on Phillyrea latifolia (Oleaceae), in the south of France resembling those on Rhamnus but with shorter and thicker appendages. As in M. itadori, there are uncertainties about the life cycle. Anholocyclic populations occur on Rh. alaternus in the Mediterranean area (Barbagallo & Stroyan 1982). Contrary to Blackman & Eastop (1994), there appear to be no authentic western European records of this species from Polygonum. Host alternation to Polygonum is however reported from Ukraine (Chumak 2004), and a form regarded as a subspecies, M. rhamni ssp. tarani, is believed to alternate between Rhamnus spp. and Polygonum in Central Asia (Nevsky 1929a, as Neanuraphis; Kadyrbekov 2009b). Aphids assigned to M. rhamni s. lat. occur on the undersides of leaves of Polygonum spp. in Central Asia, Mongolia and Korea. Specimens in BMNH collection from Polygonum in Mongolia (leg. H. Szelegiewicz) and Korea (leg. W.H. Paik) have more extensive dark pigmentation and longer cephalic hairs. Those from Mongolia have spinal and marginal abdominal tubercles, but less frequently than in European specimens, and those from Korea are without tubercles (as in M. itadori). M. rhamni is also recorded from Siberia on Bassia (= Echinopsilon) sedoides (Ivanoskaya 1977, as Neanuraphis rhamni), but the Siberian aphid has longer siphunculi. It seems likely that more than one species is involved.

Machilaphis Takahashi

Phyllaphidinae

Two oriental species on Lauraceae, resembling Phyllaphis and Diphyllaphis but differing in the form of the empodial hairs and structure of the last rostral segment. Until recently there was thought to be only a single species in the genus, but Quednau (2010) has distinguished the specimens from Japan described as machili by Takahashi (1960c) as a second species.

Machilaphis machili (Takahashi) Apterae are elongate oval or pear-shaped , pale green or yellowish; BL 1.6-2.0 mm. On undersides of leaves of Lauraceae (Machilus, Neolitsea, Phoebe) and there is also a record from Cinnamomum camphora (as Shivaphis cinnamomophila Zhang). Recorded from Japan, Thailand, China, Taiwan and India (Assam, Meghalaya). According to A.K. Ghosh et al. (1971d), a waxy secretion is produced on the host plant. Biology and life cycle have not been studied, and sexual morphs are unknown.

Machilaphis pseudomachili Quednau Colour of apterae in life unknown; BL 1.3-1.9 mm. On ?Machilaphis sp. (probably thunbergii)and Cinnamomum japonicum in Japan (Takahashi 1960c, as M. machili; Quednau 2010). A series of specimens (including the holotype and paratypes) was also purportedly collected from Quercus glauca, but this is an unlikely host and may have been an error of identification or labelling. For further information see Quednau (2010).

Macromyzella Sorin & Arakawa

Aphidinae: Macrosiphini

Two east Asian species related to Macromyzus but with shorter hairs and unsclerotised dorsum, one of which is a fern feeder like Macromyzus, but the other is described from Celtis, a host unlikely enough to require additional confirmation.

Macromyzella celticola Sorin & Arakawa The appearance of apterae in life is unknown; BL c.1.8 mm. The host is recorded as Celtis jessoensis. In Japan (Honshu). Alatae viviparae and alate males possibly belonging to this species but collected on various other plants (presumably vagrant individuals) were described (as Macromyzella sp.) in the same paper (Sorin & Arakawa 2005).

Macromyzella polypodicola (Takahashi) Apterae are yellow, orange or cream-coloured, with black siphunculi and cauda; BL c.2 mm. On ferns (Asplenium, Cheilanthes, Christella, Cyclosorus, Diplazium, Polystichum) in east and south-east Asia (Japan, Okinawa, Korea, Thailand, China, Taiwan, Philippines, Malaysia, Indonesia, New Britain). Takahashi (1963a), as Macromyzus) provided a redescription.

Macromyzus Takahashi

Aphidinae: Macrosiphini

Four medium-large Asian fern-feeding species with spiculose head, siphunculi with subapical reticulation, and a rugose dorsal cuticle with raised dark hair-bearing sclerites. The apparent relationship with Tuberoaphis would be substantiated if the host alternation reported for M. woodwardiae, with a sexual phase and a remarkably modified fundatrix on Hydrangea (Miyazaki 1972),could be experimentally confirmed. The genus was reviewed by Su & Qiao (2010a).

Macromyzus (Anthracosiphoniella) maculatus (A.N. Basu) Apterae are yellowish green with dark brown dorsal markings; BL c.2.4-2.6 mm. On undersides of fronds of various ferns (Asplenium, Athyrium, Deparia, Diplazium, Dryopteris, Eriosorus, Parathelypteris, Phegopteris, Pronephrium) in northern India and China (Su & Qiao 2010a, Su et al. 2014). Apparently anholocyclic (original description, and A.K. Ghosh 1974a).

Macromyzus manoji Raha & Raychaudhuri Apterae are presumably brown with pale-tipped siphunculi: BL c. 2.4 mm. On ferns (Asplenium, Cheilanthes), and also recorded from “Disclesrea alata”, presumably an error for Dioscorea alata, although this this is unlikely to be a true host. In India (West Bengal). Records of M. woodwardiae in West Bengal may refer to this species, including one of oviparae on an unidentified plant in January (A.K. Ghosh 1974a, and see original description).

Macromyzus spinosus Su & Qiao Apterae are elongate oval, yellow-brown; BL 1.8-2.0 mm. On undersides of young leaves of Plagiogyria japonica in Hunan province, China (Su & Qiao 2010a).

Macromyzus woodwardiae (Takahashi) Plate 22b (Fig.32b, Fig.43c) Apterae on ferns are brown with black siphunculi, legs with black distal parts to femora and yellowish tibiae; BL 2.2-2.8 mm. In Japan, Taiwan, China, Korea, Nepal and India. On undersides of fronds and new growth of ferns in many genera. Anholocyclic on ferns in most places, but in Japan it is apparently also partially heteroecious holocyclic, orange-yellow fundatrices and alate spring migrants having been described from Hydrangea (Miyazaki 1972a). Moritsu (1983) records this species from five Hydrangea spp., and also from Deutzia gracilis (Hydrangeaceae) and Weigela coraeensis (Caprifoliaceae); an odd concordance of primary hosts with Rhopalosiphoninus deutzifoliae. Fundatrices have long dorsal processes like those of Tuberoaphis on Hydrangea, and the two genera are evidently closely related, so the unknown secondary hosts of Tuberoaphis may be ferns. However the life cycle of M. woodwardiae still needs experimental confirmationwith host transfers. Records of M. woodwardiae from West Bengal are referable to M. manoji (Raha & Raychaudhuri 1978). 2n=12.

Macropodaphis Remaudière & Davatchi

Macropodaphidinae

A distinctive genus of about six Asian species with greatly enlarged fore femora and numerous dorsal tubercular processes bearing hairs and wax glands. Life cycles are unknown, and host relationships are unclear; most species are found on Potentilla, but three species have Artemisia and Carex also recorded as hosts (although the morphology is not that expected of a Carex-feeding aphid). Ivanoskaya (1981) provided a key to species then known, and (1982) discussed relationships within the genus. W. Zhang et al. (1995d), Qiao & Zhang (2003d) and D. Zhang & Qiao (2008a) reviewed the genus in China. Quednau (2010) reviewed and keyed the species, making several synonymies, and illustrated all available morphs.

Macropodaphis alexandri Ivanoskaya-Shubina (= M. tsherepanovi Ivanoskaya) Apterae are pale green or yellowish green, BL 1.5-2.5 mm. Alatae are unknown. On Potentilla spp. (chrysantha, supina), and also recorded from Artemisia frigida and Carex duriuscula (an unlikely host?) in Russia (west Siberia, Gorno-Altaisk). Closely related to M. paradoxa, but removed from synonymy with that species by Quednau (2010).

Macropodaphis dzhungarica Kadyrbekov Apterae are glaucous lilac-colured, with mainly dark appendages; BL c. 1.5-1.7 mm. Alatae are unknown. On Potentilla (=Pentaphylloides) parviflora in Kazakhstan, and redescribed from the Tibetan Plateau (D. Zhang & Qiao 2008a). Closely related to M. primigenius.

Macropodaphis paradoxa Zachvatkin and Aizenberg (= M. kuraijensis Ivanoskaya) Plate 3i (Fig.12c) Apterae are pale green or yellowish green, appendages mainly pale with dark apices; BL 1.5-2.0 mm. One alata has secondary rhinaria distributed III 26, IV 20, V 3. Described from Carex in east Siberia, but subsequently also recorded by Ivanoskaya (1965) in west Siberia from Artemisia frigida (as M. kuraijensis), and this is more likely to be the true host. M. damrosei Ivanoskaya, described from a single trapped alata, is a possible synonym (Quednau 2010).

Macropodaphis primigenius Ivanoskaya (= M. bamensis Ivanoskaya, = M. dasiphorae Ivanoskaya) Apterae are pale green or glaucous lilac, with dark head, antennae, femora, siphunculi and cauda; BL 1.6-3.8 mm. Apterae have 0-5(-7) secondary rhinaria on ANT III, and one alata has them distributed III 19, IV 2, V 0. On Potentilla fruticosa in Siberia and Kazakhstan, and on ?Carex sp. in Mongolia (BMNH collection, leg. H. Szelegiewicz). Differences in size and colour between populations are considered to be attributable to intraspecific variation (Quednau 2010). An ovipara collected in west Siberia in late August on Potentilla sp. is illustrated by Quednau (2010).

Macropodaphis rechingeri Remaudière & Davatchi (= M. kulundensis Ivanoskaya, = M. ivanoskajae Kadyrbekov) (Fig.12a,b) Apterae are pale green (original description and Kadyrbekov 1991, as M. ivanoskajae) or yellowish grey (Ivanoskaya 1963, as M. kulundensis), with dorsal tubercles very evident; BL 1.4-2.1 mm. An alata has secondary rhinaria distributed III 22, IV 10, V 0. On undersides of leaves of Artemisia austriaca var. orientalis in Iran, moving very quickly when disturbed (original description). However, Kadyrbekov (2012a) recorded it on Potentilla bifurca in Kazakhstan. M. kulundensis, collected on Salvia stepposa (= dumetorum)in Siberia, is regarded as a synonym, as also is M. ivanoskajae, found dispersed on upper and lower sides of leaves of Potentilla sp. in Kazakhstan (Quednau 2010).

Macropodaphis tubituberculata Zhang & Zhang Apterae are green, with whitish dorsal tubercles evident; BL c.2.2 mm. Alatae are unknown. On Carduus crispus in China.

Macrosiphoniella del Guercio

Aphidinae: Macrosiphini

About 115 palaearctic and 5 nearctic species, mostly having siphunculi and cauda similar in length, with siphunculi reticulated over distal half. Most species feed on Anthemideae and have a stiletto-shaped R IV+V. At least half the species feed on Artemisia and/or the closely- related Seriphidium,and 6-8 species are known from each of Achillea, Aster, Helichrysum, Centaurea, and other Cynareae, with fewer from Chrysanthemum and other Anthemidae. The numerous species colonising Artemisia with little evidence of host specificity, and often without apparent biological differences, is difficult to understand and worthy of closer study. Host alternation does not occur. Although found throughout the Northern Hemisphere the genus has a distinctly continental distribution, with about 30% described from Central Asia, Siberia, Mongolia and China. Accounts are available for western Europe (Hille Ris Lambers 1938), Fennoscandia and Denmark (Heie 1995), UK and Ireland (Blackman 2010), Switzerland (Lampel & Meier 2007), European Russia (Shaposhnikov 1964), Siberia (Ivanovskaya 1977), India (R.C. Basu & Raychaudhuri 1976b), Mongolia (Holman & Szelegiewicz 1974b, 1978), Korea (Szelegiewicz 1980, Lee et al. 2002c, Holman et al. 2006a, b), China (Tao 1963), east Siberia (Pashchenko 1998a-b, 1999a-b), Japan (Miyazaki 1971) and North America (Robinson 1987, Jensen et al. 2020)). Kadyrbekov (2018b) provided a key to apterae of the subgenus Asterobium, and Kadyrbekov (2019) provided a key to palaearctic species of Macrosiphoniella s. str., green in life and without dark dorsal abdominal sclerites.

Macrosiphoniella abrotani (Walker) Apterae are greyish green or dull grass green, wax-dusted, with a dark medial stripe with mainly pale legs and antennae, and siphunculi brownish with darker apices; BL 2.4-3.1 mm. Usually found on young stems of Artemisia abrotanum; also recorded from some other Artemisia and Seriphidium spp. and from Tripleurospermum (=Matricaria) inodora, and from two Tanacetum spp., with a single record from Achillea millefolium. In Europe, western Siberia, the Middle East, south-west and Central Asia (Kadyrbekov 2009a, 2017a), north Africa, and introduced to South Africa, Australia, north-eastern USA and Canada. A form corresponding morphologically to southern European populations has been introduced into Argentina (BMNH collection, leg. J. Ortego, and Ortego et al. 2006). Populations in east Asia have been distinguished as subspecies; A. abrotani ssp. chosoni Szelegiewicz on Artemisia spp.in Korea and China (Holman et al. 2006b, Lee et al. 2002c, Szelegiewicz 1980, Zhang et al. 1987), also recorded from Tanacetum boreale in east Siberia (Pashchenko 1998a), and A. abrotani ssp. sainshandi Szelegiewicz on A. sieversiana in Mongolia (Holman & Szelegiewicz 1978), also recorded from Altai, Russia (Kadyrbekov 2014g) and Kazakhstan (Kadyrbekov 2017a). Two species described from A. scoparia in north-west China (Zhang et al. 1999), M. hofuchui and M. tsizhongi, are also members of thisgroup and are not clearly distinct from abrotani (but the illustration of a siphunculus ascribed to M. hofuchui is of a member of the M. antennata group).Oviparae and alate males appear on A. abrotanum in Europe in September-October.

Macrosiphoniella absinthii (L.) Apterae are generally reddish brown, partly wax-powdered, with black head, antennae, legs, siphunculi and cauda, and a black spot in the centre of the abdomen (see influentialpoints.com/Gallery); BL 1.7-2.5 mm. Commmonly on upper parts of stems of Artemisia absinthium in northern, central and eastern Europe, western Siberia, Iran, Central Asia; also in north Africa and the Mediterranean area, where it occurs on other Artemisia and Seriphidium spp. There are also records from Dendranthema zawadzkii (in Poland; Heie 1995), Leucanthemum vulgare (in the Netherlands; Piron 2017), and Tanacetum praeteritum. Introduced to USA and Canada, and to South America (Argentina: Ortego et al. 2006; Chile: Nieto Nafría et al. 2018). Apterous males and oviparae occur in October in England (Blackman 2010). However, alate males have been recorded from Latvia (Opmanis 1928). 2n=12.

Macrosiphoniella achlys Zhang, Chen, Zhong & Li Apterae are shining black; BL c.2 mm. On stems of Artemisia scoparia in Xinjiang-Uygur region of China (Zhang 1999). A member of the atra group.

Macrosiphoniella aetnensis Barbagallo (Fig. 31m) Apterae are green, covered with greyish wax powder except for a bare central dorsal abdominal area; BL 1.7-2.2 mm. On Helichrysum italicum, living on stems and undersides of leaves, in Italy. Apparently the production of sexual morphs occurs late in the season; viviparae were present in November, and an ovipara was collected in December (Barbagallo 1970).

Macrosiphoniella aizhanae Kadyrbekov Apterae are pale green or pale pink with dark brown tibiae and siphunculi; BL 2.3-2.9 mm. On flower stems of Pseudolinosyris grimii on rocky mountain slopes in southern Kazakhstan (Kadyrbekov 2019a).

Macrosiphoniella ajaniae Kadyrbekov Apterae are shining black, BL 1.3-1.5 mm. In small colonies on flowerheads of Ajania fastigiata in south-east Kazakhstan (Kadyrbekov 1999a). Probably a member of the atra group.

Macrosiphoniella (Asterobium) aktaschica Nevsky 1929 Apterae are green, dusted with wax, with brownish antennae, legs, siphunculi and cauda (Kadyrbekov 2018b); BL 2.2-2.6 mm. It was originally described from Aster in Central Asia as a subspecies of M. asteris, and subsequently recorded (as M. aktashica) from Aster (= Pseudolinosyris) grimmii and Heteropappus altaicus respectively by Kadybekov (2005b) and Kadyrbekov & Aoitzhanova (2005),as well as from two species of Erigeron (Kadyrbekov 2014d). Oviparae and alate males of M. (A.) aktaschica, collected on Aster in October in south-east Kazakhstan, were described by Kadyrbekov (2018b), who also provided a redescription of the aptera.[Note that M. aktashica Nevsky 1928 is a quite different species, related to M. tapuskae.]

Macrosiphoniella aktashica (Nevsky 1928) Apterae are very pale, whitish to pale green, with siphunculi dark distally, antennae somewhat pigmented and tibial apices and tarsi dark; BL 2.0-3.1 mm. On undersides of leaves of Tanacetum sp(p). in Central Asia, and also now recorded from various Achillea spp. in Iran and Turkey, Remaudière et al. (2006) providing morphometric data and recording oviparae and apterous males from those countries. A longer-haired form on T. pseudoachillea and Achillea sp. was described as a subspecies, M. aktashica ssp. hirsuta(Daniyarova), but this is synonymised with the nominate species by Kadyrbekov (2019), who also gives the very similar M. tapuskae subspecies status.[Note: M. (Asterobium) aktaschica Nevsky 1929 from Aster is another species; see under M. asteris.]

Macrosiphoniella alatavica (Nevsky) Apterae are rather elongate-bodied, pale green with antennae and upper halves of siphunculi dusky; BL 2.0-2.8 mm. On flower stalks of Artemisia dracunculus in Central Asia. Records from other Artemisia species should be treated circumspectly as the name has been wrongly applied. M. alatavica has been placed as a member of the nitida group (e.g. Holman & Szelegiewicz 1978), close to M. dracunculi, but this does not fit well with the original description. M. lambersi (q.v.) has been erroneously placed as a synonym. M. tadshikana may be closely related.

Macrosiphoniella albiartemisiae Zhang, Chen, Zhong & Li Apterae are blackish brown; BL c.1.7 mm. On Artemisia stelleriana in Qinghai Province, China (Zhang 1999).

Macrosiphoniella altaica Ivanoskaya (= M. longirostrataHolman & Szelegiewicz; see Kadyrbekov 2019a) Apterae are pale green with dark antennae, legs, siphunculi (except pale basally) and cauda; BL 1.6-2.1 mm. On upper parts of stems of Artemisia frigida in Siberia. Kadyrbekov (2011c) assigned aphids on A. austriaca in Kazakhstan to this species, and also reported its occurrence in Mongolia (Kadyrbekov 2017a). Probably a member of the nitida group. [The specimens recorded as M. longirostrata from A. stellariana in Sakhalin by Pashchenko (1988a) were presumably M. sachalinensis (q.v.). See also comment under M szalaymarzsoi.]

Macrosiphoniella (Phalangomyzus) antennata Holman & Szelegiewicz (Fig.13h) Apterae are elongate spindle-shaped, bluish or pinkish pruinose with brown head and a dark spot between siphunculi, and with appendages (including siphunculi and cauda) mainly blackish brown; BL 2.7-4.2 mm. On Artemisia spp., living singly or in small groups on the lower sides of leaves. In Mongolia, Russia (Altai, east Siberia) Kazakhstan and Korea, and also now recorded from California, USA (Skvarla et al. 2017). M. antennata ssp. takahashii is placed as a synonym by Lee et al. (2002c). M. annulata Zhang, described from China on Artemisia capillaris, resembles antennata but is green in life, and mounted specimens have pale middle sections to the tibiae. A second sample of antennata-group aphids from China (BMNH collection, VFE 18,135), from Artemisia ?annua, also has pale middle sections to the tibiae, but the siphunculi are shorter, like antennata s. str. from Mongolia. Until this variation in pigmentation and siphuncular length has been studied further it seems best to apply the name antennata to all members of this group. 2n=12 (for VFE 18,135).

Macrosiphoniella (Papillomyzus) arctica Pashcshenko Apterae are greenish grey, sometimes with pinkish tinge, wax-dusted; BL c.2.5 mm. In small dense colonies on upper parts of Artemisia arctica in north-east Siberia (Pashchenko 1999a). Other morphs unknown.

Macrosiphoniella arenariae Bozhko Apterae are green with appendages mainly dark and with dark dorsal abdominal markings, especially on segments 2-4; BL c. 2 mm. On stems of Artemisia arenaria (= A. campestris ssp. inodora) in Ukraine, and later recorded from other Artemisia spp. (Bozhko 1976), and in Russia (north Caucasus) and Kazakhstan (Kadyrbekov 2012a). It is possibly a member of the atra group, although these are usually black in life.

Macrosiphoniella (Asterobium) argynica Kadyrbekov Apterae are shining dark brown with blackish siphunculi and cauda, legs also blackish aprat from bases of femora and middle parts of tibiae; BL 1.8-2.3 mm. On flowers and stems of Galatella sedifolia ssp. biflora on rocky mountain slopes in central Kazakhstan (Kadyrbekov 2018b).

Macrosiphoniella artemisiae (Boyer de Fonscolombe) (Fig.13m) Apterae are greyish green, wax powdered, with appendages mainly black (see influentialpoints.com/Gallery); BL 2.3-3.6 mm. A yellow mutant form was found in England (Blackman 2006). Common on upper parts of Artemisia vulgaris, especially betweeninflorescences, and frequently forming large colonies. Several other species of Artemisia are also recorded as hosts, and there are also records from Leucanthemum vulgare and Tanacetum parthenium. Throughout Europe, eastward across Siberia, south-west and Central Asia, Pakistan, Mongolia and China, and introduced to North America and Argentina (Ortego et al. 2004). Populations on Artemisia arborescens in Sicily were described as a subspecies, M. artemisiae ssp. meridionalis Barbagallo, distinguished by smaller size, shorter siphunculi and fewer secondary rhinaria and caudal hairs. Similar aphids are also found in the Canary Islands. The east Asian M. yomogifoliae is closely related and has been regarded as a subspecies. M. sibirica Ivanoskaya, on Artemisia spp. in Siberia, cannot be differentiated from M. artemisiae using its published description, and M. tanacetaria ssp. divia described from Tanacetum and Ambrosia in east Siberia (Pashchenko1999a), may also be a synonym. Oviparae and alate males appear in September-October in western Europe. 2n=12.

Macrosiphoniella (Asterobium) asteris (Walker) Apterae are brownish green with black dorsal spots, body somewhat wax-powdered, with appendages mainly dark (see influentialpoints.com/Gallery); BL 2.3-3.2 mm. On Aster tripolium, in small colonies on upper parts of stems and in theinflorescences; also recorded from Galatella sedifolia (in Spain). Throughout Europe, and also recorded from Korea (Lee et al., 2002c). Possibly it also occurs in Central Asia, if M. erigeronis ssp. villosae Smailova is a synonym. Oviparae and apterousmales occur in early October (northern Italy; Hille Ris Lambers 1938).

Macrosiphoniella atra (Ferrari) Apterae are shiny black with mainly black appendages; BL c.1.8-2.1 mm. In shoot tips and inflorescences of Artemisia alba (= camphorata) in Italy (Roberti 1958), and also found on other Artemisia spp.and in other parts of Europe (France, Netherlands, Hungary, former Yugoslavia, Bulgaria, Greece), and in west Siberia and Kazakhstan. Holman & Szelegiewicz (1978) described a subspecies, M. atra ssp. latysiphon, from Mongolia, and this form is reported from Artemisia spp. in Korea (Lee et al. 2002c, Holman et al. 2006b), Kazakhstan (Kadyrbekov 2009a, 2017a) and the Altai (Kadyrbekov 2014g), and also from east Siberia on Tanacetum boreale (Pashchenko 1988a). Another subspecies, M. atra ssp. turanica, was described from Tajikistan (Narzikulov & Umarov 1969, as M. affinis ssp. turanica), and was given full species status by Remaudière & Remaudière (1997, as M. affinis), but this seems to be based only on size-related differences. Holman & Szelegiewicz (1978) discussed morphological variation in the atra group, which includes M. atrata Umarov (on Artemisia sp. in Tajikistan), and M. nigropilosa Nevsky (described from Artemisia persica in Kazakhstan, but subsequently collected frequently on A. santolinifolia, and also recorded from other Artemisia spp., and from Thymus serphyllum in Central Asia; see Narzikulov & Umarov 1969 and Kadyrbekov 2017a). Umarov (1964) described Central Asian populations of M. nigropilosa differing from Nevsky’s as a subspecies, M. nigropilosa ssp. pamirica. A member of the atra group is also present in north-west India (Bindra & Sekhon 1969, as hikosanensis).The atra group may also include M. achlys, M. ajaniae, M. albiartemisiae, M. brevisiphona, M. himalayana and M. santolinifoliae. Kadyrbekov (1999a) provided a key to distinguish some of these species then described.

Macrosiphoniella austriacae Bozhko Apterae are pale brown (with darker transverse bars on dorsal abdomen), densely covered in white wax powder; BL c. 1.8-1.8 mm. On stems of Artemisia spp., specially A. austriaca,in Ukraine, Moldova, south-east Russia and western Siberia, and also recorded from Kazakhstan (Kadyrbekov 2004b). Close to M. teriolana.

Macrosiphoniella borealis Pashchenko Apterae are green with mainly pale brown appendages; BL c.2.6 mm. On flower stems of Artemisia arctica, A. tanacetifolia and (one sample) Tanacetum boreale, usually forming small dense colonies, in north-east Siberia(Pashchenko 1998a), and also recorded from Artemisia rupestris in Kazakhstan (Kadyrbekov 2009b). Oviparae and an apterous male were collected in Siberia in August.

Macrosiphoniella (Asterobium) bozhkoae Remaudière Apterae are dark green, wax-dusted; BL c. 2.5 mm. On Linosyris (= Galatella) villosa in Ukraine (Bozhko 1976, as M. ucrainica), and also recorded from south-east Russia and west Kazakhstan.

Macrosiphoniella brevisiphona Zhang Appearance in life is unknown, probably dark as it appears to be a member of the atra group; BL c. 2.4 mm. On Artemisia sp. in Tibet. A paratype specimen has been examined, and seems close to M. atra ssp. latysiphon.

Macrosiphoniella capillaricola Holman, Lee & Havelka Apterae are dark brown to shiny black; BL 2.0-2.8 mm. Immatures are reddish brown. On stems of Artemisia in South Korea, clearly preferring A. capillaris (Holman et al. 2006b).

Macrosiphoniella caucasica (Wojciechowski) Apterae are matt brownish black; BL 1.7-1.9 mm. On flowers and fruits of Pyrethrum sp. in Russia (Caucasus).

Macrosiphoniella cayratiae Tseng & Tao Colour in life is unknown; BL of aptera c.2.4 mm. On Cayratia japonica in China.

Macrosiphoniella cegmidi Szelegiewicz Apterae are bright greenish, with siphunculi dark at apices; c. BL 1.7-2.0 mm. On Artemisia sp. in Mongolia. (See comment under M. szalaymarzsoi.)

Macrosiphoniella (Sinosiphoniella) chaetosiphon Takahashi & Moritsu (Fig.13e) Apterae are dark red with black siphunculi and cauda, BL 2.0-2.5 mm. On Artemisia spp. in Japan and Korea. Oviparae were found on A. princeps in October in South Korea (Holman et al. 2006a). Generic placement was discussed by Holman & Szelegiewicz (1979).

Macrosiphoniella chamaemelifoliae Remaudière & Leclant (Fig.13o) Apterae are pale green, wax dusted, with head, antennae, legs, siphunculi and cauda black; BL 1.6-2.6 mm. In colonies, often very dense, on shoots of Artemisia chamaemelifolia at altitudes of 1,600-2,000m in the French Alps. It is also reported to occur in Iran (Rezwani 2010). Oviparae and both apterous and alate males appear in France from August-October, fundatrices in late June. Aphidius absinthii and Ephedrus niger are recorded as parasitoids of this species (original description).

Macrosiphoniella cinerascens Hille Ris Lambers Apterae are bright sea-green, with dorsal pattern of white powder; BL 2.1-2.6 mm. On lower leaves of Achillea californica (= millefolium), turning their tips brown. In western USA (California, Colorado, Idaho, Oregon, Washington).

Macrosiphoniella (Papillomyzus) confusa Holman, Lee & Havelka Colour in life is unknown; BL 2.3-2.9 mm. On Tanacetum boreale in Korea (Holman et al. 2006a).

Macrosiphoniella (Asterobium) crepidis Holman & Szelegiewicz Apterae are yellowish green, covered with a greyish wax powder, with dark appendages; BL 1.8-2.5 mm. On Crepis spp.in Mongolia. Other morphs have not been described.

Macrosiphoniella cymbariae Bozhko Apterae are dark yellow-brown; BL c.2.2 mm. On Cymbaria borysthenica in Ukraine.

Macrosiphonella (Asterobium) davazhamci Holman & Szelegiewicz Apterae are matt green, dusted ventrally with greyish wax powder; BL 1.9-2.6 mm. On Aster hispidus in Mongolia, on Aster canescens (Nees) Fisjun (= Kalimeris altaica) in Kazakhstan (Kadyrbekov 2012a) and Iran (see below), and on Aster sp. in China, near Beijing (BMNH collection, leg. V.F. Eastop). Immature oviparae and alate males were found in September (original description), and the sexuales have now been described (including detailed SEM studies) from material collected on K. altaica by G. Remaudière in Iran in November (Kanturski & Barjadze 2018).

Macrosiphoniella dimidiata Börner (Fig.13j) Apterae are are brown or greenish, wax powdered, with black head, siphunculi and cauda, and mainly black (except at bases) antennae and legs; BL 1.8-2.7 mm. On upper parts of shoots of Artemisia spp. (usually campestris) throughout most of continental Europe, western Siberia, south-west and Central Asia (Kazakhstan). In Israel it occurs on A. monosperma (BMNH collection). M. lena Pashchenko, described from Artemisia commutata (= pubescens) in east Siberia (Pashchenko 1998a), is possibly this species, and M. kirgisica Umarov in Central Asia also seems closely related. [There is an argument for using the earlier name fasciata del Guercio for this species (Leclant 1968b), but the application of this name is confused and we therefore here retain the more generally accepted name dimidiata.] Oviparae and apterous males occur in October in Finland (Heikinheimo 1997). 2n=12.

Macrosiphoniella dracunculi (Umarov) Apterae are green with dark siphunculi, BL c. 2.6-2.8 mm, on stems of Artemisia dracunculus in Kyrgyzstan. Probably close to M. nitida, but with shorter antennal terminal process and longer siphunculi (Holman & Szelegiewicz 1978).

Macrosiphoniella (Asterobium) elegans Pashchenko Apterae are shining black; BL c. 2.9 mm. On shoot apices and flower buds of Galatella dahurica in east Siberia (Pashchenko 1999a).

Macrosiphoniella (Sinosiphoniella) elenae Pashchenko Apterae are dark brown; BL c. 1.9 mm. Found in small sparse colonies on stems of Artemisia stolonifera, visited by ants (the short siphunculi and cauda are clearly adaptations for ant attendance). In east Siberia (Pashchenko 1999b).

Macrosiphoniella (Asterobium) erigeronis Nevsky Apterae are shining brown, with black siphunculi and cauda; BL 1.7-2.5 mm. Forming dense colonies on flower stalks of Erigeron acer in west and central Asia (Georgia, Azerbaijan, western Siberia, Kazakhstan, Uzbekistan).

Macrosiphoniella erythraea Zhang & Qiao Apterae are red in life. Described from two apterae and immatures on Artemisia sp. in Fujian province, China(Qiao & Zhang 1999c, 1999d). It is said to be near M. myohyangsani, except for its colour in life and paler, more hairy cauda.

Macrosiphoniella femorata Bozhko Apterae are dark green; BL c.1.9 mm. On Hieracium umbellatum in Ukraine. This species could not be included in the key to aphids on Hieracium on the basis of the published description.

Macrosiphoniella formosartemisiae Takahashi Apterae are shiny black with mainly blackish brown appendages, BL c. 1.8-2.2 mm. On stems of Artemisia spp. in Japan, China, Korea, Taiwan, east Siberia, Mongolia and India. Sexuales do not appear to have been described; the species is anholocyclic in Taiwan (Takahashi 1923). Holman & Szelegiewicz (1978) compared Mongolian and Japanese material. 2n=10.

Macrosiphoniella frigidae Ivanoskaya Apterae are yellowish-green with a pattern of dark dorsal abdominal pigmentation and dark appendages; BL c. 2.7 mm. In small colonies on Artemisia frigida and Artemisia sp. in western Siberia, and also recorded from A. compacta and A. frigida in Kazakhstan (Kadyrbekov 2004b). Probably it is a member of the dimidiata/teriolana group.

Macrosiphoniella frigidicola Gillette & Palmer Apteraea are bluish green, wax-powdered, with siphunculi dusky at apices; BL c.1.6-1.7 mm. On leaves of Artemisia frigida, and perhaps sometimes on other Artemisia spp.and Seriphidium tridentatum, although there is possible confusion with M. abrotani. In central and western North America; records from eastern North America are more likely to be abrotani. A record from Krascheninnikovia lanata was probably of vagrants. Oviparae and apterous males appear in October in Colorado (Palmer 1952).

Macrosiphoniella frigidivora Holman & Szelegiewicz (Fig.13l) Apterae are green or pinkish green dusted with fine grey wax powder, with blackish antennae, legs, siphunculi and cauda; BL 1.6-2.8 mm. At ends of shoots of Artemisia frigida in Mongolia, and also recorded from A. frigida and A. rupestris in Kazakhstan (Kadyrbekov 2017a), and from A. turcomanica in Iran (Sedighi et al. 2020b). The latter authors provide a redescription. A paratype specimen of M. hokkaidensis in the BMNH collectionfrom Japan is very similar, and the distinction between these two species needs to be confirmed.

Macrosiphoniella (Asterobium) galatellae Bozhko Apterae are shining brownish black with black appendages; BL 2.3-2.7 mm. In small colonies of very mobile aphids on flower stems of Aster and Galatella spp., dropping readily when disturbed (Bozhko 1976b).In eastern Europe (Hungary, Slovakia, Ukraine), Russia (north Caucasus) and Kazakhstan (Kadyrbekov 2004b, 2012a). A species described from China, M. quinifontana, is very similar and possibly a synonym.

Macrosiphoniella gaoloushana Zhang, Chen, Zhong & Li Apterae are black; BL 1.8-2.5 mm. On Artemisia japonica in Gansu Province, north-west China (Zhang 1999).

Macrosiphoniella glabra (Gillette & Palmer) Apterae are shining “shamrock green”, with dusky spots at bases of dorsal hairs, mainly black legs and antennae, and black siphunculi; BL 1.8-2.3 mm. Singly or in small colonies on leaves and flowerheads of Artemisia dracunculus in western USA from Washington to New Mexico, and there is also a record from Chrysothamnus viscidiflorus. Oviparae and apterous males appear in September-October (Palmer 1952).

Macrosiphoniella (Phalangomyzus) gmelinicola Szelegiewicz Apteraea are shiny black with blackish antennae, legs, siphunculi and cauda; BL c.3.2-3.3 mm. On upper parts of stems of Artemisia spp.in Korea and east Siberia.

Macrosiphoniella (Phalangomyzus) grandicauda Takahashi & Moritsu Apterae are pale yellow or pale green with mainly dark brown antennae and legs, black siphunculi and pale cauda; BL 2.5-3.9 mm. On Artemisia spp., especially on older leaves, and there are also host records from Ainsliaea aptera, Arctium lappa and Petasites japonicus.In Japan, China, Korea, east Siberia, India and Nepal, and also recorded from Maryland, USA (Foottit et al. 2006). A record from Algeria (Laamari et al. 2013) requires additional confirmation. 2n=12 (Chen & Zhang, 1985b).

Macrosiphoniella helichrysi Remaudière (Fig.31f) Apterae are grey with black appendages, the body covered in wax powder except for a shiny black central dorsal abdonimal patch and spots anterior to bases of siphunculi; BL 1.4-2.6 mm. On shoot apices and in flowerheads of Helichrysum spp., the adult apterae dropping readily from the plant when disturbed. In southern Europe, Turkey, Iran (Rezwani et al. 1994), and introduced to South Africa. Aguiar & Ilharco (2005) provided a detailed redescription. Sexual morphs are apparently unknown.

Macrosiphoniella (Sinosiphoniella) hikosanensis Moritsu (Fig.13g) Apterae are reddish-yellow with reddish head and prothorax, mainly pale antennae and tibiae, mainly dark femora, dark antesiphuncular spots, and black siphunculi and cauda (Moritsu 1983); BL 1.5-2.8 mm. On undersides of older leaves of Artemisia spp. in Japan, Korea, and east Siberia. Indian records of M. hikosanensis apply to other species, mostly kikungshana, but Bindra & Sekhon (1969) had a member of the atra group.

Macrosiphoniella (Ramitrichophorus) hillerislambersi Ossiannilsson Apterae are reddish with (in life) a large black dorsal abdominal spot, and dark appendages including siphunculi and cauda; BL 1.6-1.9 mm. On Helichrysum arenarium, colonising parts near or just below ground level. Attended by ants. In Denmark, Sweden, Germany, Lithuania (Rakauskas et al. 2008) and Switzerland (Heie 1995). There is also a record from Arctium tomentosum in Kazakhstan (Kadyrbekov 2014d).

Macrosiphoniella himalayana Saha & Chakrabarti Apterae are blackish; BL 1.9-2.4 mm. on undersides of leaves and in inflorescences of Artemisia spp. in India and Pakistan. Oviparae and alate males were collected in October (original description). Probably a member of the M. atra group (q.v.).

Macrosiphoniella hokkaidensis Miyazaki Apterae are yellowish green, without wax according to original description (but see Lee et al. 2002c), with head reddish brown, and antennae, legs (except base of femora), siphunculi and cauda black; BL 1.8-2.6 mm. Described from Artemisia montana in Japan, and there are also records from several Artemisia spp. in east Siberia (Pashchenko 1988a) and Korea (Lee et al. 2002c, Holman et al. 2006b). M. frigidivora Holman & Szelegiewicz on A. frigida in Mongolia is closely related and possibly the same species, although described as wax-powdered in life.

Macrosiphoniella huaidensis Zhang Apterae are green; BL c.2.5 mm. On Artemisia sp. in China. 2n=12 (Chen & Zhang 1985b)

Macrosiphoniella huochengensis Zhang, Chen, Zhong & Li Colour of apterae in life and BL are unrecorded. On leaves of Artemisia scoparia in Xinjiang-Uygur region of China (Zhang 1999). Also recorded from Euphorbia lunulata (= esula), but these were presumably vagrants.

Macrosiphoniella insignata Kadyrbekov Apterae are shining black; BL 1.3-1.7 mm. At shoot apices of an Artemisia sp. in south-east Kazakhstan (Kadyrbekov 1999a).

Macrosiphoniella (Papillomyzus) iranica Nieto Nafría & Pérez Hidalgo Colour of apterae in life is unknown, BL 2.8-3.2 mm. On an unidentified Centaurea sp. in western Iran (Nieto Nafría & Pérez Hidalgo 2013b).

Macrosiphoniella (Asterobium) ixeridis Holman, Lee & Havelka Apterae are pale green to yellowish green with dark head, dark tips to appendages and black siphunculi; BL 2.3-2.9 mm. On upper parts of Ixeris chinensis and Youngia sonchifolia in Korea (Holman et al. 2006).

Macrosiphoniella (Ramitrichophorus) janckei Börner (Fig.31g) Apterae are reddish brown to black, with green spots at bases of siphunculi, powdered with wax; BL 1.8-2.0 mm. On stalks, flowerheads and undersides of leaves of Helichrysum arenarium. In northern Germany, Poland, Lithuania (Rakauskas et al. 2008), Ukraine, Russia (west Siberia, Volga region, North Caucasus) and Kazakhstan (Kadyrbekov 2011c). Oviparae and apterous males occur in Germany in late September (Müller 1975a).

Macrosiphoniella jaroslavi Szelegiewicz Apterae are green, dusted with fine grey wax powder, with blackish antennae, legs, siphunculi and cauda; BL 1.6-2.3 mm. On tips of shoots and in inflorescences of Artemisia messerschmidtiana (var. discolor) in North Korea.

Macrosiphoniella (Asterobium) jaroslaviholmani Stekolshchikov & Khureva Apterae are green, yellow-green or green with yellow-green abdomen, dull glistening, without wax powder, with antennae dark except at base of ANT III, legs mainly pale except for tarsi and apices of femora and tibiae, siphunculi mainly dark and cauda pale to grey; BL 3.2-3.8 mm. On shoot apices and in inflorescences of Artemisia spp. (arctica, tilesii) in north-east Siberia (Chukotka). The life cycle is abbreviated; old fundatrices, apterous and alate viviparae, oviparae and apterous males were all collected in early August (Stekolshchikov & Khureva 2015, as M. jaroslavi). [The name was replaced by Stekolshchikov & Khureva (2018) because it is a secondary homonym of M. jaroslavi Szelegiewicz.]

Macrosiphoniella kalimpongense R.C. Basu & Raychaudhuri Appearance of apterae in life is unknown; BL 2.6-3.3 mm. On Artemisia vulgaris and Artemisia sp. in India (West Bengal) and Nepal (BMNH collection).

Macrosiphoniella kareliniae Kadyrbekov, Renxin & Shao Apterae are greenish with grey wax film, siphunculi brown with basal third pale; BL 2.0-2.5 mm. On stems of Karelinia caspia in central and south Kazakhstan and Xinjiang-Uygur region of western China (Kadyrbekov et al. 2002).

Macrosiphoniella kaufmanni Börner (Fig.13m) Apterae are green, lightly wax-dusted (according to Bozhko 1976), with dark siphunculi and cauda; BL c.1.9-2.2 mm. On Artemisia pontica in Czech Republic and Hungary, and also recorded from Ukraine on A. arenaria (Bozhko 1976), and from Italy on A. alba (Barbagallo & Pollini 2014). There is no adequate description of this species. Remaudière & Leclant (1972) distinguish it from M. chamaemelifoliae. Monoecious holocyclic according to Börner (1952), but sexual morphs are undescribed.

Macrosiphoniella kermanensis Mehrparvar & Rezwani Apterae are greyish green thinly covered in wax powder; BL 2.1-2.6 mm. Alatae have 5-8 secondary rhinaria on ANT III. In sparse colonies in April on shoots of Artemisia (= Seriphidium) sieberi in Iran (Mehrparvar & Rezwani 2007). Life cycle and sexual morphs are unknown. Very similar to M. seriphidii and M. terraealbae, and the characters distinguishing it from those species could be those of the fundatrix morph.

Macrosiphoniella khinganica Pashchenko Apterae are shining green; BL c.2.3 mm. On Artemisia mongolica in east Siberia. Originally described as a subspecies of M. pulvera (Pashchenko 1998b), but differing from that speciesin several morphological features, including longer antennal hairs, and given separate species status by Kadyrbekov (2019).

Macrosiphoniella kikungshana Takahashi Apterae are shining brown with mainly dark brown appendages; BL 2.1-3.2 mm. On undersides of older leaves of Artemisia spp. in China, Korea, east Siberia, Nepal and India (Sikkim). Szelegiewicz (1980) redescribed the species, and also described a population on A. sylvatica in Korea with shorter dorsal hairs and paler femora as a subspecies, A. kikungshana ssp. sylvaticae,but Holman et al (2006b) regard the subdivision as unjustified. Oviparae occur in October (Sikkim). Some of the Indian records of M. hikosanensis should be referred to this species (see R.C. Basu & Raychaudhuri 1976b). 2n=12.

Macrosiphoniella kirgisica Umarov Apterae are greyish green with dark green spots on dorsal abdomen, and dark brown antennae, legs, siphunculi and cauda; BL c. 2.0-2.1 mm. In large colonies on stems of numerous Artemisia/Seriphidium spp. (Kadyrbekov 2003c, 2017a) in Central Asia (Kazakhstan, Kyrgyzstan, Tajikistan, Xinjiang-Uygur region of China). We could not distinguish this species from M. dimidiata on the basis of the published description.

Macrosiphoniella (Asterobium) konyratica Kadyrbekov Apterae are green, wax-powdered, with blackish appendages; BL c.1.8 mm (2 specimens). On stems and flowers of Galatella sedifolia ssp. biflora on rocky mountain slopes in central Kazakhstan (Kadyrbekov 2018b).

Macrosiphoniella (Sinosiphoniella) kuwayamai Takahashi (Fig.13f) Apterae are dull green-black dusted with transverse bands of grey wax powder, reddish head and mainly black antennae, legs, siphunculi and cauda; BL 1.5-3.2 mm. On Artemisia spp., usually living on lower parts of stems and older leaves. In Japan, Korea, China and E Siberia. Szelegiewicz (1980) redescribed apt. and al. vivip. of this species from Korea.

Macrosiphoniella lambersi Verma Colour of apterae in life unknown; siphunculi pale and cauda with 7 hairs in the holotype according to the original description, but an apt. in the BMNH collection labelled as a paratype (leg. K.D. Verma) has dark siphunculi and its cauda has 11 hairs. The measurement of the base of ANT VI must also be wrong in the original description; the paratype has ANT PT/BASE ratio of 3.86. On Artemisia scoparia in India (Jammu). It has been erroneously synonymised with M. alativica in catalogues.

Macrosiphoniella lazoica Pashchenko Apterae are blackish green; BL c.2.3 mm. At stem bases of Saussurea pulchella, attended by ants. In east Siberia (Pashchenko, 1999b).

Macrosiphoniella leucanthemi (Ferrari) Apterae are yellowish green, with a darker green or reddish brown band between the siphunculi, and appendages with contrasting pale and dark sections; BL 2.3-2.7 mm. Alatae have conspicuously brown-bordered wing veins. On undersides of basal leaves of Leucanthemum spp. in Europe (Italy, Hungary, France, Netherlands), and also recorded from USA (Oregon, Pennsylvania, Maryland). Stoetzel & Miller (1999) provided a redescription of apterae and alatae.

Macrosiphoniella lidiae Umarov Apterae are green, powdered with wax, with appendages dark except for bases of ANT III and siphunculi; BL c. 1.9 mm. In large colonies at tops of stems of an Artemisia sp. in Kazakhstan.

Macrosiphoniella lijiangensis Zhang, Zhong & Zhang (= Macrosiphoniella sensorinuda Zhang, Zhong & Zhang) Appearance in life is unknown: BL 2.4-2.8 mm. On Artemisia vulgaris in China (Hengduan mountain region).

Macrosiphoniella (Asterobium) linariae (Koch) (Fig.15a,b) Apterae are brownish black, with black appendages; BL c. 2.4-2.8 mm. On stems of Aster linosyris in Germany, northern Italy, and also reported from Greece (Tsitsipis et al. 2007). Oviparae and alate males occur in October (Hille Ris Lambers 1938).

Macrosiphoniella lithospermi Bozhko Apterae are shining brown with dark brown dorsal markings, siphunculi and cauda black; BL c. 3 mm. In colonies on stems of Lithospermum arvense in Ukraine (Bozhko 1976b).

Macrosiphoniella lopatini (Umarov) Apterae are grass green powdered with greyish wax, with siphunculi dark only on reticulated part; BL c.3.4-3.6 mm. On stems of an Artemisia sp. in Tajikistan, and also recorded from Kazakhstan (Kadyrbekov 2017a). Alatae are unknown.

Macrosiphoniella ludovicianae (Oestlund) Apterae are glaucous green or yellowish green dusted with greyish white wax; BL 2.0-2.8 mm. Commonly on leaves of Artemisia ludoviciana, but it has also been recorded from A. vulgaris. A record from Tanacetum sp. may have been a misidentification, and arecord from Krascheninnikovia (=Axyris) lanata was probably of vagrants. Widely distributed in North America(Palmer 1952, Robinson 1987). Oviparae and alate males occur in October. 2n=12.

Macrosiphoniella maculata Nevsky Apterae are yellow-green or green with black dorsal spots and antesiphuncular sclerites; BL 1.8 mm. On Medicago sativa (which is unlikely to be its true host)in the Pamirs at 3,600 m, in Tajikistan.

Macrosiphoniella madeirensis Aguiar & Ilharco Apterae are pale grey mostly covered with fine whitish wax powder but leaving black central dorsal spot; BL 1.9-2.9 mm. On terminal shoots and leaf petioles of Helichrysum melaleucum in Madeira (Aguiar & Ilharco 2005).

Macrosiphoniella (Ramitrichophorus) medvedevi (Bozhko) (Fig.31e) Apterae are grey with brownish red speckling, or reddish yellow; BL 1.6-2.6 mm. On stems and flowers of Helichrysum arenarium in Ukraine and Poland, attended by ants (Szelegiewicz 1958, as M. nasti). It is also reported to occur in Iran (Rezwani et al. 1994).

Macrosiphoniella miestingeri (Börner) Colour of apterae in life is unrecorded; BL 2.7-3.0 mm. Under lower leaves of Tanacetum corymbosum, which are curved and yellowed. Only known from Germany.

Macrosiphoniella millefolii (De Geer) Plate 27a (Fig.5g) Apterae are yellowish green, powdered with grey wax except for spinal stripe on abdomen and presiphuncular spots, and with black appendages (see influentialpoints.com/Gallery); BL 2.1-3.6 mm. On Achillea spp., especially A. millefolium, forming colonies especially in the inflorescences. Also occasionally found on plants in related genera (Leucanthemum, Tanacetum, Tripleurospermum). Common throughout most of Europe, in west Siberia, Iran, Central Asia (Kadyrbekov 2017a), and introduced to North America. In east Siberia a form occurs with more secondary rhinaria in apterae (38-59, as opposed to 8-28 in European populations) and a rather shorter antennal terminal process (M. millefolii ssp. orientalis Pashchenko 1998). M. sudkharis (q.v.) could be a cold-temperature form of M. millefolli.Oviparae and alate males occur in September-October in north-west Europe (Heie 1995). Sobhani (1970) studied the biology, life cycle and morphology of M. millefolii in Germany. Morlacchi et al. (2011) studied its developmnet on A. collina. 2n=12.

Macrosiphoniella mutellinae Börner Apterae are thickly dusted with grey wax; BL 1.7-2.3 mm. On Artemisia genepi and A. mutellina in Austria. A single, presumably vagrant, aptera was found on Leontopodium alpinum in the French Alps (Remaudière 1954).

Macrosiphoniella (Chosoniella) myohyangsani Szelegiewicz Apterae are pale green, with lead-coloured posterior abdomen and blackish appendages; BL 2.7-3.7 mm. On lower parts of stems of Artemisia spp. in Mongolia, east Siberia, China and Korea (Lee et al. 2002c). Holman et al. (2006a) pointed out that this species may not be specifically distinct from M. spinipes. 2n=12 (Chen & Zhang 1985b).

Macrosiphoniella (Ramitrichophorus) nikolajevi Kadyrbekov Apterae are reddish brown; BL 2.1-2.4 mm. On flower stalks of Helichrysum maracandicum in south Kazakhstan (Kadyrbekov 1999b), and on H. arenarium in Iran (Kanturski & Barjadze 2018). Alatae have been described by Kanturski & Barjadze (2018).

Macrosiphoniella nitida Börner Apterae are light olive-brown or reddish, shiny on back but wax-powdered on abdomen, with dark antennae and tibiae, mainly pale femora and black siphunculi and cauda; BL c. 2.6-2.9 mm. On Artemisia dracunculus in Germany, former Yugoslavia, Iran (Kanturski & Barjadze 2018), western Siberia and Kazakhstan (Kadyrbekov 2012a). Aphids collected on this host in Mongolia with a longer R IV+V and shorter dorsal hairs are regarded as a subspecies, M. nitida ssp. soongarica Szelegiewicz(Holman & Szelegiewicz 1978), although should perhaps have full species status. This subspecies is also recorded from south-east Kazakhstan (Kadyrbekov & Aoitzhanova 2005). M. dracunculiappears to be closely related, as also does M. piceaphis, described from north-west China on Picea sp. (G.Zhang 1999, as Piceaphis).

Macrosiphoniella (Phalangomyzus) oblonga (Mordwilko) Plate 27c Apterae are long-bodied, pale green or apple green, with a darker green spinal stripe, and mostly pale appendages (see influentialpoints.com/Gallery); BL 3.0-5.1 mm. Usually found dispersed on undersides of lower leaves of Artemisia vulgaris, alsosometimes on other Artemisia spp., on cultivated florists’ chrysanthemum (Dendranthema indicum, morifolium, frutescens), and occasionally on plants in other related genera (Leucanthemum, Tanacetum). Oviparae and apterous males occur in September-October in western Europe. M. oblonga occurs throughout Europe and across Asia to Japan and Taiwan. A record from Alaska (International Barcode of Life Consortium 2016) requires further confirmation. Some Asian populations have been given the status of separate species; M. hidaensis Takahashi & Moritsu in Japan and east Siberia, M. paraoblonga R.C. Basu & Raychudhuriin India, and M. similioblonga Zhang in China (also recorded from Korea; Lee et al. 2002c). Further work would be needed to show that these are not all one geographically variable species. 2n=12.

Macrosiphoniella olgae (Nevsky) Apterae are brownish yellow, with dark dorsal spots, wax-dusted; BL 2.2-2.8 mm. On Gnaphalium spp. (sylvaticum, supinum) in Central Asia. Alatae are unknown.

Macrosiphoniella oronensis Szelegiewicz Apterae are shiny brown, with mainly pale antennae and legs and blackish brown siphunculi and cauda; BL 2.0-2.4 mm. On undersides of older leaves of Artemisia japonica in North Korea.

Macrosiphoniella (Phalangomyzus) pallidipes Holman, Lee & Havelka Apterae are green with black antennae, tips of legs, siphunculi and cauda; BL 3.1-3.5 mm. On Artemisia spp. in Korea (Holman et al. 2006).

Macrosiphoniella (Papillomyzus) papillata Holman Apterae are greyish green, covered with a waxy secretion, with black appendages; BL 2.1-3.2 mm. On stems and undersides of leaves of Centaurea spp., and also found on Crupina vulgaris (with heavily infested Centaurea close by). Described from Crimea, and since found in Kazakhstan (Kadyrbekov & Aoitzhanova 2005), Iran, Syria, Turkey, Bulgaria, Greece and France (Remaudière & Remaudière 1997, p.305).

Macrosiphoniella (Ramitrichophorus) paradoxa (Bozhko) Apterae are matt dark brown, with black antennae and siphunculi; BL c.1.6 mm. On shoot tips and among flowers of Helichrysum suaveolens (? an untraceable name) in Ukraine and southern Russia (Shaposhnikov 1964). It is closely related to U. janckei, and possibly a geographical or seasonal variant of that species.

Macrosiphoniella paucisetosa Robinson Colour of apterae in life is unrecorded, probably pale greenish except for blackish brown antennae (beyond base of III), tibiae and siphunculi (except basally); BL 1.8-2.5 mm. On Artemisia spp. in Manitoba, Canada (Robinson 1987).

Macrosiphoniella (Phalangomyzus) pennsylvanica (Pepper) Apterae are green with two longitudinal rows of wax markings; BL 1.9-2.5 mm. On Achillea millefolium in North America (Arizona, California, N Carolina, Pennsylvania, Tennessee). Monoecious holocyclic with oviparae and alate males in October (original description, as Macrosiphum).

Macrosiphoniella (Phalangomyzus) persequens (Walker) Apterae are green, with darker green spinal stripe, and dusky/dark apices to antennal segments, legs and siphunculi (see influentialpoints.com/Gallery); BL 4.2-5.2 mm. On undersides of lower leaves of Tanacetum vulgare in Europe, and also recorded from Pyrethrum millefoliatum (Bozhko 1976b). Oviparae and apterous males occur in September-October. 2n=12.

Macrosiphoniella procerae Bozhko Apterae are matt green, lightly wax-dusted, with dark apices of siphunculi; BL c. 1.9 mm. In small colonies on upper parts of stems of Artemisia procera (= abrotanum) in Ukraine (Bozhko 1976b) and south-west Russia (Kadyrbekov 2014g), and also recorded from A. glauca (= dracunculus) in western Siberia and A. terrae-albae in Kazakhstan.

Macrosiphoniella pseudoartemisiae Shinji Apterae are pale yellow-brown with greenish tinge (slide label, Japanese specimens, BMNH collection); BL 1.8-2.8 mm. A common species on Artemisia spp. in east and south-east Asia (Japan, Korea, Indonesia, Malaysia, India, Sri Lanka) and introduced to South America (Argentina, Ortego et al. 2006); Brazil, BMNH collection). L.K.Ghosh (1972b) described the ovipara from Bhutan and Maity & Chakrabarti (1981b) found one in Uttar Pradesh in October, but males have not been reported. There has been some confusion with another common and widely distributed east Asian species, M. yomogifoliae, which has entirely black tibiae. This is usually a reliable means of distinguishing the two species, but one population of M. pseudoartemisiae from Java in the BMNH collection also has entirely black hind tibiae. Another discriminant is the caudal hair number; 10-14 in pseudoartemisiae, 16-32 in yomogifoliae. 2n=10 (Pal & Khuda-Bukhsh 1982, Dutta & Gautam 1993) or 2n=12 (Kar & Khuda-Bukhsh 1988; perhaps this was misidentified yomogifoliae?).

Macrosiphoniella ptarmicae Hille Ris Lambers Apterae are green, wax-powdered, with olive patches at bases of siphunculi; BL 2.2-3.3 mm. On Achillea ptarmica in Europe, and eastward to Central Asia. Oviparae and alate males occur in September-October. Sobhani (1970) studied the biology, life cycle and morphology of M. ptarmicae in Germany.

Macrosiphoniella pulvera (Walker) Apterae are greyish green or greyish white, heavily wax-powdered; BL 1.9-2.9 mm. On undersides of leaves of Seriphidium maritimum, with which they are concolorous. There are also records from several other Artemisia and Seriphidium spp. Widely distributed mostly in coastal regions of Europe (except Iberian peninsula?), and eastward across Asia to Mongolia; the Mongolian aphids, collected on A. adamsi, differed in some respects from European populations (Holman & Szelegiewicz 1978). Oviparae and apterous males of M. pulvera appear in September in Europe.

Macrosiphoniella (Asterobium) quinifontana Zhang, Chen, Zhong & Li Colour of apterae in life is unrecorded; BL c.1.6 mm. On stems of Aster tataricus and Artemisia sp. (a less likely host)in Gansu Province, China (G. Zhang 1999). It’s distinction from M. galatellae needs to be verified.

Macrosiphoniella remaudierei Barbagallo & Nieto Nafría Colour of apterae in life is unknown, probably green with wax dusting; BL 1.6-2.3 mm. On Helichrysum armenium in mountainous regions of Iran (Barbagallo & Nieto Nafría 2016). Probably closely related to M. aetnensis.

Macrosiphoniella (Papillomyzus) riedeli Szelegiewicz Colour of apterae in life is unknown; BL 2.9-3.4 mm. On Carduus, Centaurea and Cuprina in south-west Asia (Syria, Lebanon, Sinai, Iran). Alatae are not described and the biology is unknown. The only difference between this species and M. papillata seems tobe in the degree of pigmentation, and it should possibly be synonymised with that species.

Macrosiphoniella sachalinensis Pashchenko Apterae are grey-green, wax-dusted; BL c.1.9-2.0 mm. In inflorescences of Artemisia stellariana in Sakhalin (Pashchenko 1998b).

Macrosiphoniella sanborni (Gillette) Plate 27b Apterae are shiny, dark red‑brown to blackish brown, broadly spindle‑shaped, with black, relatively short and thick siphunculi, shorter than the black cauda (see influentialpoints.com/Gallery); BL 1.0-2.3 mm. A cosmopolitan pest on cultivated florists’ chrysanthemum (Dendranthema indicum, morifolium, frutescens) on undersides of leaves. Of east Asian origin, where it often occurs on other Compositae (Anthemis, Artemisia, Aster). Sexual morphs are unknown. See also Blackman & Eastop (2000). 2n=12 (many samples from UK and India, and one from China; but Chen & Zhang (1985a,b) recorded 2n=10 for a sample from China).

Macrosiphoniella santolinifoliae Kadyrbekov Apterae are shining blackish brown; BL 1.6-1.9 mm. On flower stalks of Artemisia santolinifolia in south-east Kazakhstan(Kadyrbekov 1999a). A member of the M. atra group.

Macrosiphoniella (Sinosiphoniella) saussureae Holman & Szelegiewicz (= Macrosiphoniella chita Pashchenko) Apterae are reddish brown to brownish black; BL 1.5-2.2 mm. Apterae have secondary rhinaria distrbuted III 9-24, IV 0-4, and alatae have them ditributed III 20+, IV 5-6. On flower stalks of Saussurea salicifolia, attended by ants (the short siphunculi and cauda are modifications for ant attendance). Mongolia and east Siberia (Pashchenko 1999b, as M. chita).

Macrosiphoniella scepariae Bozhko Apterae are rose pink, finely dusted with greyish wax, BL c. 2.3 mm. In large, fairly dispersed colonies on undersides of leaves of Artemisia scoparia in Ukraine (Bozhko 1976b), and also recorded from several other Artemisia/Seriphidium spp. in Kazakhstan (Kadyrbekov 2017a). Kadyrbekov (2019) distinguished populations of green aphids on flower stems of Artemisia in sandy deserts of southern Kazakhstan as a subspecies, M. scepariae deserticola Kadyrbekov.

Macrosiphoniella sejuncta (Walker) (Fig.5h) Apterae are variable, often a mix of green and brown individuals, or green mottled with brown, with pale-based siphunculi (see influential points/Gallery); BL 2.5-3.1 mm. On Achillea millefolium, living on leaves close to ground level. Europe, and eastward to Iran and west Siberia. Monoecious holocyclic, with oviparae and apterous males in September-October (Heie 1995). 2n=10.

Macrosiphoniella seriphidii Kadyrbekov Apterae are pale green with silvery sheen, siphunculi dark distally; BL 2.1-2.7 mm. On Artemisia and Seriphidium spp. in Kazakhstan and Xinjiang-Uygur region of China (Kadyrbekov 2000, Kadyrbekov et al. 2002, Kadyrbekov 2017a), and also recorded from Tajikistan (Kadyrbekov 2013c) and Russia (Altai; Kadyrbekov 2014g). It is very similar to M. terraealbae.

Macrosiphoniella sibirica Ivanoskaya Apterae are greenish brown with dark antennae, legs, siphunculi and cauda: BL c. 3.1 mm. On stems and influorescences of Artemisia spp. in western Siberia and Kazakhstan. Close to or synonymous with M. artemisiae.

Macrosiphoniella sieversianae Holman & Szelegiewicz Apterae are green, powdered with whitish wax, with antennae and legs mainly dark, and siphunculi dark distally; BL 2.2-2.8 mm. On upper parts of stems of Artemisia sieversiana in Mongolia, and also on this and other Artemisia spp. in Kazakhstan and western Siberia (Kadyrbekov 2009b, 2014g, 2017a).

Macrosiphoniella sikkimartemisiae Agarwala & Raychaudhuri Appearance in life and size of apterae are unknown; antennae and legs partly pale, siphunculi and cauda dark. On Artemisia sp. in India (Sikkim).

Macrosiphoniella sikhotealiensis Pashchenko Apterae are green, wax-dusted, with dark brown to black appendages; BL c.3.6 mm. Other morphs are unknown. On upper parts of stems of Artemisia stolonifera in east Siberia (Pashchenko 1998b).

Macrosiphoniella silvestrii Roberti Apterae are covered in white wax except for shiny black areas on dorsum; BL 2.8-3.0 mm. On upper parts of stems of Achillea, Anthemis arvensis and Matricaria in southern Italy (Barbagallo 1970, Barbagallo & Stroyan 1982), and now also recorded from Kazakhstan (Kadyrbekov 2018a).

Macrosiphoniella sojaki Holman & Szelegiewicz Appearance of apterae in life is unknown, probably wax-powdered, with blackish brown antennae, legs, siphunculi and cauda; BL 1.7-2.3 mm. On Artemisia rutifolia in Mongolia (original description) and eastern Kazakhstan (Kadyrbekov 2009a), and also reported to occu in Iran (Rezwani 2010).

Macrosiphoniella (Asterobium) soosi Szelegiewicz Apterae are light green with black head and appendages; BL 1.9-3.0 mm. On Galatella sedifolia (= punctata), living onundersides of leaves, in Hungary, and also recorded from G. sedifolia ssp. biflora, G. fastigiiformis G. chromopappa and G. villosa.In Hungary, Romania and Kazakhstan (Kadyrbekov & Aoitzhanova 2005, Kadyrbekov 2017a).

Macrosiphoniella (Chosoniella) spinipes A.N. Basu (Fig.13d) Apterae are shiny green with blackish siphunculi and dusky cauda; BL 1.8-3.8 mm. On undersides of leaves and young shoots of Artemisia vulgaris in north-east India (R.C. Basu & Raychaudhuri 1976b). 2n=10 (Kar et al. 1990).

Macrosiphoniella staegeri Hille Ris Lambers Apterae are dark brown to black, dusted with whitish wax; BL c.2.7-2.9 mm. On leaves and stems of Centaurea spp. in southern Europe, Turkey, Armenia, Iran, Tajikistan and Kazakhstan (Kadyrbekov 2014e). A subspecies was described from Ukraine (M. staegeri ssp. ucrainica Bozhko). Oviparae and alate males were found in Turkey in October (Tuatay & Remaudière 1964); probably anholocyclic in mild climates, e.g Sicily (Patti 1983).

Macrosiphoniella subaequalis Börner Apterae are light brown to pinkish dusted with whitish grey wax, with dark antennae, siphunculi and cauda; legs with femora dark except at base and tibiae with pale middle section. BL 2.1-3.1 mm. On young shoots of Artemisia campestris (where it may be mixed with M. dimidiata), and also recorded from A. rupestris and A. variabilis. In continental Europe (France, Poland, Austria, Switzerland, Italy). It is also reported to occur in Iran (Rezwani 2010). Jorg & Lampel (1988) reviewed information about this species.

Macrosiphoniella subterranea (Koch) Apterae are reddish brown covered with greyish wax except on mid-dorsum and around bases of siphunculi, appendages with contrasting pale and black sections, siphunculi and cauda black; BL 2.6-3.5 mm. On undersides of leaves of Leucanthemum spp., causing yellow spots. Widely distributed in Europe (Heie 1995), in Iran (Rezwani 2010) and Kazakhstan (where it was collected on Chrysanthemum indicum; Kadyrbkov 2017a), and also in North America (Ontario, Pennsylvania, Robinson 1987; Oregon, Jensen et al. 2020). Oviparae and alate males occur in October-November. 2n=12.

Macrosiphoniella sudhakaris Banerjee, A.K. Ghosh & Chakrabarti Apterae are pale green (immatures are white), with mainly dark appendages; BL 1.3-1.6 mm. On inflorescences and young leaves of Achillea millefolium at a high altitude in Uttar Pradesh, India. This is possibly a cold-temperature form of M. millefolii.

Macrosiphoniella sunshine Jensen, Barjadze & Kanturski Apterae are pale green with darker green longitudinal spinal stripe and dark green patches around siphuncular bases, antennae black, legs mainly pale brown, siphunculi pale basally and brown to black distally, and cauda pale; BL c.1.5-2.8 mm. On flower stems and leaves of Eriophyllum lanatum (“Oregon Sunshine”) in western USA (Washington, Oregon, California). Fundatrices were collected in April and oviparae in October (Jensen et al. 2020); males are unknown.

Macrosiphoniella szalaymarzsoi Szelegiewicz Apterae are green, powdered with fine grey wax, with antennae dark distally, legs dark except for bases of femora and middle parts of hind tibiae, and siphunculi dark distally and paler at base; BL 2.0-2.7 mm. On terminal shoots of Artemisia alba ssp. saxatilis (= alba) in Hungary; also collected on A. herba-alba in Israel (BMNH collection, leg. V.F.Eastop), on Artemisia sp. in Iran (Sedighi et al. 2020b), and recorded from several Artemisia and Seriphidium spp. in Kazakhstan (Kadyrbekov 2004b, 2009b, 2014e). Sedighi et al. (2020b) provided a redescription. It is separated in the key from three similar species [cegmidi, longirostrata (= altaica) and taesongsanensis] on the basis of the published descriptions, but the discriminants given are liable to environmentally induced variation, and the group evidently requires further study. 2n=12.

Macrosiphoniella tadshikana Narzikulov Apterae are green, wax-dusted, with siphunculi dark except at bases; BL c.2.6 mm. On upper parts of stems of Artemisia ferganensis in Tajikistan. M. alativica may be closely related.

Macrosiphoniella taesongsanensis Szelegiewicz Apterae are green, powdered with fine grey wax. The head is dark brown, with a colourless, semicircular spot at posterior margin. Antennae and siphunculi are pale basally and dark distally, legs mainly pale; BL c. 1.7-2.0 mm. On terminal parts of shoots of Artemisia spp. in Korea (Lee et al. 2002c), China (Zhong & Liu 1986), east Siberia (Pashchenko 1988a) and east Kazakhstan (Kadyrbekov 2004b). (See comment under M. szalaymarzsoi.)

Macrosiphoniella tanacetaria (Kaltenbach) Apterae are grey-green, dusted with fine wax powder, with black antennae, legs, siphunculi and cauda (see influentialpoints.com/Gallery); BL 3.2-4.1 mm. The principal host plant is Tanacetum vulgare, where it colonises the flowers and flower stems in summer, but the range of reserve hosts seems greater than in most Macrosiphoniella, including records from other Tanacetum spp., species of Achillea, Anthemis, Artemisia, Aster, Bidens, Chamaemelum, Chrysanthemum, Dendranthema and Matricaria, and also Salvia officinalis (Lamiaceae). Monoecious holocyclic on T. vulgare, with alate males. Throughout Europe, Morocco, Israel, Iran, Georgia, Kyrgyzstan, Kazakhstan, Mongolia, west and east Siberia, and introduced to North and South America. Two forms in the Mediterranean region and one in Central Asia are regarded as subspecies; one of these, M. tanacetaria ssp.bonariensisBlanchard, has siphunculi longer relative to the cauda, and is the form that has been introduced into South America. Another, M. tanacetaria ssp. italica Hille Ris Lambers, occurs in Italy, Austria and former Yugoslavia and appears to be specific to Tanacetum corymbosum. The third, M. tanacetaria ssp. tianshanica Narzikulov & Umarov, is described from alatae in Tajikistan which have fewer rhinaria on ANT III and a much shorter cauda. Massonet et al. (2002) and Massonet & Weisser (2004) studied genetic variation in French and German populations, and Loxdale et al. (2011a, b) analysed variation on a smaller spatial scale using microstatellite markers. 2n = 12.

Macrosiphoniella tapuskae (Hottes & Frison) Plate 27d (Fig.5i) Apterae are pale green, in life with small darker green dorsal spots on abdomen anterior to the siphuncular bases (see influentialpoints.com/Gallery); BL 2.5-3.3 mm. On various Anthemidae (Achillea, Anacyclus, Anthemis, Argyranthemum, Artemisia, Chrysanthemum, Matricaria, Tanacetum), usually feeding on lower leaves. In Europe, and eastward to north-west, central and southern Russia, south-west and Central Asia, North Africa and North America, and more recently recorded from Argentina (Ortego et al. 2006). Monoecious holocyclic with oviparae and apterous males in September-November. [Kadyrbekov (2019) suggests that M. tapuskae should be treated as a subspecies of M. aktashica (Nevsky).] 2n=12.

Macrosiphoniella teriolana Hille Ris Lambers (Fig.13i) Apterae are yellowish green, powdered with grey wax, with appengages all black except for base of ANT III; BL c.1.7-2.0 mm. On Artemisia campestris in northern Italy. It is only known definitely from the type locality; other records refer to M. dimidiata.

Macrosiphoniella terraealbae Kadyrbekov Apterae are pale green with silvery sheen, antennae and legs brown to dark brown, siphunculi dark except basally; BL 1.5-2.4 mm. On Artemisia terrae-albae in Kazakhstan, south-west Russia and Xinjiang-Uygur region of China (Kadyrbekov 2000, 2014g; Kadyrbekov et al. 2002). Kadyrbekov (2017a) records it from several other Artemisia/Seriphidium spp. It is very similar to M. seriphidii.

Macrosiphoniella (Papillomyzus) tuberculata (Nevsky) Apterae are green, with siphunculi brown, paler at base; BL 1.8-2.9 mm. On stems and undersides of leaves of Picnomon acarna, and also recorded from Carduus and Cousinia spp. (Kadyrbekov 2017a). In the Middle East and Central Asia (Israel, Iran, Syria, Turkey, Kazakhstan, Uzbekistan). Monoecious holocyclic in Turkey with oviparae and alate males in October (Tuatay & Remaudière 1965).

Macrosiphoniella (Papillomyzus) tuberculatumartemisicola Bozhko Apterae are green, powdered with grey wax, with appendages mainly dark; BL c. 2.8 mm. In small colonies on flower stems of Seriphidium tauricum in Crimea, and also recorded from Iran (Rezwani 1994).

Macrosiphoniella umarovi Narzikulov (Fig.13k) Apterae are green, wax-dusted, with mainly brown appendages but pale spots at apices of femora, and pale cauda; BL c. 1.9-2.2 mm. On leaves and stems of an Artemisia sp. in Afghanistan. Keyed on basis of specimens in BMNH collection agreeing with the original description, collected in 1975 from Artemisia sp. in Afghanistan (leg. R. van den Bosch).

Macrosiphoniella usquertensis Hille Ris Lambers Apterae are brownish, powdered with greyish wax, with antennae and legs yellow and black, siphunculi and cauda black (see influentialpoints.com/Gallery); BL 2.4-3.2 mm. Principal host plant is Achillea millefolium, but also on other Achillea spp., and sometimes found on Artemisia campestris. It lives on the lower leaves of Achillea, which eventually turn brownish and wither (Heie 1995). Widely distributed in Europe, in Iran and Kazakhstan (Kadyrbekov 2017a), and an alata has been found in Quebec, Canada (Robinson 1987). Oviparae and alate males occur in north-west Europe in late August-October. Sobhani (1970) studied the biology, life cycle and morphology of M. usquertensis in Germany, and Sobhani & Iglisch (1972) compared morphology of males with that of M. ptarmicae.

Macrosiphoniella vallesiacae Jörg & Lampel Apterae are dark green powdered with greyish white wax, with appendages mainly pale to dark brown; BL 1.7-2.4 mm. On apices of young shoots of Artemisia vallesiaca in alpine Switzerland and Italy (Lampel & Meier 2007), and subsequently reported from Artemisia (= Seriphidium) aucheri and A. (S.) sieberi in Iran (Mehrparvar 2017).

Macrosiphoniella (Asterobium) victoriae Kadyrbekov Apterae are dark green, with mainly dark brown appendages; BL 2.2-2.4 mm. On Saussurea elegans in Kazakhstan (Kadyrbekov 1999a), and subsequently recorded from Jurinea (=Pilostemon) filifolia (Kadyrbekov 2014d).

Macrosiphoniella xeranthemi Bozhko Apterae are pale brown with black appendages; BL 2.2-3.0 mm. On Xeranthemum spp.in eastern Europe (Bulgaria, Czech Republic, Ukraine) and Kazkhstan (Kadyrbekov 2017a).

Macrosiphoniella xinjiangica Kadyrbekov, Renxin & Shao Apterae are bright green; BL 2.5-3.0 mm. On stems of Artemisia dracunculus in Xinjiang-Uygur region of western China (Kadyrbekov et al. 2002).

Macrosiphoniella yangi Takahashi Only apparently known from original collection, from “a composite”. Placed in Asterobium by Szelegiewicz (1980), but it does not seem to belong there, and it is unclear where the host record from Aster sp. originated.

Macrosiphoniella (Asterobium) yomenae (Shinji) (Fig.15c) Apterae are dirty yellow-brown to green with brown-black head and pronotum, brown presiphuncluar sclerites, and brown-black appendages; BL 2.6-3.5 mm. On Aster and related genera (Boltonia, Erigeron, Galatella, Kalimeris) in Japan, Korea, China and east Siberia. Also recorded from Hieracium sp. in east Siberia (Pashchenko 1988a), and from Artemisia and Chrysanthemum in Korea (Lee et al. 2002c, and BMNH collection), which may indicate a wider host range, but this needs further confirmation. 2n=12.

Macrosiphoniella (Sinosiphoniella) yomogicola (Matsumura) Apterae are grey-green to blackish or rust brown, with dark transverse bars on dorsal abdomen, and dark antennae, legs, siphunculi and cauda; BL 2.0-3.2 mm. On Artemisia spp., attended by ants under earthen shelters (Miyazaki 1971); ant attendance has been shown to be necessary for the aphid colony’s survival (Watanabe et al. 2016, 2019), and influences the proportion of the different colour morphs within a colony (Hasegawa et al. 2019). In Japan, China, and east Siberia (Sakhalin, Kuril Islands). Also recorded from Dendranthema indica. A record from Ixeridium dentatum in Korea is probably based on a misidentification (Holman et al. 2006a).

Macrosiphoniella yomogifoliae (Shinji) Apterae are yellow-green to deep mid‑green, powdered with grey wax, with mainly dark antennae and legs, black siphunculi and brown cauda; BL 2.0-3.5 mm. On Artemisia spp., Bidens spp., Chrysanthemum spp. (morifolium, nipponicum) and Tanacetum spp. in east and south-east Asia (Japan, Korea, China, east Siberia, India, Nepal, Taiwan, Vietnam, Malaysia, Indonesia). Also recorded from Gnaphalium multiceps (= affine) in Korea (Lee et al. 2002c). Monoecious holocyclic with sexual morphs in October; an ovipara was described from Nipponathemum nipponicum in Japan by Miyazaki (1971), and oviparae and alate males collected on Artemisia vulgaris in north-west India were described by Maity & Chakrabarti (1981b). Closely related to European M. artemisiae, but more often confused with M. pseudoartemisiae (q.v.). 2n=12.

Macrosiphoniella zeya Pashchenko Apterae are brown, BL c.1.9-2.0 mm; immatures are greenish pinky brown. On upper parts of stems of Artemisia messerschmidtiana in east Siberia (Pashchenko 1998b).

Macrosiphum Oestlund

Aphidinae: Macrosiphini

About 120 species with long antennae and legs, long siphunculi usually with polygonal reticulation on distal 5‑30% (mostly 8‑18% in apterae and 15‑25% in alatae), hairs of medium length, and little or no dorsal abdominal pigmentation. Several of the well‑known species (rosae, euphorbiae, pallidum) alternate from Rosa to herbaceous secondary hosts, but most species are monoecious holocyclic on a wide variety of herbs and shrubs, with a few on trees, including subgenus Neocorylobium which has six species associated with Betulaceae. Males are usually alate. About half the species are described from North America, although some of these have seldom or never been recognized again and may really be synonyms of better known species. About 36 species are known from Europe and the remainder are from central and east Asia. Three South American species may also belong in Macrosiphum. Sitobion, often treated as a subgenus of Macrosiphum, is now regarded as a separate genus. Accounts of Macrosiphum are available for Europe (Hille Ris Lambers 1939, Heie 1994), UK and Ireland (Blackman 2010), Germany (Muller 1969b), Switzerland (Meier 1961, Lampel & Meier 2007), Central Asia (Nevsky 1929a, Narzikulov & Umarov 1969), Japan (Miyazaki 1971), Korea (S. Lee et al. 2001b, 2009), China (Tao 1963) and India (David 1976). North American Macrosiphum were reviewed by Palmer (1952 ‑ Rocky Mountain region), Hottes & Frison (1931 ‑ Illinois), Patch (1919 ‑ eastern USA), and Soliman (1927 ‑ California). MacGillivray (1968) and Jensen (1998) gave more recent information on some North American species, and Jensen (1997) clarified the distinction between Macrosiphum and Sitobion. Colour photographs of many western North American species are available at aphidtrek.org. Fern-feeding species, which seem to form a distinct group within Macrosiphum (or possibly a separate genus with 2n=16, whereas Macrosiphum typically have 2n=10) were reviewed and keyed by Jensen & Holman (2000).

Macrosiphum adianti (Oestlund) (Fig.43f) Apterae are yellow, greenish or pink; BL 1.0-2.1 mm. On Adiantum spp., esp. A. pedatum, in North America. Monoecious holocyclic with apterous males. Records from other ferns are probably due to misidentification (Jensen & Holman 2000).

Macrosiphum aetheocornum Smith & Knowlton (Fig.30a) Apterae are greenish (see aphidtrek.org); BL c.3.0-3.5 mm. On Geranium spp., especially G. richardsonii, in western USA. This species is unusual in having some specimens (but not all) with hairs on siphunculi (Palmer 1952, BMNH collection, leg. G.F. Knowlton, and aphidtrek.org). Jensen (2022) added detail on this species, which is morphologically very similar to M. ginajo, although their respective host plants are taxonomically distant.

Macrosiphum agrimoniellum (Cockerell) Apterae are broadly spindle-shaped, light apple-green with apices of antennal segments and tibiae darker; BL 2.8-3.3 mm. Forming large colonies on flower stems of Agrimonia eupatoria, and also recorded from Potentilla sp. (aphidtrek.org). Widely distributed in North America (New Mexico, Utah, Illinois, New York, Massachusetts, New Brunswick).

Macrosiphum albertinae Hille Ris Lambers Apterae are apple-green with antennae, tibiae and siphunculi mainly dark, and cauda pale green; BL 2.4-2.9 mm. In small numbers on upper sides of leaves of Thesium intermedium in northern Italy, former Yugoslavia (BMNH collection, leg. V.F. Eastop) and Czech Republic (BMNH collection, leg. H.L.G. Stroyan), and on Th. divaricatum in France (BMNH collection, leg. G. Remaudière). Alatae are undescribed.

Macrosiphum albifrons Essig Lupin Aphid Apterae are pale bluish-grey-green, dusted with white wax (see influentialpoints.com/Gallery); BL 3.2-5.1 mm. Alatae have a brown pterothorax. On leaves, stems and flowers of Lupinus spp., and also recorded from Thermopsis montana. In North America, and introduced to Europe and South America (Brazil). Monoecious holocyclic with alate males. See also Blackman & Eastop (2000). 2n=10.

Macrosiphum alpinum Meier Apterae are pale yellowish green; BL 4.3-4.8 mm. On Senecio alpinus, feeding on undersides of middle and lower leaves, and also collected from S. fuchsii (= ovatus). Described from Switzerland, and also recorded from Cicerbita alpina in Ukraine (Chumak 2004). Monoecious holocyclic with sexual morphs in October (Lampel & Meier 2007). Very similar to, and possibly synonymous with, M. polanense.

Macrosiphum amelanchiericolens Patch Described from Amelanchier spicata in Maine (Patch 1919), but not recorded since, and the types are lost (MacGillivray 1968). Apparently a distinctive species, with numerous secondary rhinaria on ANT III in both apterae and alatae. Appearance in life and biology are unknown.

Macrosiphum atragenae Holman Apterae are whitish green with dark apices to antennal segments, legs and cauda; BL 3.7-4.55 mm. In small groups on undersides of leaves of Clematis alpina in the Carpathians (Slovakia, Romania). It is also recorded from C. glauca in Kazakhstan (Kadyrbekov 2013d).

Macrosiphum audeni Macdougall Apterae are pale green; BL 2.3-2.9 mm. On upper sides of leaves of Nuphar lutea in north-western USA (Washington) and Canada (British Columbia). This species is very similar to M. euphorbiae, but specimens in the BMNH collection (leg. A.P. Macdougall) have a relatively longer antennal terminal process and shorter hind tarsus II.

Macrosiphum badium Jensen Apterae are dark reddish brown, brick-coloured (see aphidtrek.org); BL 2.3-3.1 mm. On Maianthemum spp. in western USA. Monoecious holocyclic, with oviparae and both apterous and alate males in October (Jensen 2000).

Macrosiphum bisensoriatum Macdougall Apterae are dark green; BL c.3.4 mm. On undersides of leaves of Ribes lacustre in British Columbia, Canada. The life cycle is unknown.Macrosiphum bupleuri Kadyrbekov Apterae are yellow or yellow-green, with dark tips to siphunculi; BL 2.6-3.6 mm. On inflorescences and undersides of leaves of Bupleurum aureum in south-east Kazakhstan (Kadyrbekov 2000).

Macrosiphum californicum (Clarke) Apterae are elongate spindle-shaped, green with long, yellowish green, dusky-tipped siphunculi, and sometimes with a longitudinal dorsal stripe of dark green internal pigment (see aphidtrek.org); BL 1.9-2.6 mm. Alatae are rather pale and have only 5-10 secondary rhinaria, confined to basal half of ANT III. On young growth of Salix spp., common and widely distributed in North America, and also recorded from Honduras (Evans & Halbert 2007). Monoecious holocyclic; oviparae and alate males occur in October (Palmer 1952). Possibly there are two Salix-feeding species being confused under this name (see aphidtrek.org). 2n=10.

Macrosiphum (Neocorylobium) carpinicolens Patch Apterae are green with dark legs, antennae and distal parts of siphunculi; BL 1.8-2.6 mm. On young shoots and leaves of Carpinus caroliniana in eastern North America. Monoecious holocyclic; Hottes & Frison (1931) described the fundatrix, alatae vivipara, ovipara and alate male. The sexuales occur in October. See also MacGillivray (1968).

Macrosiphum centranthi Theobald Apterae are whitish green, yellowish green or green with darker green spinal stripe, with dark brown eyes (not red as in euphorbiae), and femora and siphunculi darker towards apices; BL 2.0-3.6 mm. Alatae have 17-26 secondary rhinaria on ANT III. On leaves and stems of Valerianaceae (Centranthus, Valeriana), sometimes in mixed colonies with M. rosae, and colonies also occur sporadically on various other plants. Europe (UK, Switzerland), Turkey (Şenol et al. 2017; requires confirmation), India, and Africa (Mozambique, Tanzania, South Africa). Indian populations seem more polyphagous, with records fron Chrysanthemum, Cineraria, Mangifera, Rosa, Tibouchina, etc. (David 1976).Monoecious holocyclic in Europe, with alate males (Meier 1961), probably anholocyclic elsewhere. 2n=10.

Macrosiphum cerinthiacum Börner Apterae are pale yellow with dark brown apices to antennae, legs and siphunculi, and dark “knees”; BL c.3.5 mm. Alatae have c.35-40 secondary rhinaria on ANT III. On undersides of leaves of Cerinthe minor in France (BMNH collection, leg. J.H. Martin), Spain, Austria, Czech Republic, Slovakia, Hungary, Ukraine and Turkey. The life cycle is unknown.

Macrosiphum cholodkovskyi (Mordvilko) (= Macrosiphum corallinum Theobald) Apterae are yellow-green to dark blue-green or vivid coral-pink to red, with femora and siphunculi dark distally (see influentialpoints.com/Gallery); BL 3.1-5.1 mm. Immatures often have a darker green mid-dorsal stripe. Alatae have 9-28 secondary rhinaria on ANT III. On upper leaves, stems and inflorescences of Filipendula ulmaria, and occasionally on Valeriana spp. In Europe, eastward to Turkey, Iran, west Siberia, Transcaucasus and Kazakhstan. Monoecious holocyclic, with alate males (Heie 1994). 2n=10.

Macrosiphum chukotense Stekolshchikov & Khureva Apterae are very variable in colour, yellow-green with a dark green spinal stripe, green, purple, reddish purple or red-brown, with appendages mainly pale; BL 2.4-3.8 mm. On shoot apices, stems and inflorescences of plants in numerous families; Asteraceae (Arnica, Artemisia, Hieracium, Petasites), Brassicaceae (Arabis, Raphanus), Caryophyllaceae (Dianthus, Eremogone), Ericaceae (Vaccinium), Fabaceae (Astragalus), ?Onagraceae (Epilobium – but possibly this is M. tinctum?), Orobanchaceae (Pedicularis), Polemoniaceae (Polemonium), Polygonaceae (Persicaria), Primulaceae (Cortusa), Ranunculaceae (Anemone, Clematis, Ranunculus), Rosaceae (Comarum, Rubus), Rubiaceae (Galium, Rubus), Saxifragaceae (Saxifraga) and Valerianaceae (Valeriana). In north-east Siberia (Chukotka) and north-west Russia (Nenets, Murmansk, Krasnoyarsk Krai; Stekolshchikov 2017, Stekolshchikov & Buga 2018, Stekolshchikov & Khureva 2020). Fundatrices were collected from plants in five of these families, confirming this species’ polyphagy. Morphologically it is close to M. euphorbiae, but there is no host alternation and the life cycle is abbreviated, with oviparae and alate males in late July-early August (Stekolshchikov & Khureva 2015). There is also a close parallel to M. valerianae, a similar species in western North America that also lives without host alternation on a wide variety of host plants.

Macrosiphum claytoniae Jensen Apterae are dark reddish or greenish brown (see aphidtrek.org); BL 1.8-2.7 mm. Immatures are dusted with purplish or bluish grey wax. On Claytonia spp., especially C. sibirica, in western USA (Colorado, Oregon) and Canada (British Columbia). Apparently entirely anholocyclic, surviving the winter in mild humid areas west of the Cascade Mountains (Jensen 2000). 2n=16.

Macrosiphum clematifoliae Shinji Apterae are rather large, spindle-shaped, pale yellow green to green with a darker blue-green spinal stripe, and black siphunculi that are pale at their bases; BL 2.8-3.5 mm. On Clematis spp., feeding on both sides of leaves. In Japan, China, Korea and Taiwan. Monoecious holocyclic in Japan (Miyazaki 1971). 2n=18 (the karyotype suggests that this species may be a Sitobion).

Macrosiphum clematophagum Zhang, Chen, Zhong & Li Apterae are green; BL c. 2.3 mm. On leaves of Clematis florida in Gansu Province (Xinglongshan mountains, 2200m), China (Zhang 1999).

Macrosiphum clum Jensen Apterae are mid-green (see aphidtrek.org); BL 1.7-2.6 mm. On leaves and stems of Clematis ligusticifolia, in low densities and usually in exposed and dry or rocky sites. In north-west USA (Oregon, Idaho, Washington). Monoecious holocyclic; oviparae were collected in October (Jensen 2015).

Macrosiphum clydesmithi Robinson (Fig.43g) Apterae are pale green (see aphidtrek.org); BL 1.7-3.0 mm. Heteroecious holocyclic, migrating from Holodiscus discolor to Pteridium aquilinum in western North America, from Mexico to Washington. In western Oregon remigration to Holodiscus occurs in October-November, a few weeks earlier than M. pteridis, which utilises the same hosts (Jensen & Holman 2000). 2n=16.

Macrosiphum constrictum Patch Colour of apterae in life is unknown, and BL unrecorded. On Pedicularis sp. and Saxifraga sp. on St Paul and St George (Pribilof Islands, Bering Sea). Monoecious holocyclic; oviparae occurred (on Pedicularis) in June (original description). The generic position is uncertain.

Macrosiphum corallorhizae Cockerell Apterae are green, with long siphunculi, blackish in middle and at apices; BL 2.5-3.0 mm. On Corallorhiza spp. in USA (New Mexico) and Canada (British Columbia; Forbes & Chan 1989).

Macrosiphum cornifoliae (Shinji) Apterae are translucent, whitish, with banded antennae; BL c. 2.2mm. Alatae are pale yellow-green with yellow-brown thorax and dusky appendages (Moritsu 1983). On undersides of leaves of Cornus spp. in Japan and Korea. Presumably monoecious holocyclic; oviparae were collected in November (Miyazaki 1971). [An undescribed Macrosiphum sp. from Cornus stolonifera in British Columbia, Canada may key out to this species, but is distinct.] 2n=14 (based on 2n male =13; Shinji 1931).

Macrosiphum corydalis (Oestlund) Apterae are green, BL not recorded. Very similar and almost indistinguishable in morphology to M. euphorbiae, but monoecious holocyclic, with apterous males. On Fumariaceae (Dicentra formosa, Corydalis aurea) in North America; so far known from British Columbia, Washington, California, Minnesota Wisconsin and Maine. In late summer it can occur in dense colonies with M. dicentrae on flower stalks of Dicentra (Jensen & Chan 2009). [This species was for many years known only as alatae, and has been treated in catalogues as a synonym of Acyrthosiphon pisum.] 2n=10.

Macrosiphum (Neocorylobium) coryli Davis Apterae have reddish brown head and prothorax, abdomen mainly green suffused with dark brown laterally and posteriorly, antennae and legs mainly dark, and siphunculi black contrasting with the pale cauda (Davis 1914, and see influential points/Gallery); BL 1.6-2.2 mm. On shoot tips and undersides of young leaves of Corylus americana and C. cornuta (incl. var california) in USA (more northerly states) and across Canada. Presumably monoecious holocyclic, but life cycle and sexuales are apparently unknown.

Macrosiphum (Unisitobion) corylicola Shinji Apterae are dull yellow to yellowish green, often suffused dorsally with dark brown, antennae, legs and siphunculi mainly black, cauda dusky; BL 2.5-3.1 mm (Miyazaki 1971). On Corylus spp. and Carpinus laxiflora in Japan, east Siberia and Korea (Paik 1972). Biology and sexuales are unrecorded.

Macrosiphum creelii Davis Apterae are pale green, pink or yellow; BL 2.3-4.0 mm. On various Leguminosae/Fabaceae (Lathyrus, Lens, Medicago, Phaseolus, Pisum, Vicia) in western North America. Halfhill (1982) studied its host plant preferences and temperature relations. Apparently monoecious holocyclic, but males are not yet described. 2n=10.

Macrosiphum cyatheae (Holman) Apterae are pale green, with dark brown antennae and tibiae; BL 1.6-3.0 mm. Singly or in small colonies on undersides of fronds of Cyathea balanocarpa and Pteridium caudatum in Cuba (original description, as Acyrthosiphon), and subsequently found on Pteridium in Venezuela and Costa Rica (Villalobos Muller et al. 2010), and on Pteridium sp. and unidentified epiphytic ferns in eastern Mexico (Jensen & Holman 2000).

Macrosiphum cystopteris Robinson Apterae are dark green to greenish black; BL 2.2-2.5 mm. On undersides of fronds of Cystopteris bulbifera in USA (Pennsylvania), and also recorded from Cystopteris sp. in Mexico (Jensen & Holman 2000). Monoecious holocyclic, with oviparae in late September (in USA).

Macrosiphum daphinidis Börner Apterae are pale yellowish or whitish green with slightly darker spinal stripe, with dark brown eyes (cf. euphorbiae), entirely pale femora, and siphunculi only slightly darker at apices (see influentialpoints.com/Gallery); BL 2.4-4.2 mm. Alatae have 26-47 secondary rhinaria on ANT III. On Daphne spp., especially D. mezereum, forming small, rather loose colonies on growing buds and shoots, and scattered on undersides of leaves (Watson 1982). In Europe, and introduced to north-western North America (Oregon, British Columbia). Monoecious holocyclic, with oviparae and alate males (Heie 1994) in September. 2n=10.

Macrosiphum dewsler Jensen Apterae are matte mid-green, paler marginally and ventrally, with pale brown appendages (see aphidtrek.org); BL 1.8-2.8 mm. Alatae have 4-9 secondary rhinaria on ANT III. On Linum lewisii in dry montane forest regions of western USA (Oregon, New Mexico). Oviparae and alate males were collected in Oregon in October (Jensen 2017).

Macrosiphum dicentrae Jensen & Chan Apterae are yellowish to mid-green dusted lightly with wax; BL 2.4-3.8 mm. Alatae have 4-13 secondary rhinaria on ANT III. Monoecious holocyclic on Fumariaceae, mostly feeding on Dicentra formosa; also found on Dicentra sp., Corydalis scouleri and C. aurea. In western North America. Oviparae and alate males appear in September-October (Jensen & Chan 2009). 2n=16.

Macrosiphum diervillae Patch Apterae are milky white with pale appendages; BL 2.9-3.4 mm. Living singly on undersides of leaves of Diervilla lonicera in north-eastern USA and eastern Canada. Monoecious holocyclic; oviparae and males (immature) were collected in September in New Brunswick (MacGillivray 1968).

Macrosiphum doronicicola Leclant Apterae are spindle-shaped, green to greenish white with dark apices to appendages: BL 2.9-4.2 mm. Forming small colonies on flower stems and flowerheads of Doronicum austriacum in south-west Europe (France, Spain). Monoecious holocyclic, apparently with an abbreviated parthenogenetic phase; fundatrices were collected in mid-June, and oviparae and alate males were present in mid-July (original description).

Macrosiphum dryopteridis (Holman) Apterae are yellowish green to bright or pale green, sometimes pale yellow or pinkish; BL 1.7-2.4 mm. On various ferns (Athyrium, Dryopteris, Gymnocarpium, Thelypteris), feeding on undersides of fronds along veins. In north and central Europe. Monoecious holocyclic with oviparae and alate males in September-October (original description). Possibly there are host-specific populations; Müller (1988) found that a bright green form on Gymnocarpium dryopteris and a yellowish green form on Athyrium filix-femina bred true in the laboratory, with hybrids having intermediate colouration.

Macrosiphum dzhibladzeae Barjadze Apterae are yellow with a green spinal stripe; BL 3.1-3.9 mm. In dense colonies on the stems and undersides of leaves of Euphorbia macroceras in Georgia (Barjadze & Gratiashvili 2010). Monoecious holocyclic; oviparae and alate males collected in mid-August were described by Barjadze et al. (2018a).

Macrosiphum eastopi Barjadze & Blackman Apterae are pale green with pale cauda and black-tipped siphunculi; BL 2.8-3.5 mm. In sparse colonies on the undersides of leaves and stems of Oberna multifida, only known from western Georgia. Alate males were found in mid-August (Barjadze et al. 2017a).

Macrosiphum echinocysti Bartholomew Apterae are forest green, with slight whitish bloom, head paler, eyes dark red, and with a dark green spinal stripe; BL 2.4-3.5 mm. On Echinocystis sp. in California, USA (San Francisco), in large numbers on the undersides of leaves and dropping readily when disturbed (original description). The author could not find this species again when he visited the site the following year, and it has not been collected since.

Macrosiphum edrossi Essig Apterae are pale green, with mainly brown appendages; BL 2.3-3.3 mm. On Baccharis latifolia in Peru (Nieto Nafría & Ortego 2018). It was originally described from an ?Urtica sp., but these were possibly vagrants. Nieto Nafría & Ortego (2018) provided redescriptions of apterae and alatae. Probably monoecious holocyclic on Baccharis, but sexual morphs are unknown.

Macrosiphum equiseti (Holman) Apterae are spindle-shaped, uniformly green or pinkish with apices of appendages dark brown to blackish; BL 1.6-2.6 mm. In small groups on undersides of branches of Equisetum spp. in Europe and Canada (British Columbia). Monoecious holocyclic on E. sylvaticum in Czech Republic, with oviparae and apterous males in September (original description). 2n=16.

Macrosiphum euphorbiae (Thomas) Potato Aphid Plate 25g Apterae are broadly spindle-shaped, usually green, sometimes yellowish, pink or magenta, with red eyes, and femora and siphunculi pale or only slightly darker towards apices (see influentialpoints.com/Gallery); BL 1.7-3.6 mm. Immatures are rather long-bodied, paler than adults but with a dark spinal stripe, and are dusted with greyish wax. Alatae are rather pale, with a pale greenish to yellow-brown thorax, only the antennae and siphunculi noticeably darker than in apterae, and have 12-21 secondary rhinaria on ANT III. Of North American origin, now almost world-wide. Heteroecious holocyclic with a sexual phase on Rosa in north-eastern USA, but elsewhere probably mainly or entirely anholocyclic on secondary hosts in more than 20 different plant families. [A very similar undescribed species is monoecious holocyclic on Oplopanax horridum in the Cascade Mountains of north-western USA (see aphidtrek.org).] Raboudi et al. (2011, 2012) studied the genetic diversity of populations of M. euphorbiae on different host plants in Tunisia, and Clarke et al. (2017) demonstrated variable susceptibility to the parasitoid Aphidius ervi. See also Blackman & Eastop (2000, p.291). 2n=10.

Macrosiphum euphorbiellum Theobald (= M. amygdaloides Theobald; synonymy established by Watson 1982, and published in Blackman 2010) Apterae are broadly spindle-shaped and very variable in colour – yellow green, mid green, pink, magenta or wine red – with black apices to antennae, femora, tibiae and siphunculi (see influentialpoints.com/Gallery); BL 1.5-3.7 mm. On stems and flowerheads of Euphorbia spp. in Europe (southern England, southern Ireland, Germany, Austria, Switzerland, Corsica, Italy and Turkey), and there is also a record from Washington, USA (Skvarla et al. 2017). Monoecious holocyclic with red oviparae and red alate males occurring in late September-October (in Switzerland; Meier 1961). 2n=10.

Macrosiphum fagopyri A.K.Ghosh & Raychaudhuri Apterae are pale yellowish with yellowish brown siphunculi; BL 4.2-4.8 mm On young growth and undersides of leaves ofFagopyrum sp., and later recorded from F. cymosum (Raychaudhuri et al. 1980), in north-east India. Males possibly of this species were described by David (1976).

Macrosiphum floridae (Ashmead) Apterae are shining palegreen, with pale siphunculi. On Rosa laevigata in Florida.

Macrosiphum funestum (Macchiati) Apterae are rather dull mid to dark green, or magenta to reddish brown, with mainly dark antennae and siphunculi (see influentialpoints.com/Gallery); BL 1.9-4.0 mm. On young shoots and leaves of Rubus spp., esp. R. fruticosus s. lat. Throughout Europe, eastward to Moldova, Turkey and Iran, and also recorded from Canada. Monoecious holocyclic, with alate males; probably anholocyclic in areas with mild winters. 2n=10.

Macrosiphum fuscicornis Macdougall Apterae are dark olive-green, with red eyes and uniformly dark tibiae; BL c.3.2 mm. Found on stems and leaf buds of Epilobium angustifolium in Canada (British Columbia), and apparently not recorded since its original description.

Macrosiphum garyreed Jensen Apterae are green or brownish pink, somewhat shiny, with antennal joints and apices of siphunculi and legs dark brown (see aphidtrek.org); BL 1.9-3.0 mm. Alatae have c.15 secondary rhinaria on ANT III. On stems and leaves of Geum triflorum in dry forest regions of western USA (Oregon, Idaho). Sexual morphs occur in early October, males being apterous (Jensen 2017).

Macrosiphum gaurae (Williams) Apterae are bright green, orange-red or pink, with dark antennae, mainly dark brown to black siphunculi and a dusky cauda (see influentialpoints.com/Gallery); BL 2.7-3.9 mm. On Gaura and Oenothera spp., widely distributed in North America. Oviparae have been collected on Gaura in Oregon in October (aphidtrek.org). MacGillivray (1968) provided a redescription. Records of the very similar M. pallidum from Onagraceae should probably all be referred to this species.

Macrosiphum gei (Koch) Apterae are spindle-shaped, mid-green to bluish green, or mauve with green mottlings to wine red, with femora and siphunculi dark at apices (see influentialpoints.com/Gallery); BL 1.9-5.4 mm. Alatae have 8-26 secondary rhinaria on ANT III. On Geum spp., especially G. urbanum, where it forms dense colonies on upper parts of flower stems, and it can also occur on certain Umbelliferae/Apiaceae (Anthriscus, Chaerophyllum, Conium, Myrrhis, Torilis), and occasionally on Caryophyllaceae, usually scattered or in small colonies on the undersides of the leaves. Dense colonies form on upper parts of stems of G. urbanum during flowering. In Europe, Iran, west Siberia, and introduced to North America (USA, Canada, Bermuda). Monoecious holocyclic on Geum, with alate males. 2n=10.

Macrosiphum geranii (Oestlund) Apterae have a glaucous appearance due to a covering of fine white powder; BL of apterae is unrecorded, BL of alatae 2.3-3.3 mm (Oestlund 1887, and BMNH collection, leg. J.O. Pepper). On Geranium maculatum in eastern USA. Records from western USA may all be referable to M. euphorbiae, and aphids from a cultivated Geranium sp. in Manitoba (Robinson & Bradley 1965) are an undescribed Macrosiphum species with shorter siphunculi (BMNH collection, leg. A.G. Robinson). 2n=10.

Macrosiphum ginajo Jensen Apterae are medium green to reddish brown; BL 2.5-3.8 mm. Apterae have 2-10 and alatae 11-19 secondary rhinaria on ANT III. Monoecious holocyclic with winged males, on Silene oregana and other species of this genus with glandular-sticky hairs. Recorded in western USA. It is morphologically very similar to M. aetheocornum, but with different host plants.

Macrosiphum glawatz Jensen Apterae are mid-green, pinkish red or red mottled with green, with siphunculi pale to dusky, darker at apices (see aphidtrek.org); BL 1.8-3.1 mm. Alatae have 18-35 secondary rhinaria on ANT III. On Potentilla gracilis throughout the growing season, in meadows and edges of dry forest (Jensen & Rorabaugh 2020). In Oregon and Washington east of the Cascade Mountains, USA . Unusually for a Macrosiphum, colonies are almost always ant-attended. Probably monoecious holocyclic, but sexual morphs are still to be found.

Macrosiphum hamiltoni Robinson Apterae are pale green with antennal joints, tarsi and tips of siphunculi dusky to black; BL 2.2-3.1 mm. Originally described from specimens collected on Humulus lupulus in Manitoba, Canada, but since collected from Cornus spp. in Manitoba and New Brunswick. From dates of collection host alternation seems unlikely, so Cornus is probably the usual host. 2n=10.

Macrosiphum hartigi Hille Ris Lambers Apterae are green, with a faint greyish wax bloom, antennae and legs yellowish brown, with apices of tibiae and tarsi brown, siphunculi green with black apices, cauda green; BL c.3.6 mm. On Silene vulgaris, feeding on upper parts of flowerstalks and under leaves (Hille Ris Lambers 1931b, as M. montanum). In southern Europe (Italy, Austria, Switzerland). Oviparae (vagrants?) have been recorded in October from Linum alpinum in Switzerland (Lampel & Meier 2007).

Macrosiphum helianthi (Tao) Apterae are yellowish green with antennae, apices of femora and tibiae, tarsi and whole of siphunculi blackish brown; BL c. 2.4 mm. Described from specimens collected on Helianthus annuus in Szechuan Province, China, in 1936, and apparently not recorded since.

Macrosiphum hellebori Theobald & Walton Apterae are yellow-green with darker marbling, with dark apices to antennal segments, femora, tibiae and siphunculi (see influentialpoints.com/Gallery); BL 1.7-4.3 mm. On undersides of leaves of Helleborus spp. in Europe, Georgia (Barjadze et al. 2009), and introduced to New Zealand, Australia (Valenzuela et al. 2009a) and USA (Washington; Skvarla et al. 2017). Monoecious holocyclic, with oviparae and alate males in September in continental Europe, but overwintering mainly as viviparae in UK. Badmin (1991) studied the population ecology of this species in south-east England. 2n=10.

Macrosiphum holmani Leclant Apterae are green or red, with brown head, antennae, legs, siphunculi and cauda; BL 2.1-3.1 mm. On flower-stalks and flower-heads of Sanicula europaea in Europe (Corsica, northern Italy, Czech Republic, Romania). Monoecious holocyclic, with oviparae and alate males in late August-October (original description).

Macrosiphum holodisci Jensen Apterae are white (for fundatrix see aphidtrek.org), BL 2.0-3.4 mm. Monoecious holocyclic on Holodiscus discolor in north-western USA (Idaho, Oregon). Oviparae and apterous males occur in September-October (Jensen 2000).

Macrosiphum impatientis Williams Apterae are shining green to dark green with strikingly black siphunculi (see influentialpoints.com/Gallery); BL 1.7-3.8 mm. In north-eastern and midwestern USA. Heteroecious holocyclic, migrating from Rosa spp. (multiflora, carolina) to Impatiens spp. (Jensen 1998). Some records of M. pallidum on Impatiens (e.g. MacGillivray 1968) may be referable to this species. Records from Turkey (e.g. Akyurek et al. 2010) are probably due to misidentification. 2n=10.

Macrosiphum inexspectatum Leclant Apterae are green, similar in colour to host plant; BL 1.6-3.2 mm. On Euphorbia insularis, and apparently specific to this host,in Corsica. Monoecious holocyclic with apterous males.

Macrosiphum insularis (Pergande) Apterae are pale brownish yellow, with appendages mainly pale; BL 2.5-3.4 mm. On undersides of leaves of Asparagaceae (genera formerly placed in Comvallariaceae; Maianthemum, Polygonatum, Streptopus)in western North America (Colorado, Arkansas, Utah, British Columbia and St. Paul Island). Monoecious holocyclic, with oviparae and alate males in September (Palmer 1952, as M. yagasogae Hottes; see also Jensen 2000).

Macrosiphum (Unisitobion) isodonis (Sorin) Apterae are yellowish green, with broad lateral longitudinal dark stripes, black antennae, legs and siphunculi, and a pale cauda; BL 1.9-2.2 mm. Immatures are whitish (Moritsu 1983; as Unisitobion ?corylicola). On undersides of leaves of Isodon japonicus in Japan. Closely related to M. perillae on related hosts in China and Korea.

Macrosiphum jasmini (Clarke) Apterae are yellowish green, with pale siphunculi. In small colonies on undersides of leaves of Jasminum sp. in California, USA. Apparently not found since original description.

Macrosiphum jeanae Robinson Apterae are green with slightly darker appendages; BL c.1.8-2.0 mm. Feeding inside the pitchers of Sarracenia purpurea in Manitoba, Canada. Monoecious holocyclic with oviparae and alate males in September (original description). A very similar, undescribed species occurs on Darlingtonia californica in California (BMNH collection, leg. D.W. Nielsen).

Macrosiphum knautiae Holman (= M. silvaticum Meier) Apterae are yellowish green to grass-green, more rarely pinkish red, with dark head and thorax and black siphunculi; BL 2.3-3.8 mm. On undersides of leaves and shoot apices of Knautia spp. in central and eastern Europe, south to Greece (Papapanagiotou et al. 2012). Monoecious holocyclic, with oviparae and alate males in October in Czech Republic (original description). M. silvaticum was shown to be a synonym by morphological and molecular studies (Rakauskas 2003a, Turčinavičiene & Rakausakas 2009). Buga et al. (2012) studied the morphological distinction between M. rosae and M. knautiae on Knautia.

Macrosiphum lambi Robinson Apterae are green: BL 2.3-3.1 mm. Apterae have 4-45 secondary rhinaria on ANT III, and alatae have 62-106. On Athyrium filix-femina in eastern North America, and also collected on Thelypteris confluens (= palustris)in Alaska (BMNH collection, leg. A.G. Robinson).

Macrosiphum lapponicum Shaposhnikov Apterae are green, yellow or red; BL c.4 mm. Apterae have 37-46 secondary rhinaria on ANT III, alatae are unknown. On Athyrium alpestre (host identity requires confirmation) in north-west Russia (Khibiny Mountains).

Macrosiphum laseri Holman Apterae are yellow to yellowish green, sometimes covered with a white waxy secretion; BL 3.1-4.2 mm. On Laser trilobum and Laserpitium latifolium, mostly feeding on upper part of stem and in inflorescences, more rarely on undersides of leaves, which turn yellowish. In Crimea, Hungary (Szelegiewicz 1966a), France (BMNH collection, leg, G. Remaudière) and Italy (Barbagallo & Massimino Cocuzza 2014).

Macrosiphum lilii (Monell) Apterae are red and yellow with dark siphunculi; BL 2.5-3.2 mm. On Lilium spp. (and possibly other Liliaceae) in eastern USA. The aphids were described from bulbs imported from Japan (Comstock 1879), but there have been no subsequent Japanese records. See also Blackman & Eastop (2000).

Macrosiphum lisae Heie Apterae are whitish or yellowish white, with antennae pale except for dark segmental apices, and legs pale; BL 2.5-3.7 mm. On Epilobium angustifolium, colonising undersides of middle and lower leaves of the smaller plants that grow in shady places and rarely flower, reproducing vegetatively. In northern Europe. Monoecious holocyclic, with oviparae and alate males appearing in August-September (original description).

Macrosiphum longirostratum Jensen & Holman Apterae are green with dark tips to antennae, legs and siphunculi; BL 2.2-2.8 mm. Feeding singly or in small groups along mid-ribs on undersides of leaves of Woodsia mollis in Mexico. Monoecious holocyclic, with oviparae and alate males in October (Jensen & Holman 2000).

Macrosiphum manitobense Robinson Apterae are pinkish green (see aphidtrek.org) ; BL 2.5-2.8 mm. On Cornus sericea (= stolonifera)in Manitoba, Canada, and also reported from a Cornus sp. in western Oregon, USA. Robinson (1965) described all morphs, including oviparae and alate males in late September, but did not rule out host alternation, as no specimens were found on Cornus in July-August. The secondary host might be Potentilla, as this species is very similar to, and could be a synonym of, M. potentillae (Jensen & Rorabaugh 2020). [Specimens of a Macrosiphum from C. sericea in British Columbia (BMNH collection, leg. C.K. Chan) appear to be another, undescribed species.] 2n=10.

Macrosiphum meixneri Börner Apterae are yellowish green, dusted with white wax; BL 3.2-4.5 mm. On Euphorbia sp. (austriaca, hyberna) in Austrian and French Alps, and in Spain (Cantabrian mountains). Monoecious holocyclic with alate males (Nieto Nafría & Mier Durante 1991).

Macrosiphum melampyri Mordvilko Apterae are green with dark distal parts of antennae , femora, tibiae and siphunculi; BL 3.2-4.0 mm. On Melampyrum spp. in northern Europe (northern England and Scotland to north-west Russia). It is also recorded from Digitalis ambigua (not D. purpurea, contrary to Blackman & Eastop 2006). Heie (1994) gave a redescription. Monoecious holocyclic; oviparae and alate males occur on Melampyrum pratense in Sweden in late August (BMNH collection, leg. F. Ossiannilsson). A related, undescribed species was found on Melampyrum sp. in Pennsylvania, USA (BMNH collection, leg. H.L.G. Stroyan).

Macrosiphum mentzeliae Wilson Apterae are light to mid green with dorsal reddish patch, or pinkish (see aphidtrek.org); BL 1.8-3.0 mm. Alatae have 13-21 secondary rhinaria on ANT III. On leaves of Mentzelia spp., in western USA and Mexico, often in hot situations. Monoecious holocyclic, with oviparae and alate males occurring in October (Palmer 1952) and fundatrices in late March (Jensen & Rorabaugh 2020).

Macrosiphum mertensiae Gillette & Palmer Colour of apterae in life is unrecorded; BL 2-3 mm. On leaves of Mertensia, including M. platyphylla (aphidtrek.org)in western USA.

Macrosiphum miho Jensen & Holman Apterae are pale yellowish green; BL 1.1-2.2 mm. On “Aspidium sp.” growing in rock walls (Hottes & Frison 1931, as M. adianti), which is probably either Dryopteris marginalis or Polypodium virginianum (Jensen & Holman 2000). In USA (Illinois, Wisconsin). Apparently monoecious holocyclic, as fundatrices were collected in May.

Macrosiphum minatii (Das, Raychaudhuri & Raychaudhuri) Apterae are pale brown, BL 1.8-4.4 mm. On Delphinium sp., with collections also from other unidentified Ranunculaceae, in Himachal Pradesh, India. Described in Uroleucon, but transferred to Macrosiphum by Chakrabarti & Medda (2004).

Macrosiphum mordvilkoi Miyazaki Apterae are green or yellowish green with shiny black head and prothorax, mainly black antennae and black siphunculi; BL 2.5-3.8 mm. Apterae have 23-36 secondary rhinaria on ANT III, alatae have 30-40. On undersides of young leaves of Rosa spp., especially R. rugosa,in east Asia (not usually on cultivated roses). A record from Turkey (Kaygin et al. 2010) requires further conformation. Monoecious holocyclic with oviparae in October (original description).

Macrosiphum multipilosum Nevsky Apterae are undescribed. Alatae have pale green or pale yellow abdomen and brownish thorax, smoky cauda, darker siphunculi, black antennae and legs; BL c. 2.7 mm. Alatae have c.50 rhin. on ANT III, and numerous long dorsal body hairs. On stems and undersides of leaves of Helichrysum punctatum (? name is untraceable) in southern Kazakhstan. This species has subsequently been collected on H. maracandicum (Kadyrbekov 2014d, as Macrosiphoniella multipilosa; it is more likely to belong in Macrosiphoniella or Uroleucon than in Macrosiphum).

Macrosiphum nasonovi Mordvilko Apterae are greyish green or red, BL 2.0-3.6 mm. Alatae have a dark brown head and thorax and green abdomen and 7-11 secondary rhinaria on ANT III. Morphologically it is very similar to M. euphorbiae. Recorded from several genera of Ericaceae (Andromeda, Chamaedaphne, Oxycoccus, Vaccinium) in northern Europe (Finland, Latvia, Russia) and France (Pyrenees). Monoecious holocyclic on Vaccinium and Andromeda, with oviparae and alate males in September(Müller 1989).

Macrosiphum nevskyanum MacGillivray Only alatae are described (Nevsky 1929a, as Macrosiphum crataegi Monell); BL 1.9-2.1 mm. On Crataegus monogyna in Kazakhstan (Kadyrbekov 2002).

Macrosiphum occidentale (Essig) Apterae are very pale yellow to whitish; BL 2.6-3.5 mm. On undersides of leaves of Oemleria cerasiformis, and possibly Prunus emarginata (see Jensen 2000),in north-western USA and Canada (British Columbia). Monoecious holocyclic on Oemleria, with alate males (Jensen 2000). 2n=16

Macrosiphum olmsteadi Robinson Colour of living apterae unknown, probably pale green; BL 2.6-3.0 mm. Described from Eurybia macrophylla in Ontario, Canada. An aptera identified as this species on E. conspicua in Idaho is illustrated by A. Jensen at http://www.flickr.com/photos/sandnine/4749821046/, who has also collected samples on Eurybia in the Rocky Mountain parts of Montana, Alberta and Washington. Illinoia (Masonaphis) magna (Hille Ris Lambers), described from unidentified Compositae in Alberta and British Columbia, may be a synonym (A. Jensen, pers. comm.). Oviparae and alate males occur in October (original description).

Macrosiphum opportunisticum Jensen Apterae are medium to dark green, sometimes pink or reddish (for colour photo go to aphidtrek.org); BL 2.1-3.1 mm. Alatae have a dark brown head and thorax and dark green or pink abdomen with paired dark pleural abdominal markings, and 9-17 secondary rhinaria on ANT III.. This species feeds on a wide range of broad-leaved shrubs, particularly in montane habitats, in north-western USA and British Columbia. Hosts are mainly Ericaceae (Menziesia, Azaleastrum, Bryanthus, Pieris, Vaccinium), but it has also been found on Liquidambar, Symphoricarpos, Holodiscus, Sorbus and Spiraea, and under certain conditions it can even colonise Tsuga (A. Jensen, pers. comm.). The aphids live dispersed on the host plants and drop readily when disturbed, an unusually large proportion of individuals being alate. Monoecious holocyclic, with alate males; all morphs are described by Jensen (2012), who also discussed the opportunistic feeding on various shrubs by this aphid. 2n=16.

Macrosiphum oredonense Remaudière Apterae are pale green, with very long legs, pale except at apices; BL 2.4-4.5 mm. On undersides of leaves, and later in small compact colonies on stems and young fruits of Lonicera nigra in south, central and eastern Europe. Monoecious holocyclic, with oviparae and reddish-brown alate males produced in August (Müller 1989).

Macrosiphum oregonense Jensen Apterae are shiny green (see aphidtrek.org); BL 2.3-3.7 mm. On undersides of leaves of Lysichiton americanum, often in semiaquatic situations. In western North America (Oregon, Washington, British Columbia). Monoecious holocyclic with oviparae and alate males in October (Jensen 2000).

Macrosiphum orthocarpus Davidson Apterae are bright pea-green, with black antennae and siphunculi; BL more than 3 mm. Among flower-spikes of Orthocarpus purpurascens (= Castilleja exserta)in California, USA.

Macrosiphum osmaroniae Wilson Apterae are pale to mid green (see aphidtrek.org); BL 2.7-3.5 mm. On spring growth of Oemleria cerasiformis, migrating in late April-May to pass the summer in small populations on Pteridium aquilinum (Jensen & Holman 2000), with a return migration to Oemleria in September-October. In north-western USA (Oregon, Washington)and Canada (British Columbia). 2n=16*.

Macrosiphum pachysiphon Hille Ris Lambers Apterae are very pale pink with stout black siphunculi; BL 3.0-3.5 mm. Alatae have a dusky sclerotic pattern on the dorsal abdomen. On Rubus spp. (especially lasiocarpus) and sometimes on other Rosaceae (Rosa, Potentilla, Spiraea). Records from other plants (Berberis, Rumex)are probably of vagrants. In India and Pakistan. A record from Turkey (Şenol et al. 2014c) requires additional confirmation. The life cycle is unknown. 2n=18.

Macrosiphum paektusani Lee & Havelka Apterae are green, BL 3.4-4.3 mm. On Rosa spp. in North Korea (S. Lee et al. 2009). Similar to M. mordvilkoi but with paler head and pale bases to siphunculi. The life cycle in unknown.

Macrosiphum pallens Hottes & Frison Apterae are pale whitish green, with pale siphunculi and cauda; BL 1.9-2.4 mm. Alatae are pale yellow-green, without marginal sclerites. On Silphium integrifolium in Illinois, USA, with records also from Ambrosia trifida and Aster (=Symphyotrichum) novae-angliae (the latter from New Brunswick, Canada; aphidtrek.org).

Macrosiphum pallidum (Oestlund) Plate 25h Apterae are green or pink, with antennae, femoral apices, tibiae, tarsi dark, and siphunculi dark except at bases; BL 2.1-4.6 mm. Apterae have 3-14 secondary rhinaria on ANT III, alatae have 14-22. On wild Rosa spp. and various other Rosaceae (Agrimonia, Fragaria, Geum, Potentilla), and also apparently able to colonise a wide range of other plants, although some records may be misidentifications of other species (e.g. records from Oenothera are probably all M. gaurae). Widespread in North America. The life cycle still seems to be unknown. MacGillivray (1968) provided a detailed redescription. A very similar but paler species on Oxalis oregona in Oregon (BMNH collection, leg. D. Hille Ris Lambers) is undescribed. A record from Turkey (Akyürek et al. 2010, repeated in Görür et al. 2012) is based on a misidentification. 2n=10.

Macrosiphum parvifolii Richards (Fig.56a,b) Apterae are whitish or pale pink, powdered with white wax, with dark tips to appendages; BL 2.8-3.4 mm. Alatae are very pale whitish except for sclerotised parts of head and thorax, with 6-12 secondary rhinaria on ANT III (Jensen 2012). On Vaccinium spp. in British Columbia, Canada (Richards 1967 and BMNH collection), and in north-western USA (Oregon, Idaho, Montana), where it is also recorded from Menziesia ferruginea (Jensen 2012). 2n=16.

Macrosiphum pechumani MacGillivray Apterae are milky white, with black head and appendages; BL 2.5-3.1 mm. On Asparagaceae (in genera formerly placed in Convallariaceae; Convallaria majalis, Maianthemum racemosa) in north-eastern USA (New York, Washington DC) and Canada (Ontario; BMNH, leg. M. Sÿpkens). Monoecious holocyclic, with oviparae and alate males in September–November (original description). Records from plants in other families (Fragaria, Hibiscus, Pyrola) are likely to be vagrant individuals, although Leonard (1968) recorded oviparae from Hibiscus.

Macrosiphum penfroense Stroyan Apterae are bright apple green, immatures have a wax bloom; BL 1.9-3.4 mm. On Silene uniflora (= maritima)in England and Wales, feeding in small groups in flowers, or singly on buds or on upper surfaces of leaves. Apparently anholocyclic, with delayed production of ovipariform apterae under laboratory conditions (G.W. Watson, unpubl. observations). Described as a subspecies of M. sileneum (= M. stellariae), but the consistent differences in morphology and host plant indicate that it should have separate species status (Watson 1982). 2n=10.

Macrosiphum (Unisitobion) perillae (Zhang) Apterae are pale yellow with broad dark lateral longitudinal stripes, black antennae , legs and siphunculi, and a pale cauda; BL 1.8-2.9 mm. On upper parts of stems and undersides of leaves of certain Lamiaceae (Isodon, Mentha, Perilla) in China and Korea. Monoecious holocyclic on Isodon inflexus, with oviparae and alate males in October (S. Lee et al. 2001b). (A record from Rosa multiflora could be a misidentification of host plant.) 2n=18 (Chen & Zhang 1985a).

Macrosiphum polanense Pašek Apterae are yellowish white to pale green; BL 3.8-4.6 mm. In large colonies on flower stems of Cicerbita alpina in Slovakia (Pol’ana). Monoecious holocyclic with oviparae and alate males in September (original description). M. alpinum is possibly a synonym.

Macrosiphum potentillae (Oestlund) Colour of apterae in life is unrecorded; BL c. 2.2 mm. On undersides of leaves of Potentilla anserina and Potentilla sp. in USA (Minnesota, Kansas, Oregon).

Macrosiphum prenanthidis Börner Apterae are green, pinkish or violet brownish, slightly wax-powdered; BL 3.2-3.8 mm. Apterae have 9-16 secondary rhinaria on ANT III, alatae have 20-30. On Prenanthes purpurea, in small colonies on stems in spring/summer, and later on undersides of leaves.In central and eastern Europe, and recently recorded from northern Italy (Barbagallo et al. 2011). Monoecious holocyclic with alate males.

Macrosiphum (Neocorylobium) pseudocoryli PatchApterae are green with antennae, tibiae and siphunculi dusky to dark, especially at apices (see influentialpoints.com/Gallery); BL 2.4-3.6 mm (MacGillivray 1968). On young growth and undersides of leaves of Ostrya virginiana and Corylus spp. in north-eastern USA and Ontario, Canada. Presumably monoecious holocyclic; oviparae were collected in Virginia in late September (BMNH collection, leg. R.L. Blackman). 2n=10*.

Macrosiphum pseudogeranii Chakrabarti & Raychaudhuri Colour of apterae in life is not recorded; BL 2.8-3.5 mm. On Geranium spp. in north-west India (Himachal Pradesh, Uttar Pradesh). Probably monoecious holocyclic; Raychaudhuri et al. (1980) described the ovipara (from Polygonum – a vagrant?), and Maity & Chakrabarti (1984) described the alate vivipara and male.

Macrosiphum ptericolens Patch Apterae are pale yellowish green to darker shiny green, with mainly pale appendages; BL 2.3-3.3 mm. Alatae are pale green with appendages also mainly pale (see influentialpoints.com/Gallery). Apterae have 3-35 secondary rhinaria on ANT III, alatae have 27-66. On Pteridium aquilinum in North America east of the Rocky Mountains, and introduced to Europe (Lawton & Eastop 1975) and South America (Brazil). Records from other ferns are probably attributable to other Macrosiphum species (see Jensen & Holman 2000). Monoecious holocyclic with alate males on Pteridium, but in the USNM collection there are alatae, oviparae and a male that are apparently M. ptericolens collected on Ilex verticillata in Pennsylvania, suggesting that host-alternation might also occur. Probing behaviour and stylet penetration were studied by Halarewicz & Gabryś (2012). 2n=16.

Macrosiphum pteridis Wilson (Fig.43h) Apterae are very pale green; BL 2.5-4.2 mm. Heteroecious holocyclic, migrating from Holodiscus discolor to Pteridium aquilinum in western USA and Canada (Jensen & Holman 2000). Detailed descriptions of all morphs occurring on Holodiscus are given by Forbes & Chan (1993, as Sitobion blackmani). Wilson’s type material included M. clydesmithi, and there is also confusion in the literature with M. rhamni (e.g. Robinson 1980). 2n=16.

Macrosiphum pulcherimum (Nevsky) Apterae are bright green, with a slightly mealy coat, siphunculi and cauda pale green; BL 1.8-2.2 mm. On stems and leaves of Lactuca saligna and Lactuca sp. in Central Asia.

Macrosiphum pyrifoliae MacDougall Apterae, according to the original description, are dusky brown, with head slightly darker; BL 2.4-3.6 mm. Described from Pyrus occidentalis (= Sorbus sitchensis) in British Columbia, Canada (MacDougall 1926). MacDougall counted 14-17 secondary rhinaria on ANT III of apterae, but apterae identified as this species from the type locality (Merritt) in MacDougall’s collection have only 3-8 secondary rhinaria. Aphids believed to be this species were collected more recently in the Vancouver area, from Sorbus aucuparia and S. sitchensis (BMNH collection, leg. C.K. Chan). These had green and red colour morphs (C.K. Chan, pers. comm.). Apparently monoecious holocyclic on Sorbus, although sexuales have not been described. Very similar to M. euphorbiae. 2n=10, 11 and 12*.

Macrosiphum ranunculi Pašek Apterae are yellowish white to pale greenish yellow; BL is unrecorded. Described from Ranunculus auricomus in Slovakia, and since recorded from other Ranunculus spp. (Holman 2009). Monoecious holocyclic (original description).

Macrosiphum raysmithi Hille Ris Lambers Apterae are very pale green to whitish green, with pale appendages except for tarsi and apices of antennal segments; BL 3.4-3.7 mm. In large, very compact colonies on undersides of leaves of Lonicera ledebourii in California, USA.

Macrosiphum rebecae Jensen & Holman Apterae are green to bright green with front of head darker; BL 1.8-2.3 mm. On pedicels and along veins on undersides of fronds of Adiantum sp(p.) in Mexico. Probably heteroecious holocyclic, migrating to an unknown primary host (Jensen & Holman 2000).

Macrosiphum rhamni (Clarke) Apterae on undersides of leaves of Rhamnus in spring are yellow-green, with a large round patch of reddish internal pigment in the centre of the dorsum (Essig 1917), and have antennae ringed with black and pale siphunculi and cauda; BL 1.8-2.4 mm. Fundatrices occur in March on R. purshiana in British Columbia (BMNH collection, leg. C.K. Chan). Heteroecious holocyclic; migration occurs to ferns (Pteridium aquilinum, Polypodium hesperium; Jensen et al. 1993). Apterae on ferns are pale yellow; BL 1.8-2.6 mm. Colonies on ferns are frequently ant-attended. Gynoparae and males mature during late September and early October, and after returning to Rhamnus, can be found until mid-November. Oviparae can be found abundantly from mid-October to late November (see aphidtrek.org). In western North America. Closely related to M. clydesmithi but ranging further west (California) and north (British Columbia). A record from the Netherlands (Piron 2017, as Sitobion rhamni) requires further confirmation. 2n=16.

Macrosiphum rosae (L.) Plate 25i Apterae are green or deep pink to red-brown or magenta, with shiny black head and prothorax, bicoloured yellow and black antennae and legs, black siphunculi and pale yellow cauda (see influentialpoints.com/Gallery); BL 1.7-4.2 mm. On young growth of wild and cultivated Rosa spp. in spring, migrating to Dipsacaceae (Dipsacus, Knautia, Succisa) and Valerianaceae (Centranthus, Valeriana), and also sometimes in summer on other plants, especially other Rosaceae (including Pyrus); also some Onagraceae, and Aquifoliaceae (Ilex). Host alternation is facultative; colonies can remain on Rosa through the summer, producing some sexuales in autumn, and anholocyclic overwintering occurs in warmer regions. World-wide, except for east and south-east Asia; records from Japan apply to M. mordvilkoi. Tomiuk & Wöhrmann (1984) studied genotypic (allozyme) variation in European populations. A host-specific parasitoid, Aphidius rosae Haliday, has ben introduced into Australia as a biological control agent (Kitt 1996, Kitt & Keller 1998). See also Blackman & Eastop (2000). 2n=10.

Macrosiphum rubiarctici Heikinheimo Apterae have dark reddish violet head and thorax, and brownish violet abdomen, with greenish brown spots at bases of siphunculi, and antennae and siphunculi black except at bases; BL 2.5-3.4 mm. On flowerstalks of Rubus spp. (arcticus, saxatilis, stellatus) in Scandinavia, Slovakia and Russia (near Moscow; Holman 2009). The life cycle is unknown.

Macrosiphum rudbeckiarum (Cockerell) ( = Macrosiphum cockerelli Hottes) Apterae are pale green, with mainly pale appendages, somewhat darker in autumn (see aphidtrek.org); BL 2.9-4.2 mm. On undersides of leaves of Rudbeckia spp. in western USA. Monoecious holocyclic with oviparae and alate males in September. Hottes (1949a) gave a very full description.

Macrosiphum salviae Bartholomew Apterae are dark chestnut brown with black antennae, legs and siphunculi, and usually a central black dorsal abdominal patch; BL 2.0-2.5 mm. On Lamiaceae (Inga, Salvia, Leonurus, Ocimum, Tetrademia) in California USA, Cuba, Honduras (Evans & Halbert 2007), Venezuela, El Salvador, Costa Rica and Puerto Rico (Villalobos Muller et al. 2010). [This species should perhaps be in the genus Sitobion (and the record from Ocimum could be misidentified Sitobion mesosphaeri, which was for some time regarded as a synonym].

Macrosiphum schimmelum Jensen Apterae are pale to mid-green, sometimes pink or yellowish, usually wax-dusted (see aphidtrek.org); BL 2.2-3.7 mm. On undersides of leaves, and later on flowers and fruits, of Lonicera utahensis in north-western USA (Idaho, Oregon, Washington) and British Columbia. Possibly also on Symphoricarpus, although the host may have been misidentified. Monoecious holocyclic, with fundatrices present in May-July and oviparae and alate males in September (Jensen 2015).

Macrosiphum (Neocorylobium) skurichinae Pashchenko Apterae are brown, with antennae and legs mainly pale, living on Corylus sieboldiana var. mandshurica in envelopes of nuts, attended by ants. In east Siberia (Pashchenko 1985).

Macrosiphum (Unisitobion) sorbi Matsumura Apterae are pale yellow to yellow-green, with brown head and variably developed brown spinal and marginal longitudinal stripes, joined in region of siphunculi, which are jet black; BL 2.3-3.8 mm. On undersides of leaves of Sorbaria sorbifolia in Japan and Korea (S. Lee et al. 2001b). Monoecious holocyclic, with oviparae and alate males in late October in Japan (Miyazaki 1971). The host was originally given as Sorbus japonica,but this may have been in error.

Macrosiphum stanleyi Wilson Apterae are spindle-shaped, green (see aphidtrek.org); BL 3.0-4.4 mm. Alatae have brown-black head and thorax. On leaves and stems of Sambucus spp. (callicarpa, cerulea, pubens) in western North America, from Colorado to Alaska. Monoecious holocyclic; oviparae and alate males occur in August in Colorado (Palmer 1952), but in British Columbia oviparae are present as early as June (BMNH collection, leg. C.K. Chan). 2n=16*.

Macrosiphum stellariae Theobald (= Macrosiphum sileneum Theobald) (Fig.51a) Apterae are yellowish green, green or red, all segments of appendages, including femora and siphunculi, having blackish apices (see influentialpoints.com/Gallery); BL 1.8-4.4 mm. In small, loose colonies on young shoots of various Caryophyllaceae (Dianthus, Gypsophila, Moehringia, Silene, Stellaria), and sometimes on certain other plants (Papaver, Ranunculus, Valeriana), although pale specimens may easily be confused with M. euphorbiae.In north-western, northern and central Europe, Georgia (Özdemir & Barjadze 2015), and introduced to Canada (British Columbia) and New Zealand. 2n=10.

Macrosiphum symphyti Barjadze & Chakvetadze Apterae are yellow or yellowish green with paler spinal stripe, dark apices to antennal segments, legs and siphunculi, and a pale cauda; BL 2.6-4.3 mm. Alatae have 18-34 secondary rhinaria on ANT III. In sparse colonies on undersides of leaves and stems of Symphytum asperum in Georgia, and also now identified from Anchusa sp. in Iran (Özdemir & Barjadze 2015) and Turkey (Şenol et al. 2017). Monoecious holocyclic with alate males and oviparae collected in early November (Barjadze & Chakvetadze 2008).

Macrosiphum tenuicauda Bartholomew Apterae are green, BL 1.7-2.6 mm. On Urtica spp. in western North America (California, Oregon, Colorado, British Columbia). C.-k. Chan (pers. comm.) was able to rear this species for two years on young celery plants in the laboratory. 2n=10.

Macrosiphum tiliae (Monell) Apterae are green or pink, with dark antennae, tibiae and siphunculi; BL 2.4-2.7 mm Colonies occur on new growth, especially suckers, of Tilia americana (Hottes & Frison 1931). Widely distributed in northern USA. Monoecious holocyclic; oviparae and alate males occur in September in Wisconsin (Davis 1914).

Macrosiphum timpanogos Knowlton Apterae are pale, BL 4.1-5.0 mm. Probably collected on “a lupine of some kind” in western USA (Utah).

Macrosiphum tinctum (Walker) (= Macrosiphum epilobiellum) Apterae are mid to blue green with a darker spinal stripe (a red form is recorded from Switzerland), femora entirely pale and siphunculi dusky at apices (see influentialpoints.com/Gallery); BL 2.0-4.0 mm. Alatae have11-26 secondary rhinaria on ANT III. On Epilobium spp., especially E. angustifolium and E. montanum, feeding in small numbers on stems, flower-buds and seed-cases. Widely distributed in Europe. Monoecious holocyclic, with oviparae and alate males in late August-September. 2n=10.

Macrosiphum tolmiea (Essig) Apterae are green with black-tipped siphunculi (see aphidtrek.org); BL c. 3 mm. On three spp. of Saxifragaceae, Tolmiea menziesii, Tellima grandiflora and Mitella caulescens, near streams or seeps in shady locations, in north-western USA and British Columbia, Canada. Probably anholocyclic in some areas (Jensen 2000), but Jensen (2012) found fundatrices on early spring growth of Tolmeia menziesii in a montane habitat in Washingon.

Macrosiphum tonantzin Peña-Martínez, Muñoz-Viveros & Jensen Apterae are pale to bright green, the abdomen often with a transparent whitish spot in the centre; BL 2.0-2.6 mm. On growing points and under young leaves of Pittosporum undulatum in Mexico (Jensen et al. 2019). This plant, the only known host, is native to Australia, so M. tonantzin probably has one or more Mexican hosts, the long hairy last rostral segment indicating that these would be likely to have densely hairy leaves. Alate males are produced in December, but no oviparae, so it is possibly heteroecious with a sexual phase on another plant.

Macrosiphum trollii Börner Apterae are pale yellowish or pale green; BL 3.0-4.3 mm. On undersides of leaves and on flowers of Trollius europaeus. InEurope, with a boreoalpine distribution. Monoecious holocyclic, but males are undescribed.

Macrosiphum tuberculaceps (Essig) Apterae are yellowish with mainly pale appendages and dark-tipped siphunculi (see aphidtrek.org); BL 3.5-4.3 mm. On Achlys triphylla in north-western USA and British Columbia. Monoecious holocyclic with alate males and oviparae in October-November (Jensen 2000).

Macrosiphum valerianae (Clarke) (= Macrosiphum kiowanepum (Hottes)) Apterae are dark reddish, or greenish with a reddish tinge, pruinose, with very long, usually dark brown to blackish, antennae, tibiae and siphunculi (see aphidtrek.org); BL 2.1-4.0 mm. A polyphagous species, colonising leaves and stems of a wide range of host plants, particularly in mountains and foothills throughout western USA and Canada. Jensen (2012) established its synonymy with M. kiowanepum (Hottes), and recorded its occurrence on hosts in nine families of flowering plants; Apiaceae (Lomatium), Asparagaceae (Camassia), Asteraceae (Hymenoxys, Rudbeckia, Taraxacum), Brassicaceae (Arabis), Melanthiaceae (Zigadenus), Onagraceae (Epilobium), Rosaceae (Geum, Potentilla, Rosa), Scrophulariaceae (Pedicularis, Penstemon) and Valerianaceae (Valeriana). Monoecious holocyclic, and perhaps utilising various overwintering hosts. Pale specimens do occur and on slides these are almost indistinguishable from M. euphorbiae (see Jensen 2012). Antolin & Addicott (1988) studied habitat selection and competition with Aphis varians on Epilobium angustifolium in Colorado. M. potentillicaulis Miller, described from Potentilla bakeri in Idaho, is a synonym (Jensen & Rorabaugh 2020).

Macrosiphum vancouveriae Jensen Apterae are very pale green to white with pale brown eyes (see aphidtrek.org); BL 2.2-3.8 mm. On Vancouveria hexandra in north-western USA. The life cycle is uncertain, but probably monoecious holocyclic on Vancouveria, alate males and oviparae occurring in October-November (Jensen 2000).

Macrosiphum (Neocorylobium) vandenboschi (Hille Ris Lambers) Apterae are pale green, with black-tipped antennae, legs and siphunculi; BL 2.1-2.2 mm. On Corylus cornuta var. californica in California (Hille Ris Lambers 1966; as Corylobium vandenboschi). Biology and sexual morphs are unknown.

Macrosiphum venaefuscae Davis Apterae are pale green; BL 2.5-2.8 mm. On Polygonaceae (Fallopia, Polygonum, Rumex) in eastern North America. Monoecious holocyclic with alate males; sexual morphs and eggs are found on Rumex (original description).

Macrosiphum verbenae (Thomas) Apterae are bright pea-green with 2-3 darker green longitudinal stripes on abdomen, antennae pale with dusky tips, eyes black, siphunculi pale, cauda rather short and whitish (original description); BL is unrecorded (“medium size”). On leaves of a Verbena sp. in Illinois, USA. Since recorded from Verbena spp. in numerous states (Smith & Parron 1978), but there are no type specimens and no further account of this species, and many of these records could be M. euphorbiae.

Macrosiphum vereshtshagini Mordvilko Apterae are green, wax-dusted, with apices of antenane and siphunculi darker; BL 3.9-4.8 mm. On Cimicifuga foetida (= Actaea cimicifuga)in Russia (Altai region).

Macrosiphum violae Jensen Apterae are very pale green, with pale brown eyes (see aphidtrek.org); BL 2.2-3.3 mm. On Viola glabella in north-western USA. Attempts to transfer to two other Viola spp.were unsuccessful (Jensen 2000). Monoecious holocyclic with oviparae and alate males in September-October.

Macrosiphum walkeri Robinson Apterae are variably coloured; yellow, green, pink or red (see aphidtrek.org); BL 1.7-3.3 mm. On numerous genera and species of ferns (Adiantum, Asplenium, Athyrium, Cystopteris, Davallia, Dicksonia, Dryopteris, Matteuccia, Nephrolepis, Osmunda, Pellaea, Polypodium, Polystichum, Woodwardia) in western North America. Anholocyclic in western Oregon, overwintering on Polypodium and Polystichum, although oviparae (without males) were found on Athyrium. However Jensen (2021) has now shown that over much of its range M. walkeri has a sexual phase on Holodiscus, and has provided decriptions of alate males and oviparae. 2n=16.

Macrosiphum weberi Börner Apterae are usually dark red or dark violet, with black siphunculi, but a green colour variant occurs in Scotland (see influentialpoints.com/Gallery); BL 1.8-3.0 mm. In small, frequently ant-attended colonies on stems of Succisa pratensis, and sometimes also on Scabiosa spp. Throughout much of Europe, frequently in alpine habitats. Records from Knautia are probably all referable to M. knautiae. Monoecious holocyclic with alate males.

Macrosiphum willamettense Jensen Apterae are shining pale greenish white, immatures are slightly waxy (see aphidtrek.org); BL 2.3-3.5 mm. On young growth of Spiraea douglasii in north-western USA and Canada (British Columbia, Oregon, Washington). Monoecious holocyclic with oviparae in October (Jensen 2000). 2n=10.

Macrosiphum wilsoni Jensen Apterae are pale yellowish white (see aphidtrek.org); BL 2.2-3.9 mm. On Disporum (= Prosartes)spp. in north-western USA (Oregon, Idaho, Washington) and Canada (British Columbia). Monoecious holocyclic, with oviparae and alate males in October (Jensen 2000).

Macrosiphum woodsiae Robinson Apterae are dark green, BL 1.3-2.5 mm. Described from Woodsia ilvensis in northern Canada, and subsequently found on other Woodsia spp., Cryptogramma crispa and Pellaea sp. in Oregon, USA, perhaps specialising on ferns at high altitudes/latitudes (Jensen & Holman 2000). Probably monoecious holocyclic on Woodsia. 2n=16.

Macrosiphum zionense Knowlton Apterae are green, yellow or orange with black antennae, tibiae and siphunculi (see aphidtrek.org); BL c.4.1 mm. On Lupinus sp. and Thermopsis spp. in western USA (Utah, Idaho, Oregon, Washington; BMNH collection, leg. S. Paulaitis and G.F. Knowlton).

Macrotrichaphis Miyazaki

Aphidinae: Macrosiphini

Two east Asian species related to Uroleucon but with siphunculi bearing hairs. First tarsal segments all have 5 hairs. Their biology is hardly known.

Macrotrichaphis rarissima Pashchenko Apterae are dark brown; BL c. 2.3 mm. On apical leaves of Saussurea sp.in east Siberia.

Macrotrichaphis yatsugatakensis Miyazaki (Fig.13a) Colour of apterae life is unknown; BL 1.9-2.3 mm. Described from an unknown host in Japan, but there are specimens in the BMNH collection from Artemisia princeps var. orientalis in Korea (leg. W.H. Paik).

Maculolachnus Gaumont

Lachninae: Lachnini

Three speciesrelated to Lachnus but assocated with Rosaceae, without any distinct pattern of pigmentation of forewings, and with dorsal hairs often placed on dark sclerites. The host plant of a fourth species (Maculolachnus paiki Seo) is unknown.

Maculolachnus rubi A.K. Ghosh & Raychaudhuri Apterae are dark brown; BL 2.7-3.0 mm. On apices of stems of Rubus sp., attended by ants, in Meghalaya, India. Alatae and sexual morphs are unknown.

Maculolachnus sijpkensi Hille Ris Lambers Apterae are yellowish brown to dark brown or blackish; BL 2.4-3.8 mm. On wild Rosa spp., in sparse colonies on stems near ground,ant-attended. Monoecious holocyclic with apterous males. In North America and Mongolia, and now also reported from Iran (Mehrparvar & Lashkari 2021). 2n=10.

Maculolachnus submacula (Walker) Plate 3e Apterae are yellowish brown to dark chestnut brown (see influentialpoints.com/Gallery); BL 2.7-3.8 mm. On stems near ground or (in summer) on surface roots of wild and cultivated Rosa spp., invariably ant-attended. In summer it has also been found on roots of Geranium spp. and Potentilla anserina (Kadyrbekov 2012a). Monoecious holocyclic with oviparae and apterous males in September-October. The male genitalia were described and illustrated by Wieczorek et al. (2012). Throughout Europe, and in south-west and Central Asia, Pakistan and India. Gottschalk (1989) gave an account of this aphid in Germany. 2n=10.

Mastopoda Oestlund

Aphidinae: Macrosiphini

One North American species with greatly reduced tarsi and 5-segmented antennae, possibly with host alternation between Viburnum and ferns like Shinji in east Asia, although this needs to be confirmed.

Mastopoda pteridis Oestlund Apterae are yellowish white; BL c. 1.7 mm. On Pteridium aquilinum in north-eastern USA and Ontario, Canada (Patch 1910a, Robinson 1966). Possibly heteroecious holocyclic, as also collected from “ant sheds on leaf petioles” of Viburnum sp. (Robinson 1966, citing J.O. Pepper). Males are recorded from Pt. aquilinum in September.

Matsumuraja Schumacher

Aphidinae: Macrosiphini

About 14 east Asian species, plus one undescribed from Australia (see below), with long capitate hairs arising from tubercles, which are often extended into fingerlike processes. They are associated with Rubus (except one species described from Ficus, which is probably not the true host). However, unusually, Rubus may be the ancestral secondary host, as the only species in which the life cycle has been clarified has a sexual phase on Clethra (the fundatrix having a very distinctive morphology, see Takahashi 1959). Takahashi & Sorin (1965) and Miyazaki (1971) keyed the Japanese species, and Lee (2002b) redescribed and keyed the two species in Korea.

Matsumuraja calorai Calilung Apterae are brownish yellow; BL 1.1-1.3 mm. On Ficus ribes var. cuneata in the Philippines (Calilung 1972). Other morphs, biology and life cycle are unknown

Matsumuraja capitophoroides Hille Ris Lambers (Fig.47c,e) Apterae are pale yellow, with antennae dark at segmental apices; BL 1.3-2.3 mm. On Rubus spp. in Pakistan, Nepal, India and Sri Lanka. According to Chakrabarti & Banerjee (1993a) there is host alternation between Rubus (as primary host) and Poa annua (as secondary host) in India, but this seems very unlikely and needs confirmation. As noted by Hille Ris Lambers (original description), this species differs from the description of M. formosana only in lengths of tubercular bases of spinal hairs, which could vary according to environment or degree of alatiformity. 2n=14.

Matsumuraja formosana Takahashi Apterae are white, with antennae dusky at segmental apices; BL c.1.8 mm. Alatae are unknown. On Rubus sp. and Fragaria × ananassa in Taiwan (Tao 1963). M. capitophoroides is a possible synonym (see above). An aptera collected on Duchesnea indica in Fujian province, China (Qiao & Zhang 1999c), and described under the name Chaetosiphon anguifragum Zhang & Qiao, is possibly this species.

Matsumuraja hirakurensis Sorin (Fig.47k) Apterae are orange, with antennae dark except for basal part of III; BL c.1.2-1.3 mm. On Rubus idaeus and Rubus sp. in Japan.

Matsumuraja nuditerga Hille Ris Lambers Colour of apterae in life is unknown; BL 1.2-1.4 mm. On Rubus sp. in Japan. There is also a record from ?Pilea sp. in West Bengal (A.K. Ghosh & Raychudhuri 1972), but the specimens had a shorter R IV+V and may not be this species. 2n=14.

Matsumuraja rubea Sorin (Fig.47i) Apterae are entirely white; BL c.1.2 mm. On Rubus sp. in Japan (Takahashi & Sorin 1965). 2n=14.

Matsumuraja rubi (Matsumura) (Fig.47b) Apterae are pale yellowish or greenish white; BL 1.0-1.6 mm. On Rubus spp. in Japan, east Siberia and Korea. Heteroecious holocyclic, with gynoparae and alate males produced in late October, returning to unknown primary host (Takahashi 1959c). 2n=14.

Matsumuraja rubicola Takahashi (Fig.47a) Apterae are yellow, with antennae dusky at segmental apices, BL c. 1.8-2.0 mm. Alatae have a large black dorsal abdominal patch and secondary rhinaria distributed III 32-36, IV 14-17, V 4-7. On Rubus sp. and Fragaria × ananassa in Taiwan (Tao 1963).

Matsumuraja rubifoliae Takahashi Plate 20a (Fig.47h,l) Apterae (on Rubus)are yellow or white, antennae with segmental apices often blackish; BL 1.3-1.7 mm. Alatae have a black dorsal abdominal patch, black clavate siphunculi and many secondary rhinaria on ANT II-V. On shoot tips and undersides of young leaves of Rubus spp. in east Siberia, China, Thailand, Taiwan, Korea, Japan and Ryukyu. Aphids collected on R. rosifolius in Queensland, Australia (BMNH collection, leg. V. F. Eastop, 15.vi.89) may also be this species, although differing by characters given in the key to apterae on Rubus. Heteroecious holocyclic in Japan (Takahashi 1959c), with sexual phase on Clethra barbinervis; the fundatrix is a very different, much larger, pale yellow aphid (BL 3-4 mm). Commonly also anholocyclic on evergreen Rubus in Japan, and probably elsewhere. Small forms occur on certain Rubus spp. in summer with small dorsal hairs and thicker, more swollen siphunculi (Takahashi & Sorin 1965). 2n=14.

Matsumuraja rubiphila Takahashi (Fig.47d) Apterae are yellow, with apices of antennal segments and of siphunculi blackish; BL c. 1.5 mm. On Rubus palmatus in Japan (Takahashi & Sorin 1965). 2n=14.

Matsumuraja sorini Takahashi (Fig.47g,m) Apterae are whitish yellow with mainly pale appendages; BL c.1.7-1.8 mm. On Rubus peltatus in Japan (Takahashi & Sorin 1965).

Matsumuraja taisetsusana Miyazaki (Fig.47j) Apterae are pale yellow, with apices of antennal segments and of siphunculi dusky; BL 1.6-1.9 mm. On Rubus idaeus var. aculeatissimus in Japan, and on R. sachalinensis and Rubus sp. inSakhalin and east Siberia (Pashchenko 1988a).

Matsumuraja sp. (Fig. 47f)Apterae are yellow-green, BL 1.1-1.6 mm. On Rubus rosaefolius in Queensland, Australia (BMNH collection, leg. V. F. Eastop, 15.vi.89). A single alata trapped at Galton Research Laboratory, Queensland, 29.iv.87 (BMNH collection, leg. J.Thomas) is probably also this species. 2n=18.

Megoura Buckton

Aphidinae: Macrosiphini

Eight species in Europe and Asia, and one recently described from New Zealand, with swollen cigar-shaped siphunculi, associated with Leguminosae/Fabaceae. Regional accounts are available from north-west Europe (Heie 1995), Japan (Miyazaki 1971), and Korea (S. Lee et al. 2002b). W. Lee et al. (2014) provided a key to the world species. Kim & Lee (2008b) studied the molecular systematics of the genus, comparing results of analysis of four mitochondrial and three nuclear DNA sequences.

Megoura brevipilosa Miyazaki Apterae are of unknown body colour; BL c. 2.6-3.0 mm. Alatae are reddish vermilion with a little white wax powder dorsally. On Lespedeza spp. in Japan and east Siberia; on L. bicolor var. japonica it was found on undersides of old leaves. Monoecious holocyclic, with oviparae and alate males in November-December (Sorin & Shinohara 1973). Molecular study indicates a close relationship to M. lespedezae (Kim & Lee 2008).

Megoura crassicauda Mordvilko Apterae are green with black head, prothorax, antennae, siphunculi and cauda; BL 3.0-3.7 mm. Tibiae are variably pigented. On stems and growing points of Vicia and Lathyrus spp. in east Asia (east Siberia, China, Taiwan, Japan, Korea), and recently introduced to Australia (New South Wales; Hales et al. 2017, Duric et al. 2022). In Korea it is also recorded from Pisum sativum and Amphicarpaea edgeworthii.[The name M. japonica (Matsumura) has been used for this species by Chinese and Japanese authors.] 2n=10.

Megoura dooarsis (A.K. Ghosh & Raychaudhuri) Apterae are green to dark green with purplish brown head, and dorsal abdomen with three rows of roundish or transversely oval whitish wax spots; BL 2.2-2.9 mm. On both sides of leaves and on twigs of Indigofera spp., especially I. heterantha,and also recorded from a Hedysarum sp. (plant species name on label is campanulatum),and Tephrosia purpurea. In India, Pakistan, Afghanistan, Kashmir and Thailand. 2n=20 (Dutta & Gautam 1993).

Megoura lathyricola Lee & Akimoto Apterae are pale green with yellowish-brown head, dark antennae and siphunculi, mainly pale legs and a pale cauda; BL 4.0-4.5 mm. On stems and young leaves of Lathyrus japonicus (ssp. japonicus) in coastal localities in Japan (W. Lee et al. 2014). Life cycle is unknown; a clone from Nagasaki reared in short-day conditions failed to produce sexual morphs so was apparently anholocyclic.

Megoura lespedezae (Essig & Kuwana) Plate 25a Apterae are greenish to yellowish with dark brown antennae, legs dark brown except for distal halves of tibiae, black siphunculi and yellow cauda (for colour photographs see Sasaki 2019a); BL 1.8-2.5 mm. On Lespedeza spp. in China, Taiwan, Thailand, Korea, east Siberia and Japan. Also recorded from Cajanus cajan, Desmodium trifolium (= triflorum?)and Indigofera teysmanni (= zollingeriana)in India (Raychaudhuri 1984, partly as Neomegouropsis cajanae), and from L. thunbergii in Switzerland (Giacalone & Lampel 1996). L.K. Ghosh (1972b) reported the occurrence of an ovipara in West Bengal (as M. cajanae). 2n=14.

Megoura litoralis Müller Apterae are pale green with brownish head and prothorax and appendages, and darker brown siphunculi; BL 3.0-4.4 mm. On young growth, flowers and fruits of Lathyrus japonicus ssp. maritimus in north-west and north Europe (not UK). Monoecious holocyclic with alate males (Heie 1995). Rakauskas et al. (2008) reviewed its distribution and habitat.

Megoura nigra Lee Apterae are dark reddish brown; BL 3.1-4.3 mm. On young stems and undersides of leaves of Vicia venosa in South Korea. Monoecious holocyclic with oviparae and alate males in mid-October (Lee et al. 2002b). A record from Turkey (Akyürek et al. 2019) requires further confirmation.

Megoura stufkensi Eastop Apterae are green with appendages mainly pale, the antennae and siphunculi darker towards apices; BL 1.6-2.0 mm. Alatae are unusual in having only 0-1 secondary rhinaria on ANT III, and dark-veined forewings with the media only once-branched. On Carmichaelia spp. in New Zealand, feeding on plant stems deep within bushes (Eastop 2011). The life cycle and sexual morphs are unknown. This is the only apparently indigenous species of Macrosiphini in New Zealand, and has unique features, but DNA data indicate that it has closest affinity with this genus (Teulon et al. 2013).

Megoura viciae Buckton Plate 25b Apterae are dark bluish green to apple-green with black head, prothorax, antennae, legs, siphunculi and cauda (see influentialpoints.com/Gallery); BL 3.0-5.0 mm. On young apical parts of stems of Leguminosae/Fabaceae, especially Lathyrus and Vicia spp. In Europe, Middle East, north and Central Asia, and Ethiopia. Some Asian populations have been given subspecies status (M. viciae ssp. abchasica Mordvilko, M. viciae ssp. turanica Nevsky), but with little apparent justification. Monoecious holocyclic with oviparae and alate males in August-September. Studies on this aphid included the classic work on photoperiodism and morph determination by Lees (1973), and the first characterisation of an aphid sex pheromone (Dawson et al. 1987). See also Blackman & Eastop (2000). 2n=10.

Megourella Hille Ris Lambers

Aphidinae: Macrosiphini

Two species related to Megoura but with longer siphunculi, a dorsal pattern of dark spinal, pleural and marginal sclerites in both apterae and alatae, and a habit of living on basal parts of their host plants. Blackman (2010) reviewed the genus.

Megourella purpurea Hille Ris Lambers Plate 25c Apterae are dirty reddish violet, pink or greenish, with black dorsal spots (see influentialpoints.com/Gallery); BL 2.1-2.9 mm. On Lathyrus pratensis, on basal parts at or near ground level. In north-west, northern, central and southern Europe, and also reported from Iran (Rezwani 2010). Monoecious holocyclic; oviparae and apterous males have been reared in October in the Netherlands (Hille Ris Lambers 1949).

Megourella tribulis (Walker) Apterae are dark green to black with black antennae, legs, siphunculi and cauda (see influentialpoints.com/Gallery); BL 2.4-3.0 mm. On Vicia sepium, feeding on base of stem at or near ground level. In north-west, northern and central Europe. Monoecious holocyclic; sexual morphs were found in late September-October in the Netherlands (Hille Ris Lambers 1949).

Megourina Hille Ris Lambers

Aphidinae: Macrosiphini

A genus for one western North American species superficially similar to Megourella but perhaps more closely related to nearctic Nasonovia (Kakimia), with 4-5 hairs on first tarsal segments and long, somewhat capitate hairs, the marginal ones being placed on unusual domed sclerites (Hille Ris Lambers 1974b).

Megourina lagacei Hille Ris Lambers Apterae are of unknown body colour; BL 2.5-3.0 mm. Alatae have black head and thorax, 47-55 secondary rhinaria on ANT III, and dark (not bordered) wing veins. On Aquilegia formosa in California. Biology and sexual morphs are unknown.

Meguroleucon Miyazaki

Aphidinae: Macrosiphini

A genus for one east Asian Macrosiphum-like species with an unusual pattern of dorsal sclerotisation and 4 hairs (including 2 sense pegs) on all first tarsal segments (Miyazaki 1971). [A second species described in this genus, M. longqishanense Zhang & Qiao (in Qiao & Zhang 1999c, 1999d), seems to be based on vagrant individuals of a Uroleucon sp.]

Meguroleucon codonopsicola Miyazaki Apterae are yellowish green with a milky tint, with brown head, black antennae and siphuculi, very large black postsiphuncular sclerites, and a pale cauda; BL 2.2-2.6 mm. Alatae have 37-44 secondary rhinaria on ANT III. On Codonopsis spp.in Japan and Korea (Lee et al. 2002c), living on stems and undersides of young leaves without causing any deformation. In Korea it has also been collected on Platycodon grandiflorum.Biology and sexual morphs are unknown.

Meitanaphis Tsai & Tang

Eriosomatinae: Fordini

Two or three little-known east Asian species with an elongate pterostigma tapering around curve of wing as in Schlechtendalia, but having 6-segmented antennae and galls with exit located at base.

Meitanaphis elongallis Tsai & Tang Galls are jujube-like in form, maximally 4.1 cm long, green, becoming red or purplish when mature with many fine longitudinal ridges. On undersides of leaflets of Rhus punjabensis var. sinica in China (Tsai & Tang 1946). Alatae (BL c. 1.3 mm) emerge through a large slit in autumn. The life cycle is unknown.

Meitanaphis flavogallis Tang Galls are pea-like to jujube-like in form with yellow hairs, on undersides of leaflets of Rhus punjabensis var. sinica in China (pictured in C. Zhang et al. 2008). The description of this species has not been seen, but the antennal rhinariation was described by Yang et al. (2009). The morphology of the antennal rhinaria and DNA analyses (Yang et al. 2010; Ren et al. 2017, 2019) all indicate that it may be more correctly placed in Kaburagia.

Meitanaphis microgallis Xiang Galls are small, papilliform, on Rhus potaninii in China (Xiang 1980/81). In the DNA analysis of Yang et al. (2010) this species clustered with Kaburagia, but a misidentification may be involved as this does not accord with the original description of the forewing having an elongate pterostigma tapering around its tip.

Melanaphis van der Goot

Aphidinae: Aphidini

About 25 palaearctic species with short siphunculi, closely related to Rhopalosiphum, and with the same association with Rosaceae (Pyroidea) and Poaceae, although only a few species have proven host alternation. Alatae have dark forewing veins with media twice-branched. Three species are European, but most of the remainder are native to east Asia and associated either with Miscanthus, or with Arundinaria and related bamboos. The taxonomy of the east Asian species is in need of further study. Accounts are available for the Iberian peninsula (Nieto Nafría et al. 2005a), Japan (Sorin 1970b), China (L. Zhang et al. 2001), Taiwan (Liao 1976), India (Raychaudhuri & Banerjee 1974, A.K. Ghosh & L.K. Ghosh 2006) and north-east India (Raychaudhuri et al. 1980b). Halbert & Remaudière (2000) compared species in the miscanthi/sacchari group.

Melanaphis arthraxonophaga Zhang, Qiao & Zhang Colour of apterae in life is unknown; BL c. 1.1 mm. On Arthraxon hispidus in China (Liaoning Province; L. Zhang et al. 2001).

Melanaphis arundinariae (Tissot) Apterae are yellowish-brown with darker brown head and sides of body; BL c. 1.6mm. Dense colonies were found in April on Arundinaria gigantea ssp. tecta in Florida, USA (Tissot 1933). The life cycle is unknown. [This species, known only from the type series, was for many years placed in Rhopalosiphum but was transferred to Melanaphis by Skvarla et al. (2019).]

Melanaphis bambusae (Fullaway) Apterae on bamboos are dark brown to black with waxy grey dorsal abdominal markings, legs and antennae mainly pale; BL 0.8-1.4 mm. Alatae (on bamboos) have secondary rhinaria distributed III 14-28, IV 7-15, V 0-7. Host-alternating in Japan between Photinia villosa and bamboos (Sorin 1962); elsewhere (and also commonly in Japan) populations are anholocyclic on bamboos (Arundinaria, Bambusa, Phyllostachys). Ant-attended. Indigenous to east Asia, now in south-east Asia, Australia, India, the Mediterranean area and southern Europe (Egypt, Turkey, Morocco, France, Italy, Greece, Spain, Portugal, Madeira), USA (Louisiana, Hawaii), and a population has been found on Phyllostachys vivax in UK under glass (R. Hammon, pers. comm.). Yi & Su (1985) studied predation by a coccinellid in China. 2n=8, 10 or 12 (the reason for the discrepancy needs further investigation).

Melanaphis daisenensis (Sorin) Apterae are blackish, with yellow antennae and tibiae and black tarsi: BL c.1.5-1.6 mm. On Miscanthus sp. in Japan. Very closely related to, and possibly not distinct from, M. miscanthi.

Melanaphis donacis (Passerini) Apterae in life are dark brown with an extensive and distinctive pattern of white wax (see influentialpoints.com/Gallery and aphids of Karnataka website); BL 1.5-2.2 mm. Alatae have secondary rhinaria distributed III 3-13, IV 0-3, V 0. On Arundo donax, and also recorded from Phragmites spp. Attended by ants. In southern Europe, the Mediterranean region, Azores, Madeira, Canaries, north Africa, Middle East, Central Asia, eastward to India and Pakistan and introduced to Argentina and Chile (Ortego et al. 2004). Monoecious holocyclic on Arundo in southern France, with apterous males (BMNH collection), but commonly also anholocyclic. 2n=8.

Melanaphis elizabethae (Ossiannilsson) Apterae are dark brown, slightly wax-powdered; BL 2.0-2.7 mm. In inflorescences and under leaves of Phragmites australis (= communis). In northern Europe (Scotland, Denmark, Sweden, Finland – Heikinheimo 1997). Monoecious holocyclic (Heie 1986).

Melanaphis graminisucta Zhang Apterae are reddish brown; BL 1.5-1.8 mm. On an unidentified grass species in Hunan Province, China.

Melanaphis indosacchari (David) Apterae are dull dark brown; BL 1.1-1.5 mm. In ant-attended colonies on exposed upper leaves of Saccharum officinarum in south India, and also sometimes on Iseilema laxum. Some specimens in mixed populations with M. sacchari on Echinochloa colonum in the Philippines (BMNH collection, leg. M.R. Gavarra) seem also to be M. indosacchari (or M. sorini, if the two are indeed distinct species). Varma et al. (1978) studied its ecology and control measures in India.

Melanaphis jamatonica (Sorin) Colour of apterae in life is unknown; BL 1.3-1.8 mm. On Miscanthus sp. in Japan. Specimens from Miscanthus sinensis in Korea (BMNH collection, leg W.H. Paik) will also key to jamatonica,but differ from the Japanese material by lacking scent glands on the hind tibiae.

Melanaphis japonica (Takahashi) Apterae are yellow or purple, dusted with wax (Moritsu 1983); BL 1.3-1.6 mm. On Miscanthus sinensis in Japan, and also recorded from Korea (Lee et al. 2002c – but these specimens might be M. jamatonica. Oviparae and alate males occur in September, and eggs are laid on undersides of leaves (Sorin 1970b). 2n = c.22.

Melanaphis koreana (Sorin) Apterae are pale yellowish grey; BL 1.5-1.8 mm. Alatae were described by W.K. Lee & Seo (1992). On Miscanthus sinensis in Korea and east Siberia (Pashchenko 1988a). [Specimens in the BMNH collection, determined by Sorin as M. koreana (leg. W.H. Paik, Yangi, Korea, 11.vi.61), have shorter siphunculi than the type material (1.0-1.1 × longer than basal width, compared with c.1.45 × basal width)] .

Melanaphis luzulella (Hille Ris Lambers) Apterae are dark green, almost black; BL 1.2-1.8 mm. On Luzula spp., in spring and early summer at stem bases and on upper surfaces of basal parts of leaves, tented over by ants, and later also onto aerial shoots. In northern, central and eastern Europe. Monoecious holocyclic with apterous males (Heie 1986). Prior (1971) described the ovipara.

Melanaphis meghalayensis Raychaudhuri & Banerjee Apterae are greenish to dark brown, often dusted with wax; BL 1.3-1.9 mm. Alatae have secondary rhinaria distributed III 18-20, IV 4-8. On undersurfaces of leaves, or on stems at ground level, of unidentified grasses and bamboos in Meghalaya and West Bengal, India (Raychaudhuri & Banerjee 1974). A population described as a subspecies, M. meghalayensis bengalensis, with fewer hairs on the cauda and genital plate, was recorded from Arundinaria sp. in West Bengal. The life cycle has not been studied. Agarwala & Mahapatra (1990) described oviparae of M. meghalayensis from an unidentified bamboo, indicating that this species may be partly holocyclic and monoecious on Poaceae. However it is not clearly differentiated morphologically from the secondary host form of M. pahanensis. M. grossisiphonellus Zhang, Qiao & Zhang, described from a Bambusa sp. in China (L. Zhang et al. 2001), is possibly this species.

Melanaphis miscanthi (Takahashi) Apterae are yellowish grey, dirty yellow or dark purple; BL 1.2-1.9 mm. On Miscanthus sp. in Taiwan. This species was redescribed by Halbert & Remaudière (2000).

Melanaphis montana (Sorin) Apterae are greenish dark brown (but see also Moritsu 1983, p.184); BL c.1.2-1.3 mm. On Miscanthus sinensis in Japan, and also found on Leersia in Thailand (BMNH collection, leg. Napompeth). Very close to, and possibly synonymous with, M. tateyamaensis.

Melanaphis pahanensis (Takahashi) Host-alternating between Pyrus spp. and unidentified Poaceae in northern India (Raychaudhuri & Banerjee 1974, Medda & Chakrabarti 1992). Apterae on Pyrus in spring are dark brown dusted with wax; BL 1.8-2.1 mm. They feed on the undersides of the leaves, causing them to curl. Alatae emigrating from Pyrus have secondary rhinaria distrubuted III 15-28, IV 0-1; return migrants (gynoparae) have them distributed III 51-58, IV 18-28, V 12-15, VI 0-3 (Medda & Chakrabarti 1992). It was originally described (Takahashi 1950, as Aphis pahanensis) from Malaya, as brownish purple apterae secreting cottony wax, BL c.1.25 mm, on an unidentified bamboo of the Arundinaria group, and has subsequently been found on Pennisetum purpureum (BMNH collection, leg. S.K. David). Swirskiaphis bambuciepula Zhang & Zhang, described from Phyllostachys in China (L. Zhang & G. Zhang 2000c) is possibly this species. [Melanaphis pyrisucta Zhang & Qiao, described from Pyrus pyrifolia in China (G. Zhang et al. 1999d) is similarly long-haired, but alate spring migrants have many more rhinaria (III 31-37, IV 9-12, V 2-6).]

Melanaphis pyraria (Passerini) Pear‑Grass Aphid Apterae on Pyrus are dark brown, with an extensive brown-black dorsal abdominal patch (see influentialpoints.com/Gallery); BL 1.3-2.1 mm. In spring on undersides of pear leaves, rolling the leaves transversely or diagonally to the mid-rib, attended by ants (Andrasfalvy 1968). Host-alternating between Pyrus communis and Poaceae. Spring migrant alatae have secondary rhinaria distributed III 8-30, IV 0-9, V 0-2. Apterae on grasses are yellowish to reddish purple; BL 1.1-1.7 mm. Their colour differs according to host species and condition; on Arrhenatherum it is reddish purple, hidden under deformed leaves, while on Poa, Brachypodium and Triticum it is a smaller, yellowish aphid. Alatae produced on grasses have secondary rhinaria distributed III 23-45, IV 12-29, V 1-21 . Widely distributed in Europe, in Madeira, the Canaries and the Azores, Mediterranean region and Middle East, Caucasus and Transcaucasia. 2n=8 (the chromosomal distribution of heterochromatin and rDNA was studied by Criniti et al. 2004).

Melanaphis sacchari (Zehntner) (= formosana Takahashi 1921, 1931 nec. Sorin, 1970) Apterae are very variable in colour according to host plant and environmental conditions; pale yellow, yellow brown, dark brown, purple, or even pinkish, often with a variably developed black dorsal patch (e.g. see influentialpoints.com/Gallery); BL 1.1 ‑2.0 mm. Alatae have secondary rhinaria distributed III 4-16, IV 0(-6), V 0. In ant-attended colonies on Poaceae, especially Saccharum; also sometimes on other grasses and cereals (Arthraxon, Echinochloa, Oryza, Oryzopsis, Panicum, Pennisetum, Sorghum, Themeda, Thysanolaema), and there are also records from Araceae (Arum, Caladium). In South Africa, India, Sri Lanka, Laos, China, Philippines, Australia, Hawaii, USA, Central and South America. Probably anholocyclic almost everywhere that it occurs, but it is reported to be monoecious holocyclic (with alate males) in China (Zhang & Zhong 1983), with Miscanthus sacchariflorus as the overwintering host (Wang et al. 1961). Peña Martinez et al. (2016) have now reported sexual morphs in introduced populations of the M. sacchari/sorghi group on three sorghum species in Mexico. It still needs to be determined whether these populations are M. sacchari or M. sorghi (see below). Zimmerman (1948) gave a general account of the biology, including natural control by parasites and predators, in Hawaii.

Most of the ecological studies of M. sacchari on cultivated sorghum in Africa and Asia should probably be referred to M. sorghi (q.v.). Nibouche et al. (2014) studied genetic diversity of samples identified as M . sacchari from many parts of the world, and showed that certain genotypes have a wide distribution, but their multilocus genotypes “A” and “E” originating from Africa and Asia showed greatest genetic divergence from the rest and were almost all collected from Sorghum bicolor; further studies combining molecular and morphological data have now shown these to be M. sorghi (Nibouche et al. 2021). In the USA, populations identified as M. sacchari which prior to 2013 were limited to sugarcane in Florida and Louisiana, have recently become a major problem on cultivated sorghum in many south-eastern and central states (Medina et al. 2017; Nibouche et al. 2018), and this has now been shown to be due to the separate introduction of M. sorghi (Nibouche et al. 2021). Holt et al. (2020) have found differences between the microbiota of sugarcane and sorghum-feeding populations in the USA. The M. sacchari/sorghi/miscanthi group is closely related to the host‑alternating M. pyraria in Europe. 2n = 8 for samples from Hong Kong and India; but Khuda-Bukhsh & Kar (1990) recorded 2n=10 from India. [A record of 2n=22 for M. sacchari in Japan (Blackman 1986) is referable to another, undescribed species.]

Melanaphis siphonella (Essig & Kuwana) Apterae are dark brown, lightly dusted with white wax; BL 1.4-1.9 mm. On Pyrus spp. (betulifolia, pyrifolia) in Japan (Essig & Kuwana 1918), Korea (Paik 1965) and China (L. Zhang et al. 2001). Biology and life cycle are unknown.

Melanaphis sorghi (Theobald) Plate 8f Apterae are white or yellow, and sometimes larger individuals have a variably-developed black dorsal abdominal patch (see influentialpoints.com/Gallery); BL 1.1-2.0 mm. Alatae have secondary rhinaria distributed III 4-13, IV 0(-1), V 0. On Poaceae, especially Sorghum bicolor, favouring the axils of the lower leaves, but large colonies extend over the whole leaf. Also sometimes on other grasses or cereals (Eleusine, Panicum, Saccharum, Zea). M. sorghi occurs in Greece (Margaritopoulos et al. 2013), Cyprus, Israel, Iran (Eastop & Hodjat 1981, as M. sacchari), probably in Turkey (Kök & Özdemir 2021, as M. sacchari), and is widely distributed in Africa, east and south-east Asia (India, Pakistan, China, Thailand, Japan, Philippines). It has now been shown to be the species now attacking cultivated sorghum in the southern USA and Mexico (Nibouche et al. 2021, and see above under M. sacchari). Probably it is mostly anholocyclic, although Setokuchi (1975) reported a holocycle on Sorghum in Japan, and David (1977) recorded oviparae in February-March in north-west India. It is also likely that the introduced populations producing sexuales on Sorghum spp. in Mexico (Peña Martinez et al. 2016) are M. sorghi. 2n=8.

Melanaphis sorini Halbert & Remaudière (= formosana Sorin 1970, not Takahashi 1921, 1931) Apterae are wine red, purple or tan; BL 1.0-1.8 mm. On Miscanthus spp. in Japan and Taiwan, and introduced to USA (Florida, California; Halbert & Remaudière 2000). It has also been found in UK on Miscanthus imported from Israel (Hammon et al. 2006). Monoecious holocyclic in Japan, with alate males (Sorin 1970b, as Longiunguis formosanus); the Florida population may be anholocyclic. This species seems closely related to and morphologically almost indistinguishable from M. indosacchari, but the form introduced into Florida would not colonise Saccharum in laboratory tests (Halbert & Remaudière 2000).

Melanaphis strobilanthi Medda & Chakrabarti Colour of apterae in life is unrecorded, probably pale with darker head; BL 1.2-1.6 mm. On Strobilanthes sp. in Uttar Pradesh, India.

Melanaphis takahashii Skvarla & Miller (replacement name for Aphis arundinariae Takahashi; Skvarla et al. 2019) Apterae are purplish brown (Liao 1976); BL 1.2-1.7 mm. Alatae have numerous secondary rhinaria distributed III 26-38, IV 14-29, V 7-19. On undersides of leaves of bamboos of the Arundinaria group (Pleioblastus, Yushania niitakayamensis) at high altitude in Taiwan. The life cycle is unknown. Specimens from India described as M. arundinariae (e.g. Raychaudhuri & Banerjee 1974) have apterae with 6-segmented antennae and several other differences from the Taiwanese species, and are more likely to be the secondary host forms of M. pahanensis. Reports of Pyrus pashia as the primary host of this species, (e.g. Chakrabarti & Banerjee 1993b, Saha & Chakrabarti 1988d, as M. arundinariae) are also likely to be referable to M. pahanensis. Other long-haired Melanaphis from northern India, but with shorter siphunculi, have been described as a distinct species, M. meghalayensis (q.v.).

Melanaphis tateyamaensis (Sorin) Apterae are reddish purple; BL c. 1.6 mm. On undersides of leaves of Miscanthus sp. in Japan. Possibly these are larger specimens of M. montana.

Melanaphis yasumatsui (Sorin) Apterae are dark purple, with white powdery wax; BL 1.2-1.5 mm. On Miscanthus sp. in Japan, and recorded from M. sinensis in east Siberia (Pashchenko 1988a).

Melanaphis zhanhuaensis Zhang, Qiao & Zhang Apterae are purplish red; BL c.1.5 mm. On subterranean part of Imperata cylindrica var. major in China (Shandong Province; L. Zhang et al. 2001). Apart from the colour and subterranean habit this species seems indistinguishable from M. sorghi.

Melanocallis Oestlund

Calaphidinae: Panaphidini

One nearctic species near Tinocallis, but with differences in the ventral structure of the head, dark marginal sclerites on abdominal tergites 5-7, a squat body shape, infuscated basal parts of wings and a host association with Juglandaceae. It was placed as a synonym of Tinocallis by Remaudière & Remaudière (1997) but revalidated as a full genus by Quednau (2003).

Melanocallis caryaefoliae (Davis) (fig. 33A) All viviparae are alate, squat-bodied, very dark green to black with small tufts of white wax and conspicuous paired black dorsal tubercles, those on abdominal tergite 2 being especially large; BL 0.9-1.9 mm. On both upper and lower sides of leaves of most Carya spp., but especially C. illinoiensis. Feeding causes yellow spots on leaves, reducing photosynthesis (Wood & Tedders 1986), and large populations can cause defoliation. Apterous oviparae and alate males are produced in October (Hottes & Frison 1931, as M. fumipenella); the ovipara is illustrated by Quednau (2003). The effect of M. caryaefoliae on the yield of pecans was studied by Wood et al. (1987), and Dutcher (1985) evaluated chemical control measures. For a general account see Bissell (1978). 2n=14.

Melaphis Walsh

Eriosomatinae: Fordini

Three or possible more nearctic species related to the east Asian genera of Fordini of subtribe Melaphidina (Nurudea, Kaburagia, Schlechtendalia), and with a similar alternation between Rhus and mosses. A recent study using both morphological and mtDNA data placed it closest to Schlechtendalia (Pang et al. 2011; although see also Ren et al. 2017). The antiquity of the host association was discussed by Moran (1989). Foottit & Maw (2017) have now shown using a combination of molecular and morphological data that at least three cryptic species in North America have been confused under the name Melaphis rhois, and describe as a new species one group of populations that can be discriminated morphologically from M. rhois.

Melaphis asafitchi Foottit & Maw Forming sac-like white or green galls (when immature – becoming red when mature) on Rhus spp. similar to those of M. rhois, in eastern Canada and north-eastern USA (see influentialpoints.com/Gallery) . The length of the ultimate rostral segment (RIV+V) of adult apterae in the galls is less than 0.067 mm. Migration occurs to unidentified mosses. Foottit & Maw (2017) provided more complex functions to discriminate most individuals of generations on Rhus from M. rhois, but a definitive diagnosis is only possible using mitochondrial COI-5P sequence. Moss-feeding populations cannot currently be discriminated from those of M. rhois.

Melaphis rhois (Fitch) Forming closed, sac-like white or green galls (when immature – becoming red when mature; fig. 129, and see influentialpoints.com/Gallery) on leaflets of Rhus spp. (glabra, typhina) in North America. The length of the ultimate rostral segment (RIV+V) of adult apterae in the galls is more than 0.067 mm (cf. M. asafitchi). Emigrant alatae (BL 1.4-1.7 mm) emerge in late August-October and fly to mosses. Apterae on mosses are pale with darker antennae and legs; BL 0.8-1.2 mm; a detailed redescription of them was provided by Pike et al. (2012; but this could possibly apply to M. asafitchi). Anholocyclic populations also occur all-year-round on mosses, and have been introduced into Europe (Sweden, UK), although following the work of Foottit & Maw (2017) the specific identity of these European populations is uncertain. Hebert et al. (1991) studied genetic variation in local populations in Ontario, Canada by enzyme electrophoresis, and found two groups differing in allele frequency profiles, which perhaps correspond to the two species recognised by Foottit & Maw (2017). The complete mitochondrial genome has been published (Ren & Wen 2017; although it is unclear whether this is M. rhois or M. asafitchii (or an undescribed species). 2n=26*.

Meringosiphon Carver

Greenideinae: Cervaphidini

One western Australian species with 6-segmented antennae, long dorsal body hairs arising from low tuberculate bases, and siphunculi with hairs on basal half as well as subapically. Alatae have forewings with twice-branched media.

Meringosiphon paradisicum Carver Apterae are probably brownish; BL 1.55-1.76 mm. The host plant is uncertain; the type specimens were swept from areas where Daviesia divaricata was the dominant vegetation (Carver 1959), but the host has now been recorded as Gastrolobium dilatatum (Brumley 2020). In Western Australia.

Mesocallis Matsumura

Calaphidinae: Pterocallidini

Eleven east Asian species principally associated with Betulaceae, resembling Tinocallis except for the lack of spinal tubercles, but probably more closely related to Pterocallis, differing from that genus in that the spinal and marginal hairs on abdominal tergites 1-4 are usually unduplicated. Accounts are available for Japan (Higuchi 1972), China (Qiao et al. 2005b, J. Chen et al. 2020), the Korean peninsula (including DNA barcoding; Y. Lee et al. 2017, 2018) and India (Chakrabarti 1988). Quednau (2003) reviewed the genus as a whole, illustrating all available morphs of species then known, and placed Paratinocallis, previously regarded as closer to Pterocallis, as a subgenus of Mesocallis.

Mesocallis alnicola Ghosh Colour in life is unrecorded, probably pale yellow. Apterous as well as alate viviparae are described; BL of apterae 1.0-1.2 mm, of alatae 1.3-1.4 mm. Alatae have 9-11 secondary rhinaria on ANT III, distributed along most of length of segment. On Alnus nepalensis in Meghalaya, India (A.K. Ghosh 1974). Biology and sexuales are unrecorded.

Mesocallis carpinicola Y. Lee Adult viviparae are all alate, pale yellow with antennae, legs and forewing vein Cu_1b having similar dark pigmentation to M. pteleae; BL 1.3-1.6 mm. On the undersides of leaves of Carpinus laxiflora in South Korea (Y. Lee et al. 2018). Morphological discrimination from the very similar M. pteleae is mainly size-related and may not be entirely reliable.

Mesocallis (Paratinocallis) corylicola (Higuchi) Adult viviparae are all alate, pale yellow with usually only ANT V and VI, apices of tibiae and tarsi pale brown; BL c. 1.5 mm. Described from Corylus sieboldiana in Japan (Higuchi 1972), and also recorded from C. heterophylla in Korea (Quednau 1979) and China (Qiao et al. 2005b). Sexual morphs and life cycle are unknown. 2n=10 (Blackman 1986).

Mesocallis fagicola Matsumura Alatae are very small, pale yellow; BL c. 0.6 mm. ANT III has about 6 secondary rhinaria spaced out along segment. Described from Fagus sylvatica in Japan (Matusumura 1919), and not recorded since. Biology and sexuales are unknown.

Mesocallis obtusirostris Ghosh Alatae are pale yellow; BL 1.2-1.5 mm. ANT III has 3-7 rhinaria, mostly on basal part. On Alnus nepalensis (with one record from Betula alnoides) in India (Uttar Pradesh, West Bengal) and Nepal (Chakrabarti 1988). Sexual morphs were reported by Chakrabarti (1988, p.69).

Mesocallis (Paratinocallis) occulta Y. Lee Alatae are pale yellow, with legs and antennae mainly pale; BL 0.9-1.5 mm. On undersides of leaves of Corylus spp. in South Korea (Y. Lee et al. 2018). Revealed by DNA barcoding to be distinct from M. corylicola (Y. Lee et al. 2017); the morphological discriminants given in the key between these two species may not be entirely reliable.

Mesocallis platycaryae Qiao Apterae are translucent white; BL c.1.0-1.2 mm. Living dispersed on undersides of leaves of Platycarya strobilacea (Juglandaceae) in Anhui Province, China (J. Chen et al. 2020). Alatae are unknown.

Mesocallis pteleae Matsumura Alatae are pale yellow, with ANT III brown-black, rest of antennal flagellum dark-banded, dark front of head, mainly brown fore tibiae and tarsi, mid and hind legs pale (except for dark spots at ends of hind femora), and a very darkly-bordered forewing vein Cu_1b (see influentialpoints.com/Gallery); BL 1.2-1.6 mm. Described originally (Matsumura 1919) from Ptelea trifoliata, which is unlikely to be a true host. Since recorded from species of Alnus, Betula, Corylus, Ostrya and Carpinus (although records from Carpinus could be referable to M. carpinicola). In Japan, Korea and China.

Mesocallis sawashibae (Matsumura) Alatae and apterae are pale yellow, with darker antennal joints, dark distal half of fore tibia, and a black spot near apex of hind tibia; BL 0.8-1.5 mm. On Carpinus cordata and C. japonica, in Japan, Korea, China and east Siberia. Oviparae occur in October in Japan (Higuchi 1972). 2n=10 (Blackman 1986).

Mesocallis taoi Quednau Colour in life is unknown, but alatae are probably pale except for black front of head, ANT I-III and distal parts of ANT IV-VI, black fore tibiae and forewing vein Cu_1b; BL 1.0-1.2 mm. Apterae also occur, and are pale except for dark antennal joints and tarsi and black spots at bases of dorsal hairs; BL 0.9-1.2 mm. On Alnus cremastogyne in China (Quednau 2003).

Mesocallis (Paratinocallis) yunnanensis Zhang Colour in life is unknown, BL c.2.1 mm. On Corylus heterophylla in China (Yunnan; Zhang & Zhong 1985c). Described as a subspecies of M. (P.) corylicola and raised to full species status by Quednau (2003).

Mesothoracaphis Noordam

Hormaphidinae: Nipponaphidini

One species on Loranthaceae and Viscaceae in south-east Asia with sclerotised, box-like body and greatly reduced antennae and legs.

Mesothoracaphis rappardi (Hille Ris Lambers & Takahashi) (Fig.60a) Apterae are brownish-black, somewhat shiny; BL 0.95-1.14 mm. On young shoots or older twigs of Dendrophthoë and Viscum spp. in Java (Noordam 1991).

Mesotrichosiphum Calilung

Greenideinae: Greenideini

A genus with nominally three species close to Eutrichosiphum, but typically with 4-segmented antennae in the apterae and sparse, mostly short hairs, which are mostly blunt in apterae and acute in alatae.

Mesotrichosiphum brevisetosum Noordam Apterae are pear-shaped, colour in life is unknown; BL c.1.2 mm. Described from a single specimen collected on Lithocarpus bennettii in Java (Noordam 1994).

Mesotrichosiphum pentaiarticulatum Zhang & Qiao Apterae are pear-shaped, with head and thorax black or blackish green, abdomen brown, and black siphunculi; BL 1.4-1.7 mm. Alatae are undescribed. On young leaves of Castanopsis hainanensis in Hainan, China (D. Zhang & Qiao 2008c). This species would probably be more appropriately placed in Eutrichosiphum, and is close to if not synonymous with Eutrichosiphum nigrum.

Mesotrichosiphum uichancoi Calilung Apterae are pear-shaped, colour in life is not recorded; BL c. 1.3 mm. The single described alata has 5-segmented antennae with 13-15 secondary rhinaria along the length of ANT III, forewings with a twice-branched media and hindwings with one oblique vein. On undersides of young leaves and branch tips of Premna odorata in the Philippines (Calilung 1967). The life cycle is unknown.

Metanipponaphis Takahashi

Hormaphidinae: Nipponaphidini

Eight East Asian species distinguished from Nipponaphis by the ornamentation of the tergum, consisting of rounded pustules which are rather evenly spaced and sized over the central part of the prosoma and which, under the light microscope, appear double-walled or ring-like (fig. 36C). The fused abdominal tergites 2-7 have marginal hairs but there are no spinal hairs on abdominal tergite 7, and abdominal tergite 8 has 2 (or rarely, 4) hairs. Two species in Japan have been shown to alternate between Distylium and Castanopsis (Sorin 1987), while most of the other species that probably belong in the genus are only known from Fagaceae. The Japanese species have now been thoroughly reviewed by Aoki et al. (2021), and as a result of their molecular phylogenetic analysis the species originally described in the genera Monzenia and Lithoaphis are now placed in Metanipponaphis. Aoki et al. (2021) provide a key to the Japanese species living on Castanopsis; A.K. Ghosh & Raychaudhuri (1973a) provided a partial key, and Noordam (1991) gave a detailed generic diagnosis. Qiao et al. (2018) reviewed and keyed three Chinese species.

Metanipponaphis assamensis Ghosh & Raychaudhuri Apterae are aleyrodiform, oval, “light brown to yellow” (darker in life?); BL 1.7-1.8 mm (A.K. Ghosh & Raychaudhuri 1973a). On Castanopsis tribuloides in Meghalaya, India. Other morphs and life cycle are unknown.

Metanipponaphis cuspidatae (Essig & Kuwana) Host-alternating between Distylium racemosum and Castanopsis spp. The galls on the twigs of Distylium are large and globular (Aoki et al. 2021). Emigrant alatae emerging in autumn (BL 1.8-2.1 mm) give rise to dark purple, lightly wax-dusted aleyrodiform apterae (BL 1.5-1.7 mm) on branches of Castanopsis spp.(Aoki et al. 2021). Records from other plants (e.g. Castanea, Cinnamonum, Ficus, Quercus, Litsea) are likely to be misidentifications. Alate sexuparae apparently return to Distylium in May, but anholocyclic populations also remain on Castanopsis throughout the year (Takahashi 1962a). In Japan, Taiwan and China.

Metanipponaphis echinata Ghosh Apterae are aleyrodiform, oval, “light to dark brown” (prepared specimens?); BL 0.7-0.9 mm (A.K. Ghosh 1974). Described from Castanopsis histrix in Meghalaya, India. The pustules on the pleuromaginal region of the prosoma are elongate, tapering to a blunt apex, and often have a hooked profile; specimens in the BMNH collection from C. carlesii and C. sclerophylla in Hangzhou, China (leg. V.F. Eastop) are rather larger but have similar ornamentation, and may be this species. The feeding position of the Indian aphids was not recorded, but the Chinese aphids were feeding on the leaves. Other morphs and life cycle are unknown.

Metanipponaphis globuli (Monzen) (New combination; Aoki et al. 2021). Small green spherical galls are formed on twigs of Distylium racemosum, usually in leaf axils. There is no host alternation. According to Sorin (1960), there are only 4 generations per year in Japan. Alate sexuparae (BL c. 1.4 mm) emerge in late October, and produce sexuales on undersides of leaves. Eggs are laid on the twigs and do not hatch until early September of the following year, the fundatrices mature rapidly and produce one generation of apterae in the galls, which then produce the sexuparae. The sexuparae, like those of other monoecious Metanipponaphis, have dimorphic embryos. There are reports that alatae emerge from galls twice a year, in June and October (Monzen 1954; but see Aoki et al. 2021, who provide further information on life cycle and biology). Itô & Hattori (1983) reported gall predation by Nola innocua (Lepidoptera). In Japan and Korea (Paik & Choi 1969). The male genitalia were described and illustrated by Wieczorek et al. (2012).

Metanipponaphis ihai (Sorin) (New combination; Aoki et al. 2021). Galls on twigs of Distylium racemosum in southern Japan are pale yellowish green, spherical, soft and rather rough-surfaced, 2-4 cm in diameter (fig. 130D). Alate sexuparae (BL c. 1.8 mm) emerge in November-December and deposit dimorphc first instars on undersides of leaves of Distylium,which develop into males and dark reddish purple adult oviparae in mid-January, and eggs are laid on the twigs (Sorin 1996). Hatching of eggs, fundatrix and gall development have not been described. This species is very similar to M. cuspidatae; differences are discussed by Aoki et al. (2021).

Metanipponaphis lithocarpicola (Takahashi) Apterae are aleyrodiform, almost circular, black, usually with a little white wax marginally; BL 1.0-1.3 mm. On undersides of leaves of Lithocarpus sp. in Taiwan (Takahashi 1933), and there is a record from Castanopsis sclerophylla in Fujian Province, China (Chen et al. 2014). Other morphs and life cycle are unknown.

Metanipponaphis rotunda Takahashi Host-alternating in Japan between Distylium racemosum and Castanopsis spp. The galls on Distylium are formed on leaf petioles, and sometimes on the fruit. They are pale green and rather slender, 1.7-2.8 cm long (Sorin 1987). Alatae migrate to Castanopsis spp., producing aleyrodiform apterae which are black, almost circular, dorsally depressed, BL 1.0-1.3 mm, and live on undersides of leaves. Alatae sexuparae are produced on leaves of Castanopsis sieboldii in April to early May (Aoki et al. 2021). This species could not be included in the key to aphids on Distylium as the emigrant alatae from the galls do not seem to have been properly described. Only known from Japan. Sorin (1996) described the inhabitants of yellowish green galls on D. racemosum in Okinawa as a subspecies, M. rotunda nakijinensis.

Metanipponaphis shiiae (Takahashi) (New combination; Aoki et al. 2021). Apterae are black, shining, almost circular, and strongly convex dorsally; BL 1.2-1.5 mm. Alatae have secondary rhinaria distributed ANT III 21-23, IV 10-11, V 8-11. Apterae are found on branches of Castanopsis cuspidata in Japan; immature alatae feed along the midribs on the undersides of leaves. The life cycle is unknown; apparently anholocyclic on Castanopsis in Japan (Takahashi 1959a).

Metanipponaphis shiicola Takahashi Apterae on Castanopsis are aleyrodiform, black, oval, dorsally not much depressed; BL 1.1-1.5 mm. On Castanopsis cuspidata in Japan, living on undersides of leaves along basal part of mid-rib (Takahashi 1959). Molecular studies by Aoki et al. (2021) now indicate that this species migrates from galls on Distylium racemosum.

Metanipponaphis silvestrii (Takahashi) Apterae are aleyrodiform, dark brown, almost circular; BL c.1.0-1.3 mm. Described from leaves of an unknown host plant in Shansi, China (Takahashi 1935). Qiao et al. (2018) provided a full redescription of immature and adult apterae identified as this species from two species of Quercus in China. A.K. Ghosh (1974) collected similar apterae in Assam, India, on Lindera sp. Probably the species is misplaced in Metanipponaphis. Other morphs and life cycle are unknown.

Metanipponaphis vandergooti Noordam Appearance in life is unknown, probably dark; BL of aptera 1.6-1.8 mm. On Lithocarpus sundaicus in Java. An alata collected in August had BL c.2.2 mm and secondary rhinaria distributed ANT III 15, IV 7, V 3 (Noordam 1991). The life cycle is unknown.

Metathoracaphis Sorin

Hormaphidinae: Nipponaphidini

One species with host alternation from Distylium to Quercus in Japan, in which the apterae have very short unsegmented antennae and prosoma partially fused with abdominal plate, and the alatae as well as the apterae lack siphuncular pores.

Metathoracaphis isensis Sorin (= distylicolens Sorin) Galls protrude from the upper sides of the leaves of Distylium racemosum and are yellowish green, semicircular or helmet-shaped, 1.0-1.6 cm in height. Alatae (with secondary rhinaria distributed ANT III 12-16, IV 7-9, V 5-7) emerge in late June and migrate to Quercus gilva. Apterae living on upper sides of leaves of Q. gilva are aleyrodiform, oval, somewhat convex dorsally, strongly sclerotised, blackish brown dusted with greyish wax; BL c.1.0 mm. Alate sexuparae produced in October are blackish brown with dark forewing veins, and have secondary rhinaria distributed ANT III 40-47, IV 15-19, V 6-9. Partly anholocyclic; apterae were found on Quercus throughout the year Sorin (2004) confirmed the life cycle by host transfers and described the ovipara.

Metopeuraphis Narzikulov & Smailova

Aphidinae: Macrosiphini

One species in Central Asia, related to Metopeurum but living on Amaranthaceae and with prothorax and abdominal tergites 2-4 usually having well-developed marginal tubercles.

Metopeuraphis atriplicis Narzikulov & Smailova Apterae are green dusted with greyish wax, BL c.1.7-1.9 mm. In small colonies on undersides of leaves of Atriplex sp(p). in Kazakhstan. Kadyrbekov (2011c) recorded it from A. cana. Biology and sexual morphs are unknown.

Metopeurum Mordvilko

Aphidinae: Macrosiphini

About ten palaearctic species on Compositae, mostly described from central and eastern Europe, with six of the species known only from Ukraine. They are related to Macrosiphoniella but their morphology, and also their physiology (see Woodring et al. 2006), are adapted for ant attendance. Mamontova & Tshumak (1994) provided a key to species.

Metopeurum achilleae Bozhko Apterae are pale reddish brown, wax-dusted; BL c.1.8 mm. On flower stalks of Achillea gerberi in Ukraine. This species is very similar to M. capillatum.

Metopeurum borystenicum Bozhko Apterae are pale bluish or reddish green; BL c.1.9 mm. In dense colonies on stems and leaves of Tanacetum vulgare in Ukraine, and also recorded from T. boreale in Kazakhstan (Kadyrbekov 2017a).

Metopeurum buryaticum (Pashchenko) (new combination) Apterae are pinkish lilac, in life with large blackish violet dorsal abdominal spot; BL c.2.8 mm. On flower stems of Tanacetum vulgare, attended by ants (Pashchenko 1999b, as Macrosiphoniella (Sinosiphoniella)). In Siberia (eastern shore of Lake Baikal), and also recently recorded from Georgia (Barjadze et al. 2010b). [Except for the long dorsal body hairs the description closely matches that of M. fuscoviride.]

Metopeurum capillatum (Börner) Colour of apterae in life is unrecorded; BL 2.1 mm (one specimen). On Achillea sp., possibly either atrata or moschata. In Austrian Alps.

Metopeurum enslini (Börner) Apterae are blackish; BL 2.6-3.0 mm. On stems and root collars of Tanacetum vulgare. In Germany, and Ukraine (Bozhko 1963).

Metopeurum fuscoviride Stroyan Plate 27g (Fig.52a) Apterae have a dark brown head and thorax, reddish, pink or green abdomen with a black central patch (disappearing in preserved specimens), and black siphunculi and cauda (see influentialpoints.com/Gallery); BL 1.8-2.9 mm. In ant-attended colonies on Tanacetum vulgare, and occasionally on other Tanacetum spp., with single records also from Achillea millefolium, Leucanthemum vulgare and Artemisia alba (the latter from Kazakhstan as ssp. kasachstanica; Smailova 1974). In Europe, west Siberia, Iran and Central Asia. Monoecious holocyclic with oviparae and apterous males in late August-September. Loxdale et al. (2011) analysed variation within and between local populations in Germany using microstatellite markers. Senft et al. (2017) studied its small-scale population dynamics. Lysiphlebus hirticornis is a host-specific parasitoid. 2n=8.

Metopeurum gentianae Mamontova & Tshumak Apterae are dark green, with yellow antennae and legs; BL not given. On Gentiana asclepiadea in Ukraine.

Metopeurum matricariae Bozhko Apterae are reddish brown with darker dorsal abdominal spots, black siphunculi and a pale cauda; BL c.2.2 mm. In dense colonies on stems of Matricaria chamomilla in Ukraine, where it has also been recorded from Tripleurospermum inodorum (Mamonotova-Solukha 1966). It is also reported to occur on Matricaria sp. and Tanacetum (=Pyrethrum) tanacetoides in Kazakhstan (Kadyrbekov 2014d).

Metopeurum millefolii Mamontova & Tshumak Apterae are ash-grey; BL 1.6-1.8 mm. On Achillea millefolium in Ukraine, and also recorded from Kazakhstan (Kadyrbekov 2011b).

Metopeurum urticae Mamontova & Tshumak Apterae are yellow with dark antennae, head, legs and siphunculi ; BL 2.1-2.4 mm. On undersides of leaves of Urtica dioica in Ukraine.

Metopolophium Mordvilko

Aphidinae: Macrosiphini

A little more than 20 species resembling Acyrthosiphon but perhaps more closely related to Sitobion, and like that genus typically with Rosaceae as primary hosts and migrating to Poaceae. The generic positions of the few species with other host associations are uncertain. Accounts are available for north-west Europe (Stroyan 1982a, Heie 1994, Blackman 2010) and northern India (Raychaudhuri et al. 1980). Blackman et al. (1989) discussed enzyme differences between closely-related European species.

Metopolophium albidum Hille Ris Lambers Apterae are pale straw yellow to pale yellowish green, without a green spinal stripe, but sometimes with green spots at bases of siphunculi (see influentialpoints.com/Gallery); BL 1.9-2.7 mm. On grasses such as Arrhenatherum elatius, sometimes in large numbers causing red and yellow discoloration of grass blades (Stroyan 1950). Throughout Europe. Monoecious holocyclic with oviparae and alate males in November (original description). 2n=16.

Metopolophium alpinum Hille Ris Lambers Apterae are dull green with pale appendages; BL 2.6-3.8 mm. Alatae have dusky narrow dorsal abdominal cross-bands. In spring colonies on Rosa spp. in alpine habitats in France, Austria and Switzerland. It is now also recorded from west Siberia (Stekolshchikov & Khureva 2020). Presumably heteroecious holocyclic, migrating to grasses, as it has been found on Poa alpina in Austria. It is also recorded from Carex. Oviparae and alate males occur on wild Rosa leaves in October (original description).

Metopolophium arcticum Stekolshchikov & Khureva Colour in life unknown, and apterous viviparae probably not produced; BL of fundatrix 1.7-2.7 mm. Host plant uncertain, but pitfall-trapped in areas where Dryas punctata was predominant vegetation. In arctic west Siberia (Stekolshchikov & Khureva 2020). Life cycle abbreviated, with oviparae and apterous males trapped along with fundatrices in July-August.

Metopolophium arctogenicolens Richards Apterous viviparae are undescribed, oviparae are pale green: BL 2.0-2.4 mm. Described from oviparae and one apterous male collected in July-August onTaraxacum arctogenum (= ceratophorum) on Ellesmere Island, North-West Territories, Canada. Specimens were maintained on this plant in the laboratory (originaldescription).

Metopolophium chandrani (David & Narayanan) (= M. graminum Raychaudhuri, L.K. Ghosh & Das, = M. simlaense Chakrabarti & Raychaudhuri; Eastop & Blackman 2005) Apterae are yellowish green; BL 1.9-2.6 mm. On grasses (Bromus, Poa) in India. Oviparae and alate males were collected (along with apterous viviparae) on Rubus opulifolius in Himachal Pradesh in December (Agarwala & Mahapatra 1990). Also recorded from Rosa sp., and from an unidentified shrub (as M. simlaense).

Metopolophium (Metopolophinum) darjeelingense L.K. Ghosh Apterae are greenish with brighter green spinal stripe, appendages pale except for apices of antennae; BL 2.2-2.5 mm. On Hypericum sp. in West Bengal; the host plant is unusual for a member of this genus, but is confirmed by a subsequent collection (Raychaudhuri, M.R. Ghosh & R.C. Basu 1978, as M. davidi). The species is also unusual in having spinulose femora and 4 hairs on first tarsal segments, justifying its placement in a separate subgenus. [Note: M. darjilingense Raychaudhuri, M.R. Ghosh & R.C. Basu (1978) is a different species, from an unknown host; see also under M. lacheni.]

Metopolophium dirhodum (Walker) Rose-Grain Aphid Plate 23h Apterae are green or yellowish green with a brighter green spinal stripe, and have antennae with dark apices to each segment (see influentialpoints.com/Gallery); BL 1.6-2.9 mm. Alatae have a rather pale abdomen, sometimes with indistinct brownish cross-bars on anterior tergites. In Europe it occurs on wild and cultivated Rosa spp. in spring, and is heteroecious holocyclic, migrating in June to numerous species of Poaceae and Cyperaceae. It is a major pest of cereals, and has become widely distributed especially in temperate regions of the world (see Blackman & Eastop, 2000). Some overwintering on grasses occurs in western Europe, and populations introduced to other parts of the world – or at least those in New Zealand (Nicol et al. 1997) and Brazil (Lopes-da-Silva & Vieira 2007) – seem to be permanently parthenogenetic. 2n=18.

Metopolophium fasciatum Stroyan Apterae are pale green with a darker green spinal stripe, frequently with yellow spots at bases of siphunculi; BL 1.7-3.4 mm. On grasses, especially Arrhenatherum elatius. Only known from UK, except for a single aptera from Iceland (BMNH collection, leg. R.N.B. Prior). Potentially heteroecious holocyclic, as there is an autumn migration and production of oviparae on Rosa canina, but males are rare in comparison with those of M. dirhodum, and spring populations on Rosa have not been identified, so it is thought to be predominantly anholocyclic in UK (Stroyan 1982a). [We have examined type specimens of M. berberinutritum Zhang, Chen, Zhong & Li, described from (vagrants on?) Berberis in China (G. Zhang 1999); this has longer antennae (1.33 × BL as opposed to c.1.0 × BL in UK populations of fasciatum, but is otherwise very similar.] 2n=18*.

Metopolophium festucae (Theobald) Apterae are rather shiny, evenly yellowish green to green or salmon pink, with antennae progressivly darker from III to VI; BL 1.4-2.2 mm. Alatae have conspicuous dark dorsal abdominal cross-bands. On many genera and species of Poaceae in Europe (including Iceland), Iran and Central Asia (Kadyrbekov 2017a), and it has been introduced to New Zealand, but is not recorded from Australia (contrary to Teulon & Stufkens 2002). Populations on cereal crops show slight morphological differences and are regarded as a subspecies, M. festucae ssp. cerealium Stroyan. There are records of aphids resembling M. festucae from Argentina, Bolivia, and western USA, and populations in Chile and USA have been identified as ssp. cerealium (Remaudière et al. 1993, Halbert et al. 2013a). Monoecious holocyclic with alate males, but anholocyclic overwintering is common in both subspecies. See also Blackman & Eastop (2000). 2n=16 (including ssp. cerealium).

Metopolophium frisicum Hille Ris Lambers Apterae are shining green, with ill-defined olive or brownish green dorsal abdominal shield, and dark antennae, siphunculi and cauda; BL 1.3-2.4 mm. On Poa spp., especially P. trivialis, feeding on upper sides of leaf-blades in shady, damp situations. In north-west, northern and central Europe. An earlier record from North America (Börner 1952) is unsubstantiated (Foottit et al. 2006), but A. Jensen (aphidtrek.org) has collected samples from Washington, Oregon and Colorado that may be this species (see also Skvarla et al. 2017). Monoecious holocyclic with oviparae and apterous males in October (but anholocyclic in UK).

Metopolophium lacheni Agarwala, Mondal & Raychaudhuri Colour of apterae in life is unknown; BL c.1.8-2.4 mm. Alatae have dark dorsal abdominal cross-bands. On Rubus sp. in north-east India (Sikkim). Oviparae and alate males were collected in December (original description, as M. darjeeligense ssp. lacheni).

Metopolophium longicaudatum David & Hameed Apterae are pale; BL 2.2-2.9 mm. On wheat (Triticum sp.) in Himachal Pradesh, India. Other morphs are unknown.

Metopolophium montanum Hille Ris Lambers Apterae are evenly green or apple green, without a darker green spinal stripe; BL 1.8-2.8 mm. Alatae have conspicuous dark dorsal abdominal cross-bands. In spring colonies on wild Rosa spp. in montane regions of Switzerland, Austria, France and Spain. Presumably heteroecious holocyclic, migrating to grasses as subsequently found on Poa alpina in Austria. Oviparae were found on wild Rosa leaves in October (original description).

Metoplophium mukhamedievi Akhmedov Apterae are shining pale green: BL 2.2-2.9 mm. On Lonicera microphylla in Central Asia. This species does not seem to be clearly distinct from M. dirhodum.

Metopolophium palmerae (Hille Ris Lambers) Plate 23i Apterae are probably green: BL c.2.7 mm. On Elymus sp. in Colorado, USA.

Metopolophium pedicularus (Richards) Colour of apterae in life is unknown; BL c 2.8 mm. On Pedicularis seudetica in arctic Canada (North-West Territories). Monoecious holocyclic; oviparae and apterous males were collected in mid-August (original description). The generic position of this species is uncertain.

Metopolophium rogeri Mehparvar Apterae are green no powered; BL 2.0–3.0 mm. Monoecious holocyclic on Rosa beggeriana in Kerman province (Iran), with oviparae and apterous males in October (Mehaparvar et al. 2022).

Metopolophium rosaesuctum Zhang Apterae are reddish green; BL c. 3.4 mm. Alatae are undescribed. On Rosa sp. in China.

Metopolophium sabihae Prior Apterae are yellow-green, apple green or blue-green; BL 1.3-2.1 mm. On Festuca rubra, Vulpia membranacea and Ammophila arenaria growing in sand dunes. In western Europe (France, Belgium, UK). Aphids identified as this species have now been found on Wrangel Island in eastern Siberia, where fundatrices were collected in July and oviparae in July-August, and this species is becoming increasingly common (Stekolshchikov & Khruleva 2014; Khruleva & Stekolshchikov 2019). Anholocyclic, or partially monoecious holocyclic in UK, with oviparae and both apterous (or brachypterous) and alate males produced in culture in November (original description).

Metopolophium taimyricum Stekolshchikov & Khureva Colour in life unknown; BL of aptera 2.5-3.3mm. Only collected by sweeping vegetation so host plant(s) uncertain, but commonest in sweeps of Hedysarum hedysaroides ssp. arcticum. In the tundra zone in arctic west Siberia (Stekolshchikov & Khureva 2020). Life cycle abbreviated, with fundatrices, apterae, oviparae and apterous males all collected in July.

Metopolophium tenerum Hille Ris Lambers Apterae are dirty green or reddish, rather shiny; BL 1.3-2.2 mm. On Deschampsia flexuosa and Festuca spp. in north-western and northern Europe. A record from Turkey (Görür et al. 2011b) requires additional conformation. Monoecious holocyclic, with oviperae and alate males in October in the Netherlands (original description).

Mexicallis Remaudière

Calaphidinae: Myzocallidini

Seven or more very small species on undersides of leaves of Quercus in Mexico. Apterae have 4- or 5-segmented antennae and long thick spiculose spinal and marginal hairs, except subgenus Anacallis which has inconspicuous spinal hairs. Alate viviparae are unknown except for one specimen trapped in Panama. The genus was extensively revised by Quednau & Remaudière (1996), and reviewed again with illustrations of all species by Quednau (1999).

Mexicallis analiliae Remaudière Apterae are white to creamy white; BL 0.7-1.1 mm. Spinal hair-bearing processes are very variably developed; at least some are usually long and cylindrical. On Quercus spp. especially Q. rugosa, in Mexico. Alate males and an immature ovipara were described by Quednau & Remaudière (1996) from a population including apterous viviparae on Quercus sp. in October. Certain characters, such as the length of the last rostral segment, vary greatly between populations, and two subspecies have been described from Mexican populations, M. analiliae pumilus on a dwarf Quercus (?frutex) (Remaudière 1982b), and M. analiliae caulifer on Q. rugosa (Quednau & Remaudière 1996).

Mexicallis (Anacallis) areolatus Remaudière Apterae are whitish with dark green marginal spots on each segment; BL 0.8-1.2 mm. Placed in a separate subgenus (Anacallis) on account of the peculiar development of the frontal and marginal processes. and lack of spinal hairs except on abdominal tergites 7 and 8. On Quercus rugosa (and one specimen only found on Q. mexicana) in Mexico (Remaudière 1982b).

Mexicallis (Anacallis) brevituberculatus Quednau & Remaudière Colour of aptera in life is unknown; described from one specimen, BL 1.1 mm, on an unidentified Quercus sp. in Mexico (Quednau & Remaudière 1996).

Mexicallis calvus Remaudière Apterae are whitish: BL 0.7-0.9 mm. Spinal hairs on abdomen are of remarkably different sizes; those on abdominal tergites 2, 6, 7 and 8 are very long with tuberculate bases, whereas those on abdominal tergites 1,3,4 and 5 are minute. On Quercus sp. (?mexicana) in Mexico (Remaudière 1982b).

Mexicallis longicaudus Remaudière Apterae are similar to M. spinifer, and previously described as a subspecies of that species. On an unidentified Quercus sp. in Mexico.

Mexicallis panamensis Quednau & Remaudière Apterae are white; BL 0.67-0.8 mm. On an unidentified Quercus sp. in Panama (Quednau & Remaudière 1996), and on Q. oocarpa in Costa Rica (Villalobos Muller et al. 2010).

Mexicallis spinifer Remaudière Apterae are dark green, or pale greenish grey with two darker green irregular lateral bands running from pronotum to abdominal tergite 5 or 6; BL 0.7-1.0 mm. Dorsal chaetotaxy shows considerable variation (Remaudière 1982). On Quercus spp. (including crassipes, mexicana, rugosa) in Mexico. Quednau & Remaudière (1996) described the alate male found on an unidentified Quercus sp. in December, and Wieczorek et al. (2011) described and illustrated the male genitalia.

Micraphis Takahashi

Aphidinae: Macrosiphini

One species in east Asia of uncertain affinities; possibly related to Coloradoa, except that the eyes have distinct posterior ocular tubercles and although feeding on Artemisia, R IV+V is not stilletto-shaped.

Micraphis artemisiae (Takahashi) Apterae are yellow or greenish, BL c.1.3 mm. On leaves and stems of Artemisia capillaris in China, Taiwan and Japan. Alatae have 12-18 secondary rhinaria on III and 4-7 on IV. The life cycle is unknown.

Microlophium Mordvilko

Aphidinae: Macrosiphini

Four or five species mostly on Urtica,with siphunculi similar to Aulacorthum but with smooth, very large and somewhat divergent antennal tubercles.

Microlophium carnosum (Buckton) Plate 23e Apterae are various shades of green, pink or reddish purple (see influentialpoints.com/Gallery); BL 3.1-4.3 mm. Alatae have dark marginal sclerites but only faint spino-pleural markings. On undersides of leaves and up stems of Urtica spp., especially U. dioica. Also recorded from Girardinia cuspidata (Tao 1999). In Europe, Asia east to Pakistan and Mongolia, Africa (Burundi, Rwanda), North America and Chile (Fuentes-Contreras et al. 1997); a record from Argentina (Heie 1994) could not be substantiated. Monoecious holocyclic with alate males. Perrin (1976) studied its population ecology and natural enemies. 2n=20. [In UK another, undescribed species occurs on U. dioica which can be distinguished morphologically by the characters given in the key, and has 2n=16. This is possibly the species with 2n=16 from Crimea listed as M. evansi Theobald by Kuznetsova & Shaposhnikov (1963).]

Microlophium rubiformosanum (Takahashi) Apterae are green, with antennal and leg segments blackish distally, and siphunculi blackish except at bases; BL c.3.5 mm. Alatae are without dorsal abdominal markings. On Rubus spp.in Taiwan and Japan. 2n=12.

Microlophium sibiricum (Mordvilko) Apterae are brownish green or dark green; BL 3.4-4.4 mm. Alatae have dark spino-pleural as well as marginal markings. On undersides of leaves and up stems of Urtica spp., especially U. urens. In Europe (not UK) and across Asia to east Siberia, and possibly also in Japan (Miyazaki 1971, as Acyrthosiphon carnosus); also in USA, where populations occur mainly on U. dioica ssp. gracilis and are regarded as a subspecies, M. sibiricum ssp. tenuicauda Hille Ris Lambers, although there seem to be no consistent morphological differencesfrom the Old World form. Monoecious holocyclic, with apterous males.

Micromyzella Eastop

Aphidinae: Macrosiphini

About 12 mostly African fern-feeding species related to Micromyzus but with 2-3 hairs (one sense peg) on first tarsal segments, and alatae have more normal wing venation, with wing veins dark but not heavily bordered, radial sector more moderately curved and media once- or twice-branched. Apterae often have dark siphunculi and dusky or dark dorsal sclerotisation. Remaudière & Autrique (1985) revised the genus, providing keys to apterae and alatae.

Micromyzella anisopappi Remaudière Apterae are pale with black siphuncular apices and cauda; BL c.1.5-1.7 mm. Alatae have secondary rhinaria distributed III 10-15, IV 2-4, V 0(-2), and variably developed dorsal abdominal markings. On Anisopappus africanus, often in company with Sitobion hirsutirostris. In Burundi.

Micromyzella davalliae Remaudière Apterae are dark with black appendages incl. cauda; BL 1.4-2.5 mm. Alatae have secondary rhinaria distributed III 26-42, IV 12-23, V 4-11, and dorsal abdomen with dark cross-bands. Common in the rainy season on the epiphytic fern Davallia chaerophylloides (= denticulata) growing on oil palms in Burundi (Remaudière & Autrique 1985). Monoecious holocyclic, producing oviparae and alate males in late February-April.

Micromyzella eliei Remaudière Apterae are black, except for basal parts of antennae and femora, tibiae and cauda; BL 1.6-2.1 mm. Alatae have secondary rhinaria distributed III 30-34, IV 22-26, V 10-11 (one specimen). On Helichrysum odoratissimum in Burundi.

Micromyzella filicis (van der Goot) Apterae are grass-green, sometimes with a black dorsal abdominal spot, black antennae and siphunculi; BL c. 1.7 mm. Alatae have 8-14 secondary rhinaria on ANT III only and no dark dorsal abdominal spot. On undersides of fronds, or in furled fronds, of ferns in many genera (Antrophyum, Asplenium, Blechnum, Pityrogramma, Polypodium, Pteris, Sphenomeris, Thelypteris). In Java (Noordam 2004), and introduced to Australia (Brumley 2020) and New Zealand, where it was collected on Platycerium sp. (BMNH collection, leg. V.F. Eastop). 2n=36 (in New Zealand).

Micromyzella judenkoi (Carver) Apterae are undescribed; BL of alatae 1.2-1.7 mm. Alatae have secondary rhinaria distributed III 9-18, IV 0-8, V 0-4. Described from alate viviparae and an alate male caught in yellow trays in Australia and Sri Lanka. Subsequently apterae collected on ferns (Asplenium, Athyrium, Cheilanthes)in northern India have been assigned to this species, but without a published description. Alatae have also been trapped in Hong Kong (described by Qiao & Jiang 2005) and the Philippines (BMNH collection).

Micromyzella kathleenae Remaudière (Fig.43q) Colour of apterae in life is unrecorded, probably rather pale, with dark siphunculi and black cauda; BL 1.7-2.0 mm. Alatae have secondary rhinaria distributed III 15-23, IV 12-16, V 8-12, and paired dark patches on abdominal tergites. On Asplenium aethiopicum (= praemorsum) in Burundi.

Micromyzella pterisoides (Theobald) (Fig.43p) Apterae are dark, with antennae and legs mainly dark, black siphunculi and paler cauda ; BL 1.8-2.2mm. Alatae have 7-15 sec. rhin. on III only and an often extensive dusky or dark dorsal abdominal patch. On ferns (Arthopteris, Asplenium, Davallia, Drynaria, Nephrolepis, Osmunda, Pleopeltis) in eastern and southern Africa. Oviparae and alate males occur in colonies in small numbers in rainy season (Remaudière & Autrique 1985).

Micromyzella sleonensis (Eastop) (Fig.43o) Apterae are brown; BL 1.4-1.6 mm. Alatae have secondary rhinaria distributed III 2-9, IV 0-2. On stems of ferns of family Adiantaceae (Adiantum, Pellaea), described from Sierra Leone, and also found in Kenya and South Africa (BMNH collection).

Micromyzella sophiae Remaudière Colour of apterae in life is unrecorded, probably rather pale with variably developed dark sclerotic dorsal markings, antennae, siphunculi, and distal parts of femora dark, and a pale cauda; BL 1.5-1.9 mm. Alatae have secondary rhinaria distributed III 14-24, IV 8-15, V 2-6 and paired dark patches on abdominal tergites. On Asplenium aethiopicum (= praemorsum) in Burundi, in mixed colonies with M. kathleenae.

Micromyzodium David

Aphidinae: Macrosiphini

Eight Asian species on a range of host plants including ferns, and related to Micromyzus and Micromyzella,but with long dorsal body hairs. Su et al. (2012) reviewed the species in China. Life cycles are unknown except for one species, M. kuwakusae, recently shown to have a sexual phase on Loropetalum (Hamamelidaceae).

Micromyzodium dasi Verma Apterae are bright yellow with dark head, antennae and siphunculi; BL c. 1.9-2.0 mm. Alatae have secondary rhinaria distributed III 20-35, IV 10-20, V 0-2. Described from undersides of fronds of an unidentified fern, and subsequently recorded (including an alate male) from Adiantum caudatum and Cheilanthes sp. (Saha & Chakrabarti 1988d). In north-west India.

Micromyzodium filicium David Plate 17b Apterae are shining black, antennae dark beyond III, legs dark at femoral apices, siphunculi dark at base and apex but with middle part white; BL 1.5-2.0 mm. Alatae have 12-16 secondary rhinaria on ANT III only. In large numbers on undersides of fronds or on young growth of various ferns (Adiantum, Asplenium, Cheilanthes, Eriosorus, Nephrolepis, Pityrogramma, Polypodium, Pteris, Thelypteris). Also recorded from plants in some angiosperm families; Didymocarpus and Streptocarpus (Gesneriaceae), Aristolochia (Aristolachiaceae), Cypripedium (Orchidaceae) and Geranium (Geraniaceae). The number of such records suggests that these were not all vagrants, but this extended host range needs further confirmation. In Pakistan, north and south India, and Nepal. 2n=12.

Micromyzodium kuwakusae (Uye) Apterae are yellow with mainly dark thorax and a large black dorsal abdominal patch; BL 1.4-1.8 mm. Alatae have a black patch and secondary rhinaria distributed III 29-37, IV 12-22, V 5–9. On Fatoua villosa (Moraceae) in Japan, and now also recorded from Pilea notata and Urtica fissa (Urticaceae) in China (Su et al. 2012) and Korea (Y. Lee et al. 2014a). This species was described in Macrosiphoniella and regarded as a nomen dubium in catalogues, but rediscovered by M. Miyazaki (pers. comm. 1979), who donated specimens to BMNH collection. Sugimoto (2017b) has now shown that it is heteroecious in Japan, with Loropetalum chinense as the primary host, and described oviparae, males and the spring generations. Oviparae and fundatrices are yellow without a black dorsal abdominal patch, and the patch is also reduced in early spring generations. [Note: the aphids from Pilea notata listed by Su et al (2012) under the name M. kuwakusae have the same collection data and identification number as specimens described as Kaochiaoja pileophaga Zhang in Zhang et al. (1992a), so Su et al. presumably consider the latter name to be a synonym of M. kuwakusae]

Micromyzodium levipes (R.C. Basu & Raychudhuri) Colour of apterae in life is unrecorded; BL c.1.5 mm. Described from Chromolaena (=Eupatorium) odorata in India (Arunachal Pradesh).

Micromyzodium nipponicum (Moritsu) Apterae are dark brown (immatures are pale bluish grey); BL 1.4-1.6 mm. On Perilla frutescens var. crispa and Perilla sp. in Japan, Taiwan, Korea (Y. Lee et al. 2014a) and mainland China (Hunan), where it is also recorded from Agastache rugosa and Cirsium japonicum (an unlikely host) (Su et al. 2012). The colonies on Perilla are formed on the roots.

Micromyzodium polypodii Takahashi (Fig.43j) Apterae are yellow with blackish antennae, siphunculi and cauda; BL c. 2 mm. Alatae have forewing veins broadly bordered with fuscous, and black dorsal abdominal markings including a large central patch. On unidentified ferns in Japan, and now also recorded from China, with the host identified as Parathelypteris glanduligera (Su et al. 2012).

Micromyzodium spinulosum Miyazaki Apterae are shining black, with yellowish brown antennae and legs, black siphunculi and a pale cauda; BL 1.1-1.5 mm. Alatae have 7-13 secondary rhinaria on ANT III. On stems of grasses (Digitaria adscendens, Oplismenus compositus) in Japan. 2n=10.

Micromyzodium strobilanthi L.K. Ghosh Apterae are blackish brown, with antennae and legs mainly yellowish, with distal parts of femora dark brown, and dark siphunculi; BL 1.7-2.1 mm. Alatae are undescribed. On Strobilanthes dalhousianus (Acanthaceae)in northern India. One greenish black aptera also identified as this species was collected from Ruellia tuberosa (Raychaudhuri et al. 1980).

Micromyzus van der Goot

Aphidinae: Macrosiphini

About 10 species of mostly east Asian fern-feeding aphids close to Micromyzella but with usually 4 hairs (incl. 2 sense pegs) on first tarsal segments, and alatae having dark-bordered wing veins and a strongly curved radial sector.

Micromyzus diervillae Matsumura Apterae are pale green or green; BL c.1.9 mm. Alatae have forewings with a pale pterostigma (cf. Neotoxoptera weigeliae)and veins narrowly bordered with fuscous, and dorsal abdomen with dark cross-bands. On Weigela spp. in Japan, Korea and east Siberia. Probably heteroecious holocyclic, with migration to an unknown secondary host (a fern?). The fullest description is that by Takahashi (1965b, as M. weigelae).

Micromyzus hangzhouensis Zhang Apterae have 1-3 rhinaria on ANT III, siphunculi longer than head width across eyes, and first tarsal segments with 3 hairs; BL c. 2.6 mm. On Glochidion puberum in China (Zhang & Zhong 1980a). Probably the species is incorrectly placed in Micromyzus, which are typically small fern feeders.

Micromyzus katoi (Takahashi) Plate 17c (Fig.43r) Apterae are black (original description) or shining orange-brown, paler spinally and darker on margins and at bases of siphunculi, antennae and legs banded black and white, siphunculi blackish-brown, cauda pale (Noordam 2004); BL 1.5-2.1 mm. Alatae have secondary rhinaria on ANT III only, 2-6 (Taiwan) or 8-14 (Java, Australia), and dark-bordered wing-veins. On undersides of fronds of ferns in certain genera (Microsorium, Platycerium, Polypodium). Described from Taiwan, and subsequently recorded from Indonesia, Australia, Hawaii and New Caledonia (Mille et al. 2020). However, at least some of the non-Taiwanese material may be a distinct species with different colour in life, a relatively longer R IV+V, and alatae with more secondary rhinaria.

Micromyzus mawphlangensis A.K. Ghosh Apterae are pale with a brown spinal patch on abdominal tergites 1-3, dark brown siphunculi and a pale cauda; BL c.2.1-2.2 mm. Alatae have 3-9 secondary rhinaria on ANT III only, brown-bordered wing veins and dusky segmental spino-pleural abdominal sclerites. On a Polypodium sp. in northern India.

Micromyzus niger van der Goot (Fig.43k) Apterae are shining brown-black, antennae, femora, tibiae and siphunculi pale with black tips, cauda very dark grey, almost black, with white wax on distal part (Noordam 2004); BL 0.9-1.3 mm. On undersides of fronds of ferns, which may be curved inward. Recorded from ferns in several genera (Adiantum, Eriosorus, Pityrogramma, Pteris). Specimens from the grass Dactyloctenium aegyptiacum in the BMNH collection are presumably vagrants. In Indonesia, Singapore, India, Pakistan and Sri Lanka.

Micromyzus nikkoensis Miyazaki Apterae are whitish to pale yellow with antennae black, tibiae mainly dark, siphunculi pale with dark tips, cauda pale; BL 1.5-1.7 mm. On Athyrium (= Deparia) pycnosorum (Miyazaki 1971), Osmundastrum asiaticum (Moritsu 1983) and Gymnogramma totta (BMNH collection, leg. R.L. Blackman)in Japan. 2n=12.

Micromyzus osmundae Takahashi Apterae are yellow with black antennae and tibiae, and pale siphunculi and cauda; BL c. 2 mm. Alatae are undescribed. On Osmunda spp. in Japan.

Micromyzus platycerii Mróz & Depa Apterae are brown, shiny, with reddish eyes; BL 2.1-2.6 mm. Alatae have 11-14 secondary rhinaria on ANT III and dark-bordered wing-veins. On young shoots of Platycerium coronarium, occurring in large numbers on undersides of leaves, in Thailand.

Micromyzus pojanii (Cermeli & Smith) Apterae are shiny brown with dusky to black appendages; BL 1.6-1.8 mm. Alatae have strongly bordered wing veins and secondary rhinaria distributed III 13-16, IV 1-4. On an unidentified broad-leaved fern in Venezuela (Cermeli & Smith 1979, as Picturaphis), and also recorded from Phlebodium pseudoaureum in Costa Rica (Zamora Mejías et al. 2012).

Micromyzus vandergooti Noordam Apterae are orange brown or brownish red, with black siphunculi and a pale cauda; 1.3-1.7 mm. On ferns of genus Pyrrosia (as Cyclophorus, Drymoglossum), found living (in one case) on the upperside of a leaf “protected by a roof” (Noordam 2004). In Java.

Microparsus Patch

Aphidinae: Macrosiphini

About 11 nearctic and neotropical species on Leguminosae. They are mostly brightly-coloured aphids and alatae have dark-bordered wing veins. Subgenus Picturaphis have clavate siphunculi. Apterae have marginal tubercles on abdominal tergites 2-4. Cermeli & Smith (1979) provided keys to the species of Picturaphis.

Microparsus (Picturaphis) brasiliensis (Moreira) Plate 19e Apterae are shiny greenish dark brown to black (immatures are paler, brownish yellow); BL 1.6-2.0 mm. Alatae are dark brown to black with wing veins heavily black‑bordered. On various Leguminosae/Fabaceae (Centrosema, Dolichos, Indigofera, Phaseolus, Vigna); widely distributed in the Caribbean, Central and South America, and also recorded from Florida, USA. Biology is unknown; presumably mainly anholocyclic.

Microparsus desmodiorum Smith & Tuatay Apterae are pale yellowish green to green with head dark yellow to brownish orange, antennae, legs and siphunculi mainly dark, cauda yellow-orange; BL 1.5-2.0 mm. On Desmodium spp., in eastern USA, feeding on terminals and seed-pods without distorting growth. Oviparae and alate males occur in October-November (original description).

Microparsus olivei Smith & Tuatay (Fig.26c,d) Apterae are dark green, with head orange to brownish, antennae and legs bicoloured yellow and black, siphunculi black and cauda orange; BL 1.4-1.9 mm. On Desmodium spp. in eastern USA, feeding on leaves and terminals and causing some curling, and also recorded from Mexico, Cuba, Panama, Venezuela, Puerto Rico and Costa Rica (Quirós et al. 2009; Villalobos Muller et al. 2010). Oviparae and alate males occur in September-October (original description).

Microparsus (Picturaphis) puertoricensis (Smith) Apterae are shining reddish brown; BL 1.3-1.9 mm. On Phaseolus adenanthus in Puerto Rico. Alatae were described by Smith & Gaud (1974).

Microparsus rhynchosiae Remaudière & Peña Martinez Apterae are shining green, with antennae black except at base, dark apices to femora, and tibiae, tarsi and distal halves of siphunculi, and a dark band on abdominal tergite 8; BL 1.7-2.1 mm. On a Rhynchosia sp. in Mexico, colonising undersides of leaves of prostrate stems at soil level. Monoecious holocyclic, with oviparae and alate males in October (original description).

Microparsus singularis (Hottes & Frison) Apterae are bright green with orange-brown head, black antennae except for base of segment III, black and yellow legs, black siphunculi and yellow-orange cauda; BL 1.9-2.2 mm. On Lespedeza spp., especially L. repens, in eastern USA. Monoecious holocyclic with oviparae and apterous males in October (Smith & Heie 1963, as Megouroparsus kislankoi).

Microparsus tephrosiae (Smith) Apterae are amber to reddish brown, BL 2.3-2.7 mm. Described from Tephrosia virginiana, on which large colonies may occur and cause stunting of growth. There are also records from Astragalus and Cracca (but the Cracca species involved are now placed in Tephrosia). In eastern USA (Florida, Mississippi, North Carolina). Oviparae and alate males occur in October (original description).

Microparsus variabilis Patch Apterae are green with head and cauda yellow to dark orange, antennae and legs variably pigmented, siphunculi wholly dark or paler towards bases; BL 1.6-2.0 mm. On Desmodium spp. and Amphicarpaea monoica in eastern USA. It tightly curls the leaves of Desmodium, and lives on stems and undersides of leaves of Amphicarpaea (Hottes & Frison 1931). Oviparae and alate males were found in mid-August (Smith & Tuatay 1960).

Microparsus (Picturaphis) venezuelensis (Cermeli & Smith) (Fig.26a,b) Apterae are dull greenish to bluish black (venezuelensis s. str.), or dull reddish brown (M. venezuelensis ssp. meridensis); BL 1.8-2.6 mm. On Desmodium spp. in Venezuela, and also recorded from Argentina (Ortego et al. 2004). The two subspecies in Venezuela are separated by differences in pigmentation, and they possibly also colonise different species of Desmodium.

Microparsus (Picturaphis) vignaphilus (Blanchard) Apterae are dark brown to blackish; BL 1.9-2.4 mm. On several genera of Leguminosae/Fabaceae (Cologania, Phaseolus, Vigna)in South America (Argentina, Brazil, Colombia), Mexico, and Panama (Quirós et al. 2009).

Microsiphoniella Hille Ris Lambers

Aphidinae: Macrosiphini

Four North American species on Anthemideae, related to the palaearctic genus Microsiphum, but with much longer dorsal hairs. Three of the species have a dark sclerotic tergum. Smith & Knowlton (1938) and Palmer (1952) reviewed the genus (as Microsiphum).

Microsiphoniella acophorum (Smith & Knowlton) Apterae are pear-shaped, shiny brownish black to black, with pale ANT III and mainly pale tibiae (see aphidtrek.org); BL 1.0-1.5 mm. Colonising stems and leaves of Seriphidium tridentatum and Artemisia longifolia in western USA. Oviparae and alate males occur in September-October (Palmer 1952).

Microsiphoniella artemisiae (Gillette) Plate 27h Apterae are blackish brown to cinnamon brown, with base of ANT III and most of tibiae pale; BL 1.6-2.0 mm. On stems and leaves of Artemisia and Seriphidium spp. Widely distributed in North America. Oviparae and apterous males occur in September-October (Palmer 1952).

Microsiphoniella canadensis (Williams) Apterae are pale green, BL c. 2.4 mm. On stems and leaves (causing some leaf-curl) of Artemisia and Seriphidium spp. in Nebraska, USA, and there are also records from Oregon and Kansas (Smith & Parron 1978). This species, originally placed in Cryptosiphum, has not been adequately described, and its generic placement is problematic.

Microsiphoniella oregonensis (Wilson) Apterae are shining chocolate brown, with ANT III-IV pale and tibiae mainly pale; BL c.1.2-1.4 mm. On leaves and stems of Seriphidium tridentatum in western USA (Palmer 1952). It has also been collected on Picrothamnus desertorum (= Artemisia spinescens) (aphidtrek.org)Oviparae and alate males occur in October.

Microsiphum Cholodkovsy

Aphidinae: Macrosiphini

About 10 palaearctic species forming ant-attended colonies on stems of Compositae, related to Macrosiphoniella but with much reduced siphunculi (which still have some polygonal reticulation) and a broad-based triangular cauda. The species in north-west Europe were reviewed by Heie (1995), and those in eastern Europe by Bozhko (1963). Kadyrbekov (2006) revised the genus and provided a key to species. They all seem to be strictly monophagous.

Microsiphum diversisetosum Kadyrbekov Apterae are brown, pear-shaped; BL 1.7-2.4 mm. On Artemisia abrotanum in north and west Kazakhstan (Kadyrbekov 2006).

Microsiphum giganteum Nevsky Apterae are broadly pear-shaped, pale rose-red or pale green with dark head, a broad brown band across the prothorax, and black antennae and legs; BL 2.1-3.2 mm. On stems of Artemisia spp., especially A. dracunculus, in Ukraine and Central Asia. Apart from its colour in life this species is difficult to distinguish from M. procerae.

Microsiphum jazykovi Nevsky Apterae are pear-shaped, shiny brown on head and thorax with abdomen deep green-black, and antennae and legs mainly dark brown-black; BL 1.7-2.2 mm. In dense colonies on upper parts of flower stems of Artemisia absinthium, attended by ants (original description), and since recorded from several other Artemisia spp. (Kadyrbekov 2017a). In Ukraine and Central Asia. Populations with a relatively shorter R IV+V in northern and central Europe were treated as a subspecies, M. jazykovi ssp. wahlgreniHille Ris Lambers by Kadyrbekov (2006). Fundatrices of this subspecies were described from Switzerland (Hille Ris Lambers 1947a; as M. wahlgreni).

Microsiphum millefolii Wahlgren Plate 27i (Fig.5d,e) Apterae are pear-shaped, yellowish or greyish red to dark brown with dark head, antennae, legs, siphunculi and cauda; BL 1.8-2.5 mm. On lower parts of stems of Achillea millefolium, attended by ants. Throughout most of Europe, and also recorded from Kazakhstan (Kadyrbekov 2011b). Monoecious holocyclic, with apterous males (Heie 1995).

Microsiphum nudum Holman ( = minus Bozhko; synonymy by Kadyrbekov 2006) Apterae are pear-shaped, greyish or greenish black; BL 1.75-2 mm. On Achillea spp (millefolium, nobilis), living on basal parts of stem and rootstock, in ant shelters.In eastern Europe (Bulgaria, Hungary, Slovakia, Ukraine). Monoecious holocyclic, with oviparae and apterous males in October (original description).

Microsiphum procerae Bozhko Apterae are pear-shaped, shining brown with dark antennae and legs; BL c. 1.9-2.1 mm. On upper parts of flower stems of Artemisia procera (= abrotanum)in Ukraine, northern Caucasus, Kazakhstan.A population from the same host plant in the southern Caucasus is regarded by Kadyrbekov (2006) as a subspecies, M. procerae ssp. subalpicumMamontova.

Microsiphum ptarmicae (Cholodkovsky) (= heptapotamicumKadyrbekov; synonymy by Kadyrbekov 2006) Apterae are dark brown; BL c.1.5-1.8 mm. Described from Achillea ptarmicae in Ukraine, and also recorded forming ant-attended colonies on roots and at bases of stems of Achillea millefolium in Kazakhstan (Kadyrbekov 2000, as M. heptapotamicum). Kadyrbekov (2006) discussed geographical variation in certain characters previously used in species differentiation.

Microsiphum pyrethri Bozhko Apterae are brown tinged with red; BL c.1.7 mm. On Pyrethrum millefoliatum (= Tanacetum millefolium)in Ukraine.

Microsiphum woronieckae Judenko Apterae are pear-shaped, dusky pink on thorax, abdomen reddish black, antennae black, legs black except for yellowish basal halves of femora; BL 2.2-3.0 mm. In compact ant-attended colonies on lower parts of stems of Artemisia vulgaris. In northern, central and eastern Europe, west Siberia, northern Caucasus and Kazakhstan. Oviparae and apterous males occur in September-October (Hille Ris Lambers 1947a). 2n=12.

Microunguis Tao

Hormaphidinae: Nipponaphidini

One species related to Thoracaphis but the aleyrodiform apterae are without dorsal hairs except on abdominal tergite 8, and have very short unsegmented antennae. It was erroneously transferred to Neothoracaphis by Eastop & Hille Ris Lambers (1976).

Microunguis depressa (Takahashi) Apterae are aleyrodiform, blackish brown, narrowly yellowish brown marginally, without wax; BL c. 0.9 mm. On undersides of leaves of an unidentified Quercus sp. in Taiwan (Takahashi 1933; Tao 1966, 1969), or possibly according to Tao (1999), on Phoebe sp. Other morphs and life cycle are unknown.

Mimeuria Börner

Eriosomatinae: Pemphigini

One palaearctic species related to Paraprociphilus, but the apterae on the secondary host have one-segmented tarsi and a very different association with the host plant.

Mimeuria ulmiphila (del Guercio) (fig.18B) Terminal leaf nests are formed on Acer spp., especially A. campestre, by inhibition of shoot growth, twisting and folding of leaves. Fundatrices are olive green-grey, covered with white wax wool; BL 3.5-4.5 mm. They give rise to numerous dark brown alatae (BL 2.6-3.3 mm), which fly over an extended period (June-November). Apterous exules live mainly on roots of Ulmus; they are yellow, thickly wax-powdered, BL 1.3-2.3 mm, encased singly in brown microrrhizal cysts (Marchal 1933, Krzywiec 1964). Such cysts have also been found on Rubus roots close to Ulmus (Vernon 1957). Sexuparae are produced in autumn and return to Acer, but anholocycly on Ulmus roots often seems to predominate especially in western Europe. Krzywiec (1964) suspected that anholocyclic overwintering of immature stages might also occur on bark of Acer. Recorded from Britain, France, Germany, Hungary, Greece, Italy, Netherlands, Poland, Russia, Turkey and also in Kazakhstan (Kadyrbekov 2017a). For a full account of the life cycle in Poland and descriptions of all stages see Krzywiec (1962, 1964).

Mindarus Koch

Mindarinae

Nine or more species of conifer-feeding aphids, four nearctic and four or more palaearctic, in which apterae have fused head and pronotum, eyes usually with 3 facets and well-developed wax glands. Alatae have forewings with an elongate pterostigma, tapering to a point at the wing apex, and a bluntly triangular cauda. Heie (1980) reviewed the European species, the biology of which was studied in detail by Nüsslin (1910). There are more recent accounts of this genus from China (G. Zhang & Qiao 1997d, G. Zhang et al. 1999c) and USA (Voegtlin 1995), and a world review with illustrations of all available morphs by Quednau (2010). Favret (2009) reported on mitochondrial DNA sequence data that suggest the existence of two undescribed species in the Rocky Mountain region. The complete mitochondrial genome of one species (M. keteleerifoliae) has been published (Y. Wang et al. 2015).

Mindarus abietinus Koch Plate 2c, d, (fig. 15B) Apterae are yellowish green, covered with wax wool, with antennae and tibiae distinctly darker (see influentialpoints.com/Gallery); BL 1.2-2.0 mm. Alatae (BL 1.5-2.7 mm) have dark dorsal abdominal cross-bands. On young shoots of Abies spp. (especially alba, nordmanniana), feeding between the needle bases. There are only 3 generations per year, alate and apterous progeny of the fundatrix producing very small apterous sexuales. The eggs, black covered with short white strands of wax, are laid in June-July, but do not hatch until the following spring. The male genitalia were described and illustrated by Wieczorek et al. (2012). Throughout Europe, the Middle East (Lebanon, Turkey), common on Abies sibirica in Kazakhstan (Kadyrbekov 2017a), and on Abies pindrow in northern Pakistan (Naumann Etienne & Remaudière 1995). There are also records from India, Tibet, China and Thailand, although some of these may be due to confusion with M. japonicus. Zhang & Qiao (1997d) described some alatae from China (Yunnan) as a subspecies, M. abietinus triprimesensori, but this was synonymised with M. abietinus by Quednau (2010). Records from North America should be referred to M. pinicolus (Favret 2009). M. abietinus may cause serious damage or kill young shoots, or cause deformation and loss of needles (Klein 1983). During heavy infestations many nymphs may fall onto non-hosts in the forest undergrowth, often reaching the adult stage on them (Naumann Etienne & Remaudière 1995). 2n=12.

Mindarus guatemalensis Favret & Nielsen Appearance in life is not described, but probably similar to M. abietinus; BL not recorded. On Abies guatemalensis on Guatemala, causing significant feeding damage, particularly by distortion of growing shoots (Favret & Nielson 2008).

Mindarus japonicus Takahashi (fig. 15C) Apterae are not described, but probably similar in life to M. abietinus. BL of alata is c. 2 mm (Takahashi 1931). On Abies spp., and also recorded from Cephalotaxa drupacea. An ovipara from Abies firma is illustrated by Quednau (2010). The male genitalia were described and illustrated by Wieczorek et al. (2012). In east Asia (India, Nepal, east Siberia, Japan, Korea, China).

Mindarus keteleerifoliae Zhang Appearance in life is unknown. Described from shoots of Keteleeria evelyniana in Yunnan, China (Zhang & Zhong 1984). Also recorded from Cunninghamia sp., and from Picea sp. (as M. piceasuctus Zhang & Qiao 1997d; synonymy by Quednau 2010). Alatae have secondary rhinaria distributed III 14-22, IV 0-5. The complete mitochondrial genome of this species has been published (Y.Wang et al. 2015).

Mindarus kinseyiVoegtlin Apterae are pale green covered with light wax, increasing with age, when posterior and marginal regions may extrude long wax filaments; BL 1.1-2.3 mm. Alatae have variably developed dorsal abdominal sclerotic cross-bands. Small (BL 0.7-1.0 mm) yellow-gold oviparae and blue to blue-green dwarf apterous males (BL 0.5-0.7 mm) are produced in June through to autumn, but populations also continue to reproduce parthenogenetically through the summer and into autumn. On Abies concolor in California (Sierra Nevada), and also found on this host in eastern Canada (Quednau 2010). A record from Turkey (Şenol et al. 2014b) is likely to be based on a misidentification. Voegtlin (1995) described all morphs and Ehler & Kinsey (1995) provided a detailed account.

Mindarus obliquus (Cholodkovsky) Apterae are yellowish green, covered in white wax wool; BL 1.2-1.9 mm. Alatae are similar to those of M. abietinus, but smaller (BL 1.0-1.6 mm); differences between the two species were described by Carter & Eastop (1973). On Picea spp. (engelmannii, glauca, sitchensis, but not abies), feeding between needles on new shoots in spring, but without deforming needles. In Europe, east to Turkey (or China), Canada and Alaska (BMNH collection). Presumably, on account of the host species colonised, M. obliquus is of nearctic origin. The life cycle is like that of M. abietinus, with small apterous sexuales in June-September. 2n=12, according to Robinson & Chen (1969), but a sample from P. glauca in British Columbia, Canada (leg. C.K. Chan) had 2n=8 (R.L. Blackman; unpublished data), indicating that there may be more than one species on Picea in Canada.

Mindarus pinicola (Thomas) Balsam Twig Aphid Appearance in life and biology are essentially as in M. abietinus, with which it has been long confused in the North American literature. Voegtlin (1995) observed that fundatrices and their apterous progeny of North American populations always had more wax glands than European populations and suggested that these records may all apply to a native species. He also noted that fundatricesproduced a much greater proportion of apterous progeny than the European species. The separate identity of the aphid on native American Abies (balsamea, fraseri, grandis) was then confirmed by Favret (2009) using multiple morphometric analysis of forewing characters, and the name pinicola was reinstated. This is therefore the aphid studied under the name abietinus by various workers in North America, e.g. Amman (1963; natural enemies in North Carolina), Saunders (1969; damage and control), Bradbury & Osgood (1986; chemical control), Mattson et al. (1989; increased susceptibility of infested trees to spruce budworm), Fondren et al. (2004; biological control by predators), and Doherty et al. (2017; possible effects of global warming).

Mindarus remaudierei Voegtlin Appearance in life is unrecorded; BL of aptera 2.0-2.4 mm. On Abies religiosa in Mexico (Voegtlin 1995). Quednau (2010) illustrated an ovipara collected in mid-April.

Mindarus victoria Essig (fig. 15A) Apterae are a soft, jade-green, covered with thick white wax wool; BL c. 1.9 mm. Alatae green, wax-covered, with narrow dark dorsal abdominal cross-bands; BL c. 2.5 mm. On shoot tips of Abies grandis in British Columbia, Canada. Oviparae and very small dusky apterous males occur in June. Essig (1939) described and illustrated all the morphs.

Misturaphis Robinson

Aphidinae: Aphidini

One species in Canada apparently related to Aphis but without marginal tubercles on abdominal tergites 1 and 7. Antennae are 5-segmented in both apterae and alatae.

Misturaphis shiloensis Robinson Apterae are greyish or silvery green to pale green, BL 1.05-1.7 mm. On Artemisia caudata (= A. campestris) in Manitoba, Canada. Alatae have secondary rhinaria distributed III 10-14, IV 1-3. Sexual morphs are unknown.

Miyazakia Stekolshchikov

Aphidinae: Macrosiphini

One East Asian species related to Sappaphis but with a strongly sclerotised tergum and stiff hairs on dorsal body and appendages.

Miyazakia ranunculi (Miyazaki) (Fig.45b,d) Apterae are dark reddish brown dorsally, yellowish brown to dark brown ventrally, with appendages mainly dark; BL 1.4-2.0 mm. Alatae have secondary rhinaria distributed III 48-87, IV 18-35, V 12-22. On Ranunculus japonicus (= grandis) in Japan, and also in Korea (BMNH collection). Heteroecious holocyclic, with a sexual phase on Photinia villosa; Stekolshchikov (2014b) described alate gynoparae, oviparae and alate males collected on this plant in Japan by M. Sorin.

Mollitrichosiphum Suenaga

Greenideinae: Greenideini

About 16 species. mostly feeding on Fagaceae, or on Alnus, but Meliosma also seems to be a favoured host, and several species may have more polyphagous tendencies (R. Zhang et al. 2012). Body often elongate, with long siphunculi. The genus is characterised by a row of transverse (?stridulatory) ridges on the hind tibia. The tergum is usually ornamented with conical spinules or nodules. Raychaudhuri (1956) reviewed the few species then recognised (as Metatrichosiphon), Takahashi (1962b) reviewed the Japanese species, and accounts are available for India (A.K. Ghosh & Agarwala 1993), Nepal (Das & Raychaudhuri 1983), Java (Noordam 1994) and China (D. Zhang & Qiao 2010). R. Zhang et al. (2011, 2012) studied phylogenetic relationships of seven species in China using molecular methods, and Liu et al. (2013) demonstrated co-evolution with Buchnera endosymbionts. Several species were described from unidentified hosts, so the treatment of the genus here is incomplete.

Mollitrichosiphum glaucae Takahashi Colour of apterae in life unrecorded, presumably similar to M. nigrofasciatum, as it was originally described as a subspecies of that species; BL c. 2 mm (Takahashi 1962b). On Quercus glauca in Japan, and a single aptera collected on Q. neglecta (= Q. myrsinifolia) in Hong Kong may also be this species (Martin & Lau 2011).

Mollitrichosiphum godavariense Das & Raychaudhuri Apterae are pale to dark brown, elongate, collected on undersides of young leaves of an unidentified Quercus sp. in Nepal. Siphunculi of apterae pale, 0.56-0.59 × BL. Alate oviparae, with siphunculi 0.75-0.8 × BL, were collected in October (Das & Raychaudhuri 1983). Possibly a synonym of M. tenuicorpus.

Mollitrichosiphum lithocarpi (Takahashi) Apterae are elongate, yellowish green, with antennae and legs pale yellowish brown; BL c. 2.5 mm (Takahashi 1931a). Siphunculi of apterae are long, yellowish green, c. 0.6 × BL. On undersides of young leaves of Lithocarpus spp. (glabra, uraiana) in Taiwan. Alatae and other morphs have not been described.

Mollitrichosiphum luchuanum (Takahashi) Apterae are pear-shaped, yellowish brown; BL 2.0-2.3 mm. Alatae black, with long black siphunculi 0.8-0.9 × BL (siphunculi of the apterous type specimen are missing). Described from an unidentified Quercus sp. in the Ryukyu islands (“Loochoo”; Takahashi 1930), and since recorded from mainland China (Fujian province; D. Zhang & Qiao 2010), where it has also been collected from Prunus persica and Meliosma rigida (R. Zhang et al. 2011). M. yamabiwae is possibly a synonym.

Mollitrichosiphum montanum van der Goot Apterae are pale, elongate-bodied; BL 1.8-2.4 mm. Siphunculi are long, slender, yellow to yellowish brown, 0.52-0.85 × BL in apterae and 0.8-0.9 × BL in alatae. On Alnus spp. (nepalensis, nitida), feeding on young shoots, attended by ants. In India (Assam, Uttar Pradesh, West Bengal), Bhutan (Chakrabarti & Das 2014), Nepal (BMNH collection, leg. D.Hille Ris Lambers), Tibet (Zhang & Zhong 1981, as alni) and China (Zhang & Zhong 1985, as alni). It is also recorded from Betula alnoides (Singh & Singh 2017c; possibly a misidentification of the host plant). A.K. Ghosh & Agarwala (1993) compared the description of montanum with specimens from the type locality and types of alni Ghosh, Ghosh & Raychaudhuri and alnifoliae Chakrabarti & Raychaudhuri, and concluded that they were all synonyms. Saha & Chakrabarti (1986) described the alate oviparae and males (as alnifoliae), collected on A. nepalensis in September-October.

Mollitrichosiphum nandii Basu Apterae are elongate pear-shaped, light brownish green mottled with darker green, antennae pale dusky, darker at apices of segments, legs pale brown; BL 1.9-2.6 mm. Siphunculi of apterae are pale to dark brown with darker tips, 0.4-0.5 × BL; siphunculi of alatae are blackish, 0.6-0.75 × BL. On tender parts of shoots of Alnus nepalensis, particularly the leaf axils, often in large numbers, attended by ants (A.N. Basu 1964). Described from West Bengal, India. It is also recorded from Betula alnoides (Singh & Singh 2017c; possibly a misidentification of the host plant). After examining type material, two other species, M. acutihirsutum Maity & Chakrabarti (1980), described from Uttar Pradesh, and M. buddleiae A.K. Ghosh, Banerjee & Raychaudhuri (1971c), are considered to be synonyms. M. buddleiae was described from a presumably casual occurrence on Buddleja in Sikkim, but specimens from Alnus nepalensis in Nepal (Das & Raychaudhuri 1983), and from Alnus sp. in China (Zhang & Zhong 1985f), were subsequently identified as this species, and in China it has subsequently been collected from Alnus cremastogyne, Fagus longipetiolata and Psidium guajava (R. Zhang et al. 2011). Saha & Chakrabarti (1986) described the alate oviparae and males, collected in September-October. 2n=16*.

Mollitrichosiphum nigriabdominalis Agarwala, Mondal & Raychaudhuri Colour of apterae in life is not recorded, abdomen with a central dark patch; BL 2.2-3.0 mm. Siphunculi are pale, 0.5-0.6 × BL. On Quercus sp. (identified as Q. rubra, in which case non-native) in Sikkim, India (Agarwala et al. 1982). Other morphs and biology are unknown. From the description M. trilokum Agarwala & Ghosh (in A.K. Ghosh & Agarwala 1993) would appear to be large specimens of nigriabdominalis.

Mollitrichosiphum nigrofasciatum (Maki) Apterae are pear-shaped, pale yellow-green with an extensive and very distinctive brown-black mark covering the pleural areas of the thorax and abdomen on each side and joined medially across abdominal tergites 3-4 (for illustration see Moritsu 1983); BL 1.4-2.0 mm. Alatae have a broad brown-black patch across abdominal tergites 3-5. Siphunculi of both morphs are black; c. 0.35 × BL in apterae and 0.5-0.6 × BL in alatae. On Quercus spp., and also recorded from Lithocarpus spp. (Takahashi 1931a, R. Zhang et al. 2011). Records from Castanopsis spp. (Q. Liu et al. 2013) require further confirmation. In Japan, China and Taiwan. Sexuales and life cycle are unknown.

Mollitrichosiphum nigrum Zhang & Qiao Apterae are black with thin dusting of white wax, elongate pear-shaped; BL 2.2-2.6 mm. Siphunculi are black, in aptera 0.6-0.7 × BL, in alata 0.8-1.1 × BL. On Meliosma spp., and also collected from Ailanthus altissima and Elaeagnus pungens, in China (D. Zhang & Qiao 2010).

Mollitrichosiphum niitakaensis (Takahashi) Apterae are elongate-bodied, yellow, with pale antennae, legs and siphunculi; BL 1.9-2.7 mm. Siphunculi of apterae are 0.3-0.45 × BL; siphunculi of alatae are dark brown, 0.5-0.6 × BL. On undersides of leaves of Quercus sp(p). in Taiwan (Takahashi 1937b), and on Castanopsis sp. in Java (as javanicum Raychaudhuri). Antennae of apterae can be either 5- or 6-segmented; apterae with 5-segmented antennae were originally described as Eutrichosiphum elongatum (Takahashi 1940), which is now regarded as a synonym. Sexuales and life cycle are unknown.

Mollitrichosiphum rhusae A.K. Ghosh Apterae are greenish to pale brown, rather elongate-bodied; BL 2.2-2.5 mm. Siphunculi of apterae are dark brown, curved outwards, c. 0.5 × BL, and of alatae are dark brown to black, c.0.7 × BL. On undersides of leaves of unidentified Rhus sp(p). in India (Meghalaya; A.K. Ghosh 1974b). Aphids collected from Helicia hainanensis in China have also been identified as this species (D. Zhang & Qiao 2010). Sexuales and life cycle are unknown. Samples collected in China were very similar to M. luchuanum in morphology and mitochondrial DNA sequences suggesting possibly synonymy (Q. Liu et al. 2013), but a previous study using a different method of analysis on the same samples and gene sequences found a distinction between these two species (R. Zhang et al. 2011).

Mollitrichosiphum syzygii Noordam Apterae have pale brown head, thorax and marginas of abdomen, and usually a shiny brown or brownish black central abdominal patch, darker and more extensive in small specimens; BL 1.1-2.3 mm. Siphunculi are brown, darker in small specimens, 0.25-0.47 × BL in apterae and 0.48-0.63 × BL in alatae. On undersides of young red leaves and developing shoots of Syzygium antisepticum at 1400 m in Java (Noordam 1994). Sexuales and life cycle are unknown; viviparae were collected throughout the year in Java.

Mollitrichosiphum taiwanum (Takahashi) Apterae are broadly pear-shaped, yellow, antennae and legs yellow with darker apices; BL 1.7-2.0 mm. Siphunculi of apterae are yellow with black apices, 0.4-0.5 × BL; siphunculi of alatae are black, c. 0.8 × BL. On Meliosma spp. (rigida, rhoifolia) in Taiwan (Takahashi 1921, Raychaudhuri 1956). Closely related to M. yamabiwae on Meliosma in Japan, but apparently separable by the key characters given. Sexuales and life cycle are unknown.

Mollitrichosiphum tenuicorpus (Okajima) Plate 11e, f Apterae are elongate pear-shaped, pale brown with pale legs and antennae; BL 1.9-3.0 mm. Siphunculi of apterae are dark brown, 0.6-0.8 × BL. Alata are brown-black,very narrow-bodied, with black siphunculi 0.8-1.0 × BL. On young shoots of Castanopsis, Litsea and Lithocarpus spp. in India, Java, China, Taiwan, Thailand and Japan. There are also single records from Castanospermum sp. in Sikkim, India (A.K. Ghosh & Raychaudhuri 1968a), from Castanea crenata in the Philippines (BMNH collection, leg. V.J. Calilung), and from Meliosma rigida and Alnus cremastogyne in China (R. Zhang et al. 2011). There are also a few records from Quercus spp., but Quercus does not seem to be favoured as a host plant. A.K. Ghosh & Agarwala (1993) compared M. shinjii Raychaudhuri, Ghosh, Banerjee & Ghosh, which was described from an unidentified Quercus sp. in north-east India, with tenuicorpus, and concluded that they were synonyms. Alate oviparae and males are produced in northern India in late September (A.K. Ghosh & Agarwala 1993), and in Japan in autumn and winter (Takahashi & Sorin 1959). Agarwala et al. (1987) described a new species of parasitoid from this aphid.

Mollitrichosiphum tumorisiphum Qiao & Jiang Apterae are yellow green in life, with pairs of emerald green dorsal markings and pale brown siphunculi; BL 1.9-2.4 mm. Alatae are yellow green with green dorsal markings, dark brown forewing veins and dark brown siphunculi. Siphunculi are 0.3-0.4 × BL in apterae and 0.5-0.6 × BL in alatae. On undersides of young leaves of new growth of Fagus longipetiolata in Taiwan (Jiang et al. 2015). Life cycle and sexual morphs are unknown.

Mollitrichosiphum yamabiwae Suenaga Apterae are yellow brown; BL c. 2.1 mm. Alatae have dark brown dorsal abdomen and large paired ventro-lateral black marks on abdominal segments 3-6 (Takahashi 1962b). Siphunculi are dark, c. 0.6 × BL in apterae and 0.7-0.8 × BL in alatae. On Meliosma spp. in Japan (Suenaga 1934). Alate sexuales are produced in summer (Takahashi 1962b). The male genitalia were described and illustrated by Wieczorek et al. (2012). The distinction between this species and M. luchuanum, also now recorded from Meliosma in China, needs to be verified.

Monaphis Walker

Calaphidinae: Calaphidini

A genus for one large palaearctic species with many distinctive features, feeding on Betula. See Stroyan (1977) for a generic diagnosis. However, DNA barcoding has revealed that specimens from B. schmidtii in Korea there may be a second, cryptic species (Lee et al. 2017)

Monaphis antennata (Kaltenbach) (fig. 32A) All adult viviparae are alate, robust, ventrally flattened, green, with very long thick black antennae and an elongate dark pterostigma in the forewing (see influentialpoints.com/Gallery); BL 3.3-4.3 mm. They live solitarily on Betula spp. in Europe, eastward across Asia to east Siberia, China and Japan, and also (1996) recorded from Ontario, USA (Foottit et al. 2006). Immatures are cryptic, and usually press themselves close along the mid-ribs on the upper sides of leaves (Hopkins & Dixon 1997, 2000). They migrate to the undersides of the leaves for the final moult, and the adults produce their offspring at an unusually rapid rate (Hopkins 1998). Monoecious holocyclic; green apterous oviparae and red alate males occur in September-October (Ostanin 1976). 2n=20.

Monellia Oestlund

Calaphidinae: Panaphidini

Four North American species on Carya, resembling Monelliopsis but folding their more rounded wings flat over the abdomen when in repose, and probably more closely related to the Tilia-feeding genera Eucallipterus and Tiliaphis (Quednau 2003). Bissell (1978) comprehensively revised the genus, including much morphological data and keys to alate viviparae and oviparae.

Monellia caryella (Fitch) All viviparae are alate, pale lemon-yellow to greenish yellow, with banded antennae, seasonally variable; generations from midsummer to autumn bear a continuous broad black band running around front and sides of head and down sides of body as far as abdominal tergite 3, as well as a broad brown-black band along the anterior margin of the forewing (see influentialpoints.com/Gallery). BL 0.9-2.2 mm. On leaves of Carya spp., especially C. illinoiensis and C. cordiformis. Monoecious holocyclic; sexuales occur from mid-October to early December (Mansour & Harris 1988). Oviparae are dark with black dorsal abdominal cross-bands (Quednau 2003). Widespread in USA (see Bissell 1978), in Ontario, Canada, and introduced into Spain (Nieto Nafría & Mier Durante 1988), Israel (Mansour & Harris 1988), Greece (Papapanagiotou et al. 2012), Turkey (Kaya Apak & Akşit 2016) and Argentina (Ortego et al. 2004). In Israel and Turkey it is a serious pest of pecan; biocontrol has been attempted (Mansour et al. 1988), and the impact of native natural enemies has been studied in Israel (Mansour 1998, 1993)and Turkey (Kaya Apak & Akşit 2016). Alverson & English (1990) studied population dynamics, and Edelston & Estes (1987) and Watterson & Stone (1982) studied its natural enemies in South Carolina and Texas respectively. Integrated control of this and other pecan aphids in USA was reviewed by Harris (1983). According to Bissell (1978), Davidson’s (1914) account of M. caryella in California applies to a species of Monelliopsis, perhaps bisetosa, and Richards’ (1965) description and figures under the name caryella apply to Monelliopsis nigropunctata. 2n=18.

Monellia hispida Quednau All viviparae are alate, pale yellow or colourless with banded antennae, body often with 4-6 faint yellow spots internally, and later generations with a broad dusky stripe running around front and sides of body, as in M. caryella but much paler; BL 1.2-2.0 mm (Bissell 1978). Monoecious holocyclic on Carya spp. in eastern North America. Sexuales occur in late September-November.

Monellia medina Bissell All viviparae are alate, bright yellow with banded antennae, and summer/autumn generations with a heavy black band running around front and sides and along leading edge of forewing as in M,. caryella; BL c. 2.3 mm. Recorded from Carya spp. in Maryland and Ohio, USA (Bissell 1978). Very similar to M. caryella. An ovipara collected in early November is illustrated by Quednau (2003).

Monellia microsetosa Richards All viviparae are alate, pale yellow, summer generations developing 2-3 pairs of conspicuous spots of intense yellow internal pigment, especially on abdominal tergites 1-2 and 5-6 (Bissell 1978). Antennae are banded and leading edge of forewing is spotted with black. BL 1.4-2.0 mm. On leaves of Carya spp., especially C. glabra, in eastern North America from Quebec to Florida. Sexuales occur in September-November. 2n=18.

Monelliopsis Richards

Calaphidinae: Panaphidini

Ten North American species associated with Juglandaceae, mainly associated with Juglans; visits to Carya spp. by some species are possibly only “casual”. Similar to Monellia, but holding their wings vertically at rest (as do most aphids). Quednau (2000) keyed the species, and Quednau (2003) reviewed the genus and provided a key and illustrations of all species.

Monelliopsis bisetosa Richards (fig. 33F) All viviparae are alate, yellowish with mainly dark antennae, dusky brownish tibiae and tarsi, and a dark dorsal spot near apex of femur (Richards 1966); BL 1.2-1.5 mm. Described from Juglans sp., “probably rupestris”, in Mexico, and subsequently recorded ?Carya sp., from western USA on J. major (Quednau 2003) and, perhaps as a casual occurrence, on Pterocarya fraxinifolia (Walker et al. 1978). An ovipara collected in Mexico in October on ?Carya sp. is illustrated by Quednau (2003).

Monelliopsis bisselli Quednau All viviparae are alate, and very similar in life to M. nigropunctata except for the black mark on the hind subcoxa, with similar seasonal variation in pigmentation; BL 1.5-2.0 mm. On undersides of leaves of Carya spp. in eastern USA (Pennsylvania, Maryland, Georgia). Biology and sexuales have not been described.

Monelliopsis californica (Essig) Adult viviparae are all alate, yellow with dark antennae and tibiae, and with a narrow black marginal stripe on each side of prothorax and along costal margin of forewing; BL 1.7-2.0 mm. Immatures have banded antennae (Quednau 2003). Main host is Juglans californica in California (Richards 1968b) and Utah (BMNH collection); there are also records (perhaps “casual” occurrences) from Carya illinoiensis and Pterocarya spp. (Walker et al. 1978). (This species was wrongly placed in Chromaphis by Blackman & Eastop 1994).

Monelliopsis caryae (Monell) (fig. 33B) All viviparae are alate, yellow with banded antennae and, in fully pigmented specimens, a dusky median streak on the head and small dusky spots at bases of dorsal abdominal hairs (see influentialpoints.com/Gallery); BL 1.2-1.8 mm. Immatures have black patches at bases of dorsal and marginal hairs. On leaves of Juglans nigra, widely distributed in North America, and introduced in about 1984 to Europe (France, Hullé et al. 1998; Portugal, Pita & Ilharco 1998; Spain, Mier Durante & Pérez Hidalgo 2002). Oviparae were collected in Utah in October (BMNH collection, leg. G.F. Knowlton). 2n=18.

Monelliopsis nigropunctata (Granovsky) (fig. 33G,H) All viviparae are alate, yellow. Antennae ringed with black, and well-pigmented specimens in later generations have dark sides to prothorax, dark spots at bases of dorsal abdominal hairs, and a dark ventral spot near apex of hind femur (Richards 1965); BL 1.2-1.5 mm. Immatures have dark spots at bases of dorsal hairs. On leaves of Juglans spp. and Carya spp., widely distributed in North America. Most records of this species on Carya illinoiensis should probably be referred to M. pecanis (see below). Sexuales do not appear to have been recorded. 2n=10.

Monelliopsis pallida Quednau All viviparae are alate, colour in life unrecorded, with dark antennal joints but otherwise little pigmentation; BL 1.8-2.4 mm On Juglans spp., especially hindsii, in California USA. Oviparae were collected on J. hindsii in November (Quednau 2000).

Monelliopsis pecanis Bissell All viviparae are alate, yellow, with darker markings (when developed) similar to M. nigropunctata, but with fewer dark spots per segment (see influentialpoints.com/Gallery); BL 1.2-1.7 mm. On leaves of Carya illinoiensis in USA, Mexico, and introduced to Egypt (BMNH collection, leg. A.A. Attia), Sicily (Barbagallo & Suma 1999) and South Africa. Dickey & Medina (2010) record it also in Texas on C. aquatica (see below). Sexuales, and differences from M. nigropunctata, with which it was confused for many years, were described by Bissell (1983) and Quednau (2003). Alverson & English (1990) studied population dynamics, Edelson & Estes (1987) studied natural enemies in South Carolina, and de Villiers & Viljoen (1987) investigated control with systemic insecticides in South Africa. Dickey & Medina (2010, 2011) found selection for genetically divergent host-associated populations on C. illinoiensis and C. aquatica in Texas.

Monelliopsis pleurialis Richards All viviparae are alate, yellow, very similar to M. pecanis; BL c. 1.2 mm. On undersides of leaves of Carya spp. in eastern North America. The subtle differences between this species, M. nigropunctata and M. pecanis are discussed by Quednau (2003), who also illustrates an ovipara of M. pleurialis collected on C. ovata in Ontario in late August.

Monelliopsis quadrimaculata Quednau All viviparae are alate, pale (colour in life unrecorded), with black antennal joints and marginal processes on abdominal tergites 4 and 5 usually pigmented so that there are two pairs of dark marginal spots on abdomen; BL 0.9-1.6 mm. On Juglans major in Arizona, USA (Quednau 2000). Sexuales have not been described.

Monelliopsis tuberculata Richards (fig. 33D) All viviparae are alate, yellow, dark markings when developed as in M. nigropunctata; BL 1.0-1.2 mm. Described from Juglans sp., “probably rupestris, in Mexico (Richards 1966), and also recorded from J. nigra in New Mexico, Idaho and Utah (BMNH collection). Oviparae were collected in Utah in October (BMNH collection, leg. G.F. Knowlton).

Monzenia Takahashi

Hormaphidinae: Nipponaphidini

A genus for one oriental species monoecious on Distylium. Two other previously placed in Monzenia have now been transferred to Metanipponaphis, and M. minuta might slso belong in that genus (Aoki et al. 2021)..

Monzenia minuta Yeh Galls on twigs of Distylium racemosum in Taiwan are closed, spherical, rough-surfaced (Yeh & Ko 2017). Alate ?sexuparae (BL 1.0-1.2 mm) were collected in September (described as alate viviparae, but the embryos appear to lack stylets). The life cycle in unknown.

Mordvilkoiella Shaposhnikov

Aphidinae: Aphidini

Two species on Poaceae currently placed in Aphidini because of a possible relation to Hyalopterus,but with short 5-segmented antennae, siphunculi reduced to pores and no marginal abdominal tubercles. It could have a closer affinity to Holcaphis and belong in Macrosiphini.

Mordvilkoiella jacutensis Pashchenko Apterae are pale yellowish green; BL c. 2.3 mm. On Festuca pratensis in east Siberia.

Morvilkoiella skorkini (Mordvilko) Colour of apterae in life is unknown; BL unrecorded. In leaf sheaths of Phragmites australis in Russia and Ukraine (Shaposhnikov 1964).

Mordwilkoja Del Guercio

Eriosomatinae: Pemphigini

One North American species migrating from Populus to Lysimachia, with its secondary host generations very similar to those of Thecabius subgenus Parathecabius. Alatae from galls have antennae with the antennal terminal process longer than the base of the last segment and bearing “clear spots” of unknown function.

Mordwilkoja vagabunda (Walsh) Host-alternation occurs between Populus spp. of the deltoides group and Lysimachia spp. Large, irregular, multi-lobed galls are formed from the stipules of cottonwood leaves in North America (fig. 131D, and Ignoffo & Granovsky 1961b). Emigrant alatae (BL 1.8-2.4 mm) emerge from galls in May-June and found colonies on stems, leaves or roots of Lysimachia (Smith 1971). Apterae on Lysimachia are dirty yellowish white, with wax secretion; BL 2-3 mm. Alate sexuparae returning to Populus in September-November are often larger than the spring migrants (2.1-2.9 mm) and have a shorter, more normal PT. Ignoffo & Granovsky (1961a) described the gall generations and Smith (1971) gave a full account of the life cycle. Presumably anholocyclic secondary host generations have been found on Lysimachia roots in Germany (Zwölfer 1957, as Parathecabius stammeri) and Japan (BMNH collection, leg. S. Aoki). There is, however, a report of gall populations occurring on P. nigra in Turkey (Yildiz & Toper Kaygin 2010). 2n=20.

Moritziella Börner

Phylloxeridae

The two or three species on Fagaceae placed in this genus by various authors including Blackman & Eastop (1994) are now placed in Phylloxera, as there is no valid distinction from the numerous North American species in that genus (Favret et al. 2016).

Muscaphis Börner

Aphidinae: Macrosiphini

Nine species host-alternating between Pyroideae and mosses, or known only from either the primary or the secondary host. They are Myzus-like aphids perhaps closest to Nearctaphis; fundatrices are densely hairy, and give rise in the second generation to alatae which have irregular dorsal abdominal markings and the forewing media either once-branched or with a second branch close to the wing apex. The hind tibiae of some species have a row of widely-spaced Toxoptera-like ?stridulatory pegs. The host alternation has been experimentally confirmed for two species, M. mexicana (Remaudière & Muñoz Viveros 1985b) and M. escherichi (Stekolshchikov & Shaposhnikov 1993 – see below). Most attempts to transfer other species from Pyroideae to mosses have so far been unsuccessful. Primary host generations on Pyroideae are very different from the very small aphids on mosses, and were previously described as Toxopterella. Remaudière & Muñoz Vivieros (1985b) keyed the primary host morphs. Heie (1992) and Blackman (2010) reviewed the European species.The North American genus Glendenningia may be related.

Muscaphis canadensis (Hille Ris Lambers) Apterae (fundatrices) are broadly-oval, colour in life not recorded but probably dark, with dark antennae and legs and black siphunculi; BL 2.1-2.8 mm (Hille Ris Lambers 1960c). Rolling leaves of unidentified Crataegus sp(p). in eastern North America (recorded from New Brunswick and Ontario, Canada, and North Carolina, USA). The second generation are alatae (BL 1.6-2.0 mm), and migrate to an unknown secondary host (probably a moss).

Muscaphis cuspidata (Stroyan) Apterae are dark greenish brown with brown antennae and legs and shiny black dorsum and siphunculi, ; BL 0.9-1.3 mm. Alatae have protruberant transversely oval secondary rhinaria distributed ANT III 15-24, IV 4-13, V 4-7. On mosses (Calliergonella cuspidata, Drepanocladus aduncus, Brachythecium rivulare) close to or below water-level. In Europe (England, Germany, Czech Republic), This aphid can live submerged, apparently because the papillate sculpturing of the cuticle is able to trap a layer of air around the body (original description, and Müller 1975b, as Aspidaphium cuspidata). Anholocyclic, with a specialised overwintering fourth instar (Müller 1973b). However, Albrecht (2015) pictures an ovipara identified as this species on Brachythecium rivulare.

Muscaphis escherichi (Börner) Plate 16i (Fig.44h) Apterae (fundatrices) are plump-bodied, matt dark brown to purplish black with rather shiny black siphunculi; BL 2.7-4.4 mm. They are found in spring in curled leaflets of Sorbus spp., usually with one fundatrix and its progeny in each leaflet (MacGillivray & Bradley 1961, as Toxopterella drepanosiphoides). The second generation are almost all alatae (BL 1.7-2.7 mm), and leave Sorbus from late June to early August in northern Russia. Alatae have markedly protruberant transversely oval secondary rhinaria distributed ANT III 15-20, IV 7-11, V 5-6. Stekolshchikov & Shaposhnikov (1993) succeeded in transferring some migrants from S. aucuparia to the moss Plagiothecium laetum. Only two of the larvae deposited by migrants reached maturity, but the very small adult apterae were morphologically indistinguishable from those of European populations of M. escherichi that live throughout the year on mosses, thus appearing to establish synonymy with the Sorbus-feeding aphid previously known as M. drepanosiphoides. However, M. escherichi is recorded in Europe from many species of moss, yet all attempts at transfer to species other than Plagiothecium have been unsuccessful, and gynoparae and males have never been collected from mosses. Molecular work is needed to determine whether the anholocyclic moss-feeding population in Europe is isolated genetically from the Sorbus-feeding generations, in which case it should perhaps continue to be regarded as a separate species.

M. escherichi is of holarctic distribution; across Canada (British Columbia, New Brunswick), UK, Finland, Norway, Estonia, Lithuania, Russia (St Petersburg), Czech Republic, and North Korea (Lee & Havelka 2001b). The sexual phase on Sorbus occurs in northern continental Europe, Czech Republic, Canada and North Korea. Gynoparae and males appear on Sorbus in late August in northern Russia, and mature oviparae are present in September-October. Differences have been noted between the spring forms in Canada, north-east Europe and Korea (Shaposhnikov 1963, Lee & Havelka 2001b), and Shaposhnikov gave the Canadian and north-east European populations sub-species status (as ssp. irae). 2n=12 (for “ssp. irae”).

Muscaphis japonica Sorin Only the alatae are known, and are blackish brown, BL c.1.5 mm. On Sorbus alnifoliae in Japan (Sorin 2011). In the key to aphids on Sorbus it will possibly key to M. escherichi, but the alatae are without dark dorsal abdominal markings..

Muscaphis mexicana Remaudière & Muñoz Viveros Apterae (fundatrices) are plump-bodied, velvety dark brown to reddish brown with black head and siphunculi; BL 1.6-2.4 mm. Their immature progeny are light brown. On Crataegus pubescens (=mexicana) in Mexico, distorting and reddening young leaves. It is also recorded from Argentina (Ortego et al. 2004). Heteroecious holocyclic; the second generation are alatae (BL 1.7-2.1 mm) and migrate to unidentified mosses. Apterae on mosses are yellow or yellowish green, often dark anteriorly, with reddish patches at bases of siphunculi, or (in older adults) almost black; BL 0.6-0.8 mm. Gynoparae and males fly back to Crataegus in November-December; Remaudière & Muñoz Viveros (1985b) described the life cycle, and recorded heavy predation of spring colonies on Crataegus.

Muscaphis musci Börner (Fig.44a,b,j) Apterae are greyish yellow, pale brown, or shiny dark olive-green, with reddish brown siphunculi; BL 0.5-1.0 mm. Alatae are larger (BL 1.1-1.5 mm), and have protruding, transversely oval secondary rhinaria distributed ANT III 18-24, IV 10-16, V 6-11. On mosses in the genera Acrocladium, Amblystegia, Calliergon (Amblystegiaceae); Barbula, Tortula (Pottiaceae); Brachythecium, Eurhynchium, Pseudoscleropodium (Brachytheciaceae); Bryum (Bryaceae); Catharinaea, Polytrichum (Polytrichaceae); Hylocomium (Hylocomiaceae); and Mnium (Mniaceae). Europe (UK, Belgium, Denmark, Germany, Poland, Switzerland, Czech Republic) and North America (New Brunswick, North Carolina, Idaho). The life cycle is uncertain; alate males have been collected in Germany (BMNH collection) and Poland (Wilkaniec & Borowiak-Sobkowiak 2009), and it is possibly heteroecious holocyclic with an unknown primary host, but collection of active stages in autumn/winter (Müller 1973b, as M. stammeri; Tinguely 1993) indicates that it is at least partially anholocyclic.

Muscaphis smithi (Hille Ris Lambers) Apterae (fundatrices) are very plump-bodied, dark brownish black with a reddish tinge, with jet black siphunculi; BL 3.0-4.0 mm. In tightly curled and reddened leaves of Malus angustifolia in North Carolina, USA, attended by ants. Immature progeny are reddish, all developing into reddish emigrant alatae (BL 1.7-2.1 mm), which fly away in July to colonise an unknown secondary host – probably a moss.

Muscaphis stroyani (Smith) Apterae (fundatrices) are black; BL c. 2.5 mm. On Crataegus punctata in North Carolina, USA (Smith 1980), and also recorded from two Crataegus spp. in North Korea (Lee & Havelka 2001b). Dark reddish alatae (BL 1.6-2.0 mm) are produced in the second generation, in June-July. The life cycle is unknown; probably host-alternating to mosses.

Muscaphis utahensis (Smith & Knowlton) (Fig.44i) Apterae are dark brown; BL 0.7-1.1 mm. On mosses in western USA (Idaho, Utah, Washington); the only recorded host is Cratoneurum filicinum.

Myzakkaia A.N. Basu

Aphidinae: Macrosiphini

Four nominal species in India and east Asia, three described from Polygonum, closely related to Trichosiphonaphis, but with a dark sclerotic tergum, black hairless siphunculi with a well-developed flange, and cauda with a distal constriction. The species are very similar and their distinctness needs to be confirmed by further work. The species described as Vesiculaphis polygonii by Bhattacharya & Dey (2001) probably also belongs in this genus.

Myzakkaia kuwanais (A.K. Ghosh, Basu & Raychaudhuri) Colour in life unrecorded, probably black with antennae and legs mainly pale; BL c.2.2-2.4 mm. Described (in Vesiculaphis) from an unknown host in Assam and subsequently recorded from Sikkim on Lyonia ovalifolia (A.K. Ghosh 1977); this requires confirmation, as it is likely to be the host of a Vesiculaphis rather than a Myzakkaia.

Myzakkaia niitakensis (Takahashi) Apterae are black, BL c.1.5-1.6 mm. On Polygonum sp. in Taiwan.

Myzakkaia polygonicola A.N.Basu (Fig.42b,c) Apterae are probably black, BL 1.6-1.9 mm. On Polygonum runcinatum and Polygonum sp. in West Bengal, India.

Myzakkaia verbasci (Chowdhuri, R.C. Basu, Chakrabarti & Raychaudhuri) Plate 20e Apterae are brownish black to black, a little shiny; BL 1.3-1.6 mm. Alate have secondary rhinaria distributed III 19-31, IV 0-4, V 0 (BMNH collection, leg. A.N. Chowdhuri). On Polygonum alatum and Polygonum sp(p), feeding on young shoots and under leaves, which are rolled inward (A.N. Basu 1969, as M. himalayensis). India (West Bengal, Kashmir, Himachal Pradesh), and alatae possibly of this species have been trapped in the Philippines (BMNH collection, leg. V. Calilung). The original description included specimens collected on Verbascum thapsus, but this is unlikely to be a true host. 2n=12.

Myzaphis van der Goot

Aphidinae: Macrosiphini

Eight species on Rosaceae (Rosa, Potentilla). Apterae are rather elongate oval, and dorsoventrally-flattened, with short appendages, and dorsal cuticle characteristically sculptured with numerous small, rounded depressions. They have 5 hairs on first tarsal segments, and alatae have a dark dorsal abdominal patch. The genus was comprehensively reviewed by Kanturski et al. (2018b) including scanning electron microscope studies, especially of sensillae. They transferred one species (avariolosa) to Ericaphis and erected a new genus (Richardsaphis) for another (canadensis).

Myzaphis bucktoni Jacob Apterae are pale yellow to pale green with brown dorsal markings consisting of a brown head, two large brown patches on pronotum, and paired broad stripes extending from mesothorax to base of cauda, converging between siphunculi (see influential points/Gallery); BL 1.0-1.9 mm. Alatae have a pale brown central dorsal abdominal patch, and secondary rhinaria distributed ANT III 10-14, IV 0-3. On Rosa spp., usually dispersed along mid-ribs on upper sides of leaves. In Europe, Turkey, Kazakhstan (Kadyrbekov & Aoitzhanova 2005), Afghanistan (Kanturski et al. 2018b) Pakistan (Naumann-Etienne & Remaudière 1995), Mongolia (Holman & Szelegiewicz 1972), North Africa (Algeria, Morocco), and introduced to USA and Argentina (Ortego 1998a). Monoecious holocyclic in UK, with oviparae and small dark apterous males in November (original description). All morphs were redescribed by Kanturski et al. (2018b). 2n=13 (one sample from Portugal).

Myzaphis juchnevitschae Kadyrbekov Apterae are dark greyish brown; BL 1.4-1.8 mm. On Rosa alberti in south-east Kazakhstan. Alatae have 10-11 secondary rhinaria on ANT III. Kanturski et al (2018b) provided a redescription of the type material.

Myzaphis oezdemirae Kanturski & Barjadze Colour of apterae in life unknown; BL c. 1.1 mm. Alatae are undescribed. On an unidentified Rosa sp. in Turkey and Iran (Kanturski et al. 2018b). Monoecious holocyclic, with oviparae and alate males collected in October-November; these were originally identified and briefly described by Tuatay & Remaudière (1964) as the sexual morphs of M. turanica.

Myzaphis rezwanii Kanturski & Barjadze Colour of apterae in life unknown; BL 1.1-1.6 mm. Alatae have 8-11 secondary rhinaria on ANT III. On unidentified wild Rosa sp(p). in montane localities in Iran (Kanturski et al. 2018b). Monoecious holocyclic; oviparae and apterous males were collected in October.

Myzaphis rosarum (Kaltenbach) Plate 15g Apterae are yellow-green to green (see influential points/Gallery); BL 1.2-2.4 mm. Alatae have 14-32 secondary rhinaria on ANT III. On wild and cultivated Rosa (especially rambler roses) and frequently also on shrubby Potentilla spp. They feed mainly along mid-ribs on both upper and undersides of young leaves. Occasionally found on Fragaria spp. In Europe, and now almost cosmopolitan (Asia, Morocco, South Africa, New Zealand, North and South America). Monoecious holocyclic on Rosaceae in Europe, with oviparae and small dark apterous males in November. Kanturski et al. (2018b) provided a redescription of all morphs. Anholocycly occurs in New Zealand, and probably elsewhere. 2n=4.

Myzaphis tianshanica Kadyrbekov Apterae are yellow-green; BL 1.4-2.0 mm. On Rosa albertii in south-east Kazakhstan. Kanturski et al. (2018b) provided a redescription of fundatrices, apterae and alatae, the latter having 3-18 secondary rhinaria on ANT III.

Myzaphis tuatayae Kanturski & Barjadze Colour of apterae in life unknown; BL 2.1-2.3 mm. On an unidentified Rosa sp. in Turkey (Kanturski et al. 2018b). Alatae and sexual morphs are unknown.

Myzaphis turanica Nevsky Apterae are green; BL 1.4-1.7 mm. Alatae have a paler and more irregularly-shaped dorsal abdominal patch than M. rosarum, and 6-14 secondary rhinaria on ANT III. Described from Rosa bruniana in Central Asia (Nevsky 1929a, as ssp. of M. rosarum), and subsequently recorded from wild and cultivated Rosa sp(p). in Sweden, Israel, Italy, Portugal, Turkey, Central Asia, India, Mongolia and Brazil. Monoecious holocyclic with apterous males; the alate males briefly described from Turkey by Tuatay & Remaudière (1964) were assigned to M. oezdemirae by Kanturski et al. (2018b), who redescribed all morphs. The ovipara was described from India by L.K. Ghosh (1986).

Myzocallis Passerini

Calaphidinae: Myzocallidini

About 40 species of often small, delicate, usually yellowish aphids with a knobbed cauda and bilobed anal plate, mostly monoecious holocyclic on Fagaceae. Immatures usually have long, capitate dorsal hairs. Adult viviparae are all alate, except in a few species. Forewings are variably pigmented, but there is generally at least a dark spot at the base of the pterostigma. Quednau & Remaudière (1994a) revised the subgeneric classification, Quednau (1999) revised, keyed and illustrated the world fauna, and the main regional accounts are by Stroyan (1977, Britain); Heie (1982, Fennoscandia and Denmark); Nieto Nafría et al. (1998, Iberian peninsula); Higuchi (1972, Japan); W. Zhang & Zhang (1994, China, but note nomenclatural changes in Quednau 1999); Richards (1966, Canada); Boudreaux & Tissot (1962, the subgenus Lineomyzocallis on North American red oaks), and Quednau & Remaudière (1987, the subgenus Castaneomyzocallis on North American chestnuts). Quednau (1999) transferred the subgenus Nippocallis to Tuberculatus.

Myzocallis (Californicallis) agrifolicola Richards Alatae have black head and thorax, abdomen either reddish brown or yellowish with a pattern of paired brown or black markings, antennae and legs mainly pale, and forewings with a pattern of broad fuscous bands which only partly follow the veins; BL 2.0-2.3 mm. Immatures are yellowish with dark spots at bases of dorsal and marginal hairs. On undersides of leaves of Quercus agrifolia and Q. wislizeni in California, USA (Richards 1966, 1968f; Hille Ris Lambers 1966a). Sexuales are undescribed, and biology is unstudied.

Myzocallis (Neomyzocallis) asclepiadis (Monell in Riley & Monell) Alatae have distinctive black dorsal markings and forewings with an irregular pattern of dark pigmentation (see influentialpoints.com/Gallery). On undersides of leaves of Asclepias spp.in eastern USA and Canada. Oviparae occur in October (Hottes & Frison 1931, Quednau 1999).

Myzocallis (Lineomyzocallis) atropunctata Quednau Colour in life is not recorded, probably as other Lineomyzocallis, but with three pairs of black marginal spots on abdomen; BL 1.8-2.3 mm. Immatures have a series of dark spinal and marginal sclerites. On Quercus spp. in western USA (California, Arizona) and Mexico (Quednau 1997).

Myzocallis (Lineomyzocallis) bella (Walsh) Alatae are yellow with broad black bands down sides of thorax to wing insertions, and continuing along leading edges of wings; antennae ringed with black, and legs black except for bases of femora, trochanters and coxae. BL 2.5-3.0 mm. Immatures yellow, with legs yellow except for black tarsi, and dorsum distinctively marked with 5 pairs of black patches forming an oval shape. On various Quercus spp. throughout eastern North America. Monoecious holocyclic; sexuales occur in October-December (Boudreaux & Tissot 1962).

Myzocallis boerneri Stroyan Alatae are yellowish, with head and thorax sometimes partly dusky, dorsal abdomen with small paired spinal and marginal specks of brown pigment (visible with a hand-lens), and antennae ringed with brown-black (see influentialpoints.com/Gallery); BL 1.3-2.2 mm. Immatures are yellow with paired, dusky, rather indistinct spinal and marginal spots. On undersides of leaves of Quercus spp. (cerris, ilex, infectoria, petraea, suber, variabilis) widely distributed in Europe, the Middle East (Israel, Iran, Lebanon), and introduced on European oaks to New Zealand (Rohitha 1982), California and most recently, Argentina (Ortego et al. 2006). However the high level of genetic divergence found between specimens from Italy and New Zealand indicates (assuming correct identification) that they are not the same species (Lee et al. 2017). Monoecious holocyclic; sexuales occur in October-November in the northern hemisphere. Very similar to, and often confused with, M. komareki or M. schreiberi (see Hille Ris Lambers & Stroyan 1959; López & Nieto 1983). Population dynamics was studied by Kindlmann & Dixon (1998). 2n=14*.

Myzocallis carpini Koch Alatae are pale yellow to yellowish white, without dorsal abdominal markings. The antennae are ringed with black, and the forewing has a black spot at base of pterostigma (see influentialpoints.com/Gallery); BL 1.3-2.2 mm. On undersides of leaves of Carpinus betulus, especially when used for hedging. In Europe, Mediterranean area, Middle East, and introduced into New Zealand and North America (Oregon, British Columbia). There is also a record from north-east China (Qiao et al. 2005b). Monoecious holocyclic; oviparae collected in early November in Britain, and in late March in New Zealand (BMNH collection, leg. A.D. Lowe). The male genitalia were briefly described by Wieczorek & Świątek (2007). Bihar (1963) studied honeydew production. 2n=14*.

Myzocallis (Castaneomyzocallis) castaneae (Fitch) Alatae are yellow with black ANT III-VI, tibiae and tarsi. The forewing has the pterostigma with a black spot at its base and a line of pigment following its posterior margin, and vein Cu_1b and base of Cu_1a are black-bordered. BL 2.1-2.6 mm. On undersides of leaves of Castanea dentata and Castanea sp(p). in North America (Quednau & Remaudière 1987). Monoecious holocyclic; sexuales occur in October in California (Essig 1917).

Myzocallis (Castaneomyzocallis) castaneoides (Baker) Alatae are yellow, with ANT III-VI and tibiae entirely black or with paler sections, and forewings as in castaneae; BL 1.5-1.9 mm. On Castanea spp. in eastern USA and Mexico. Sexuales are undescribed; Tissot’s (1932a) description of sexual morphs under this name should be applied to M. tissoti (Quednau & Remaudière 1987).

Myzocallis (Agrioaphis) castanicola Baker (fig. 35E) Alatae are yellow, distinctively marked with a dark median stripe on head and thorax and paired black spinal and marginal patches on dorsal abdomen (see influentialpoints.com/Gallery); BL 1.6-2.3 mm. Antennae dark beyond the basal half of ANT III, tibial apices and tarsi dark, forewing veins rather dark ending in fuscous spots, and siphunculi dark. Immatures are yellowish with brown-black dorsal markings. On undersides of leaves of numerous Castanea spp. and Quercus spp. In Europe, Mediterranean region, Middle East, southern Africa, St Helena, Australia, New Zealand, South America (Brazil, Chile, Argentina) and western North America (California, British Columbia). Monoecious holocyclic; sexuales occur on both Castanea and Quercus in October in Britain, and in May in Australia. Cottier (1953) gave a detailed account of this aphid in New Zealand, including descriptions of sexual morphs. Quednau & Remaudière (1994) distinguished a subspecies, leclanti, specific to Castanea sativa in southern Europe (France, Andorra, Spain, Corsica, Greece, Turkey). 2n=14.

Myzocallis (Pasekia) cocciferina Quednau & Barbagallo Alatae are pale yellow, with head and thorax pale brownish laterally, abdomen with rather large, pale brown, paired dorsal patches, antennae and legs mainly pale, and siphunculi pale; BL 1.4-2.4 mm. Immatures are pale yellowish with brown dorsal markings. On undersides of leaves of Quercus calliprinos, Q. coccifera and Q. ilex in the Mediterranean region (Israel, Lebanon, Sicily, France, Andorra, Spain). Sexuales are unknown; it seems to be anholocyclic, at least in Sicily (Quednau & Barbagallo 1991). 2n=14*.

Myzocallis coryli (Goetze) Alatae are pale yellow to yellowish white, antennae ringed with black, and forewing with a black spot at the base of the pterostigma (see influentialpoints.com/Gallery); BL 1.3-2.2 mm. On undersides of leaves of Corylus spp. In Europe, south-west and central Asia, Pakistan (Naumann-Etienne & Remaudière 1995), Africa (Atlas Mountains; Heie 1982), Japan (Shinji 1941), Australia, New Zealand, western North America, and South America (Argentina, Chile). However DNA barcoding has indicated that there may be two cryptic species confused under this name, one of which forms a sister group with M. carpini (Lee et al. 2017). If confirmed this could have repercussions on the interpretation of past research on this well-known species. Monoecious holocyclic; sexuales in October-November in western Europe and in April-June in New Zealand (Cottier 1953). The male genitalia were briefly described by Wieczorek & Świątek (2007). Messing & Aliniazee (1991) studied post-diapause egg development. M. coryli has developed resistance to carbamate insecticides in North America (Aliniazee 1983), and biocontrol was attempted with a host-specific race of Trioxys pallidus (Messing & Aliniazee 1988). Aruta & Carvillo (1989) reported a fungal attack in Chile. 2n=14.

Myzocallis (Neomyzocallis) discolor (Monell) Alatae are yellow, with yellow-brown head and prothorax, and variably-developed brown dorsal abdomen markings. The antennae are ringed with brown, the forewing membrane usually has a pattern of infuscation, but is sometimes almost hyaline, and the siphunculi are always dark. BL 1.5-2.0 mm. Immatures yellow, pink or brownish with darker dorsal abdominal spots (Richards 1968f). Living singly on undersides of leaves of Quercus spp. (mostly on white oaks), widely distributed in North America, and in Cuba (Holman 1974) and Costa Rica (Villalobos Muller et al. 2010). Monoecious holocyclic; sexuales occur in September-October.

Myzocallis (Lineomyzocallis) durangoensis Remaudière & Quednau Appearance in life not observed but probably yellow with dark pigmentation as in M. granovskyi; BL 1.7-2.5 mm. Immatures have paired dark spinal and marginal sclerites but pale tibiae. On Quercus durifolia and Quercus sp(p), in Mexico (Remaudière & Quednau 1992). An ovipara collected on Q. emoryi in November is illustrated by Quednau (1999).

Myzocallis (Lineomyzocallis) elliotti Boudreaux & Tissot Alatae are yellow, distinctively marked with black, and with black legs, as in M. bella; BL not recorded, and not measurable on mounted specimens, but said to be the smallest of the dark-legged species of Lineomyzocallis (Boudreaux & Tissot 1962). Immatures have paired, quadrate black patches on abdominal tergites 1-5, similar to those of bella. On Quercus schumardii and Quercus sp. in Louisiana, USA. Monoecious holocyclic; sexuales occur in November-December.

Myzocallis (Lineomyzocallis) ephemerata Richards Alatae are yellow with black markings very like M. walshii; BL c. 2 mm. Immatures are yellow, with dark tarsi and antennae ringed with black. On Quercus rubra and Q. shumardii in eastern North America (Quednau 1999). Sexuales and life cycle are unstudied.

Myzocallis (Lineomyzocallis) exultans Boudreaux & TissotAlatae are yellowish, with black spot behind eye, broad black bands down sides of thorax and along leading edges of wings, antennae ringed with black, fore tibiae black, mid and hind tibiae dusky; BL c. 1.5 mm. Immatures are yellow, dusted with a greyish wax powder. On leaves of Quercus spp. (red or pin oaks) in eastern USA (Boudreaux & Tissot 1962, Richards 1968f), and also found in Oregon and Idaho (aphidtrek.org). Monoecious holocyclic; sexuales occur in September-October (in Pennsylvania).

Myzocallis (Lineomyzocallis) frisoni Boudreaux & Tissot Alatae are yellow, marked with black as in M. bella, except that the antennae are entirely black beyond the basal part of ANT III; BL c. 1.9 mm. Brachypterous adults are sometimes present, and may have paired dark patches on abdominal tergites 3-5. Immatures have black antennae, pale legs and dark dorsal abdominal patches. On Quercus spp. (palustris, texana) in eastern USA. Monoecious holocyclic; sexuales occur in October (Boudreaux & Tissot 1962).

Myzocallis glandulosa Hille Ris Lambers Apterae are yellow, with variably developed, paired spinal dark patches or cross-bars, and smaller marginal patches, from pronotum to abdominal tergite 8; BL 0.65-1.30 mm (Hille Ris Lambers 1948). Alatae rather squat-bodied, with banded antennae having 2-5 secondary rhinaria on thicker basal part of ANT III, dark lateral spots on the prothorax, and variably pigmented spinal and marginal abdominal sclerites; BL c. 1.3 mm. The alata has not been properly described, but is illustrated by Quednau (1999); Richards’ (1968f) description under this name applies to M. komareki. On undersides of leaves of Quercus ithaburensis in Israel, and also recorded from Q. persica in Iran and Quercus sp. in Turkey (Quednau 1999). The apterae are consistently the commonest morph (BMNH collection). Canakçioglu (1975) records this species from Q. cerris and Q. aegilops in Turkey, but does not specify which morph(s) were collected, and perhaps had boerneri or komareki. The life cycle requires clarification; sexuales occur in late autumn and winter (Bodenheimer & Swirski 1957), and fundatrix-like alatae in April (BMNH collection, leg. B. Halperin). 2n=14*.

Myzocallis (Lineomyzocallis) granovskyi Boudreaux & Tissot Alatae are yellow with very broad black bands of dark pigment running from posterior margins of eyes along sides of thorax and leading edges of wings (see influentialpoints.com/Gallery). Antennae are ringed with black, femora black on distal third, tibiae and tarsi black. BL 1.8-2.3 mm. Immatures have a grey waxy bloom, dark dorsal patches and dark tibiae. On numerous Quercus spp., usually on upper sides of leaves (Boudreaux & Tissot 1962). Throughout eastern USA, in Ontario, Canada Richards 1965), and also collected once in Oregon (BMNH collection, leg. D. Hille Ris Lambers). Monoecious holocyclic; sexuales occur in October.

Myzocallis (Pasekia) komareki (Pasek) Alatae are yellow, with head and thorax variably marked with black (typically there are fine, black lines on the prothorax, roughly in the shape of a pair of rectangles), and dorsal abdomen with a series of large, paired quadrate spinal sclerites, which have dark margins and paler central areas, and similar-sized but paler and more uniformly-pigmented marginal sclerites. BL 1.1-2.2 mm. On Quercus spp. (e.g. aegilops, cerris, coccifera, ilex, ithaburensis, petraea, pyrenaica, suber) in southern Europe, Mediterranean area and Middle East (Bulgaria, Czech Republic, Greece, Israel, Iran, Italy, Moldova, Turkey, Ukraine, Canary Islands, Andorra, Spain). Monoecious holocyclic; sexuales occur in November in Iran (BMNH collection, leg. S.H. Hodjat). 2n=14*.

Myzocallis (Lineomyzocallis) longirostris Richards Alatae are yellow, antennae ringed with black, pigmentation of body and wings variable; fully-pigmented specimens have black sides to thorax, dark fore tibiae and dark bands along leading edges of wings, but some specimens are rather pale (Richards 1965). BL 1.4-1.8 mm. Immatures are bright yellow with antennae ringed with black. On red oaks in eastern North America, and in Mexico (BMNH collection, leg. G. Remaudière). Quednau (1999) illustrated an ovipara collected from Q. ilicifolia in Pennsylvania in early October. In Mexico, a subspecies (tepehuanensis; Remaudière & Quednau 1992) occurs on Q. urbani var. parvifolia.

Myzocallis (Lineomyzocallis) longiunguis Boudreaux & Tissot Alatae are yellow, distinctively marked with black as in M. bella; BL 1.5-2.0 mm. Immatures are very pale, almost translucent yellow, with antennae ringed with black. On Quercus spp. (mostly red or pin oaks) in eastern North America (Boudreaux & Tissot 1962, Richards 1968f). Quednau (1999) illustrated an ovipara collected on Q. cinerea in Florida in December.

Myzocallis (Pasekia) mediterranea Quednau & Remaudière Apterae are pale yellow to creamy white, with dark markings; BL 1.2-1.6 mm. On Quercus pubescens and Q. ilex in southern France and Spain (Quednau & Remaudière 1994a), and on Q. pubescens and related species in Italy (Sicily, Patti & Barbagallo 1997; Tuscany, Barbagallo et al. 2011). Specimens recorded as M. komareki from Q. pubescens and Q. canariensis in Sicily (Barbagallo & Stroyan 1982) are also referable to this species. A record from Turkey (Şenol et al. 2017) requires additional confirmation. Sexuales occur along with alate viviparae from November to January, their appearance probably depending on the timing of leaf senescence (Barbagallo & Stroyan 1982, as M. komareki).

Myzocallis (Lineomyzocallis) melanocera Boudreaux & Tissot Alatae are yellow, distinctively marked with black as in M. bella, and with almost wholly black antennae; BL 2.4-2.9 mm. Immatures are yellow without any wax bloom, and have pale legs and dark spinal, pleural and marginal sclerites (Boudreaux & Tissot 1962, Richards 1965). Quednau (1997) provided a redescription. On Quercus spp. (nigra, lyrata, laurifolia) in southern USA (Louisiana, Florida). Monoecious holocyclic; sexuales occur on Q.nigra in December. Aphids with paler antennae previously regarded as a northern form of M. melanocera were distinguished as a separate species (neoborealis)by Quednau (1997).

Myzocallis (Lineomyzocallis) meridionalis Granovsky Both apterous and alate morphs occur. Alatae are yellow marked with black as in M. bella; BL 1.7-1.9 mm. Apterae are yellow with black antennae, dark tibiae, and black sclerites on abdominal tergites 3-5 coalesced spinally to form a 6-lobed central patch; BL 1.7-2.1 mm. On undersides of leaves of Quercus spp. (especially nigra, schumardii) in south-eastern USA and Panama. The life cycle is unclear; oviparae occur in December (with a black central dorsal patch like that of apterous viviparae), but parthenogenetic populations persist through winter on trees which keep their leaves in the southernmost states (Boudreaux & Tissot 1962).

Myzocallis (Lineomyzocallis) multisetis Boudreaux & Tissot Alatae are pale yellow, distinctively marked with black as in M. bella, except that the mid and hind tibiae are pale with black apices; BL 1.9-2.3 mm. Immatures are very pale yellow, with black pigment usually only at apices of antennal segments. On Quercus spp. (mostly of red oak group, but not on Q. nigra), widely distributed in eastern North America. Monoecious holocyclic, with oviparae collected on Q. palustris in Pennsylvania in late November (Quednau 1999), but populations may persist into winter on trees retaining leaves in southern states (Boudreaux & Tissot 1962).

Myzocallis (Agrioaphis) myricae (Kaltenbach) Alatae are yellow or orange, with black longitudinal markings on head and thorax, and paired black spinal and marginal sclerites on abdominal tergites 1-7 (see influentialpoints.com/Gallery); BL 1.8-2.0 mm. Brachypterous specimens are common. On undersides of leaves and young growth of Myrica gale in northern and western Europe (Heie 1982), and recorded from one locality in Spain (Nieto Nafría &Mier Durante 1998). Monoecious holocyclic; sexuales occur in October. 2n=14.

Myzocallis (Castaneomyzocallis) nanae (Tissot) Alatae are yellow or greenish yellow with dark or banded antennae and tibiae, light brown pterothorax, forewings as in castaneae, and paired dusky spinal and marginal abdominal patches; BL 1.4-2.0 mm. On undersides of leaves of Castanea nana in Florida, USA. Sexuales and life cycle are unknown.

Myzocallis (Lineomyzocallis) neoborealis Quednau Appearance of alatae in life unknown, probably like M. melanocera but with banded antennae; BL 1.6-2.9 mm. Immatures are pale in spring but have dark spinal, pleural and marginal sclerites in later generations. On Quercus spp. (rubra, palustris, coccinea) in eastern USA and Canada. Sexual morphs occur on Q. rubra in September-October (Quednau 1997).

Myzocallis occidentalis Remaudière & Nieto Nafria Alatae are pale lemon yellow with dark sides to prothorax, and small dusky spinal and rather larger marginal sclerites on abdominal tergites 1-7; BL 1.5-2.2 mm. On Quercus pyrenaica, in south-west France, Spain and Portugal (Remaudière & Nieto Nafria 1974), and on Quercus sp. in Serbia (Vučetić et al. 2014) and Greece (Tsitsipis et al. 2007); it is recorded from Italy on Q. macrolepis (Nieto Nafría & Mier Durante 1998) and there is also one specimen in the BNMH collection from Q. pubescens (leg. G. Remaudière). Monoecious holocyclic; Nieto Nafría & Mier Durante (1998) describe oviparae and males collected in Spain in late September-October.

Myzocallis (Globulicaudaphis) pakistanica (Hille Ris Lambers) Apterae are “basically pale greenish marked with a darker olive green dorsal pattern, and with what appear to be white paired waxy markings running laterally from the head about two-thirds of length of body. Legs pale with black knees, antennae pale, annulated with dark markings at joints” (Hille Ris Lambers 1966); BL 1.4-1.8 mm. Alatae have 2-3 rather large rounded secondary rhinaria near base of ANT III. In colonies on undersides of leaves of Quercus dilatata in Pakistan (Murree hills and Kalam valley; Naumann-Etienne & Remaudière 1995) and India (Himachal Pradesh; BMNH collection, leg. S.K. David). Biology and life cycle are largely unknown; apterous viviparae have been collected in June, November and December (BMNH collection). In the Murree hills it is common especially in autumn and early winter between 1800m and 3000m (Naumann-Etienne & Remaudière 1995).

Myzocallis (Lineomyzocallis) pepperi Boudreaux & Tissot Alatae are yellow with banded antennae and the distinctive black markings of the subgenus; BL 1.5-1.6 mm. Immatures are pale yellow without any dorsal abdominal pigmentation. On Quercus laurifolia in Florida, USA, and on Quercus sapotifolia in Costa Rica (Villalobos Muller et al. 2010). Populations on Q. castanea and Q. crassipes in Mexico were described as a subspecies (M. pepperi iturbide; Remaudière & Quednau 1992). Sexuales and life cycle are unknown.

Myzocallis (Pasekia) persica Quednau & Remaudière Colour of alata in life is unknown; BL 1.6-2.3 mm. On Quercus persicae in Iran and Turkey (Quednau & Remaudière 1994a). Sexual morphs occur in October.

Myzocallis (Neodryomyzus) polychaeta David Colour of apterae in life is unrecorded, probably yellow; antennae banded, legs pale, dorsal markings variable but when fully developed they are quite characteristic, comprising paired spinal and marginal quadrate patches with the numerous pleural hairs in between borne on separate small rounded scleroites. BL 1.3-1.8 mm. Alate viviparae are recorded (Chakrabarti 1988), but undescribed. On Quercus spp. (dealbata, semicarpifolia) in India (Himachal Pradesh, Uttar Pradesh), Pakistan (one trapped alata; Naumann-Etienne & Remaudière 1995) and Nepal (BMNH collection). Monoecious holocyclic; oviparae and males occur in October (Chakrabarti & Raychaudhuri 1974). 2n=12 (according to Khuda-Bukhsh & Pal 1983b).

Myzocallis (Neomyzocallis) pseudodiscolor Quednau Alatae are of unknown colour, with antennae ringed with black, legs mainly pale, forewings with a variable pattern of infuscation along and between veins, abdomen with paired dark spinal and marginal sclerites often fused on abdominal tergites 4 and 5 into a large irregularly shaped patch; BL 1.6-2.5 mm. On Quercus macrocarpa in north and central USA and Canada (Manitoba). Oviparae and males occur in September-October (Quednau 1996).

Myzocallis (Neomyzocallis) punctata (Monell) Alatae are yellow, antennae ringed with black, legs mainly pale, dark dorsal body markings often present but very variable (see Quednau 1999 and influentialpoints.com/Gallery). Forewings very variably pigmented, sometimes almost hyaline in early spring populations except for dark spots at ends of veins, but later with a distinctive pattern of patchy infuscation; BL 1.8-2.5 mm. Immatures have a variably developed pattern of dark spinal and marginal sclerites. On numerous Quercus spp., especially of the white oak group in eastern North America – where it is sometimes very common in trap catches (Schultz et al. 1985) – and introduced into western states. In California it has been collected on many non-native oaks, and also on Lithocarpus densiflora (BMNH collection, leg. H.G. Walker). Monoecious holocyclic; sexuales occur in September-October in north-east USA, but in California viviparae are present through the winter. 2n=14.

[It has been pointed out (Lee et al. 2017) that some Genbank sequences of M. punctata may be misidentified M. asclepiadis.]

Myzocallis schreiberi Hille Ris Lambers & Stroyan Alatae are pale yellow, with head and thorax slightly darker; the prothorax often has short lateral streaks of dark pigment, the dorsal abdomen has small transversely oval, often dusky to dark, spinal spots, and paler marginal sclerites, the legs are mainly pale and the siphunculi usually dark (see influentialpoints.com/Gallery); BL 1.3-2.2 mm. Immatures are whitish to pale straw yellow with variable pigmentation of dorsal sclerites (Hille Ris Lambers & Stroyan 1959). On undersides of leaves of Quercus ilex in Europe (England, Wales, France, Italy, Greece, former Yugoslavia, and also Malta – Mifsud et al. 2009); rarely on other oaks. Very similar to M. boerneri; the distinction between these two species was discussed by Barbagallo & Stroyan (1982). Sexual morphs are unknown; apparently it is anholocyclic, at least in England and Sicily. Kennedy (1986) investigated the method of attachment of the tarsi to the smooth leaves of Q. ilex.

Myzocallis (Lineomyzocallis) spinosa Boudreaux & Tissot Alatae are yellow, with antennae ringed with black, broad black bands running from posterior margins of eyes down sides of thorax and along leading edges of wings, and all tibiae black; BL 1.6-2.0 mm. Immature alatae are pale yellow with paired quadrate patches on abdominal tergites 1-4 or 1-5. On leaves of Quercus spp. of the red oak group in eastern North America. Monoecious holocyclic; sexuales occur in September-October.

Myzocallis (Pasekia) taurica Quednau & Remaudière Colour of alata in life is unknown, pronotum with four longitudinal dark stripes and dorsal abdomen with four longitudinal rows of dark sclerites; BL 1.2-1.8 mm. On Quercus spp. (calliprinos, ?robur) in Turkey and Greece. Sexuales occur in October-November (Quednau & Remaudière 1994).

Myzocallis (Paramyzocallis) tenochca Remaudière & Quednau Colour of alatae in life is not recorded, probably yellowish with rather pale legs. The forewings have a dark band along the costa, the dorsal abdomen has variably developed spinal and marginal sclerites, and the siphunculi are dark; BL 2.0-2.5 mm Immature alatae have four longitudinal rows of dark dorsal abdominal sclerites (Remaudière & Quednau 1985). On Quercus crassipes in Mexico. Life cycle and sexuales are unknown.

Myzocallis (Castaneomyzocallis) tissoti Quednau & Remaudière Colour of alatae in life is not recorded, probably yellowish, with dusky, dark-banded antennae and tibiae; forewings as in M. castaneae, but with a rather truncate pterostigma only about five times longer than broad; BL 1.7-2.3 mm. On Castanea nana in Florida, USA. Sexuales occur in November (Tissot 1932; as M. castaneoides).

Myzocallis (Neomyzocallis) tuberculata Richards Alatae are yellow or pinkish with very variable dark dorsal body and forewing markings (see influentialpoints.com/Gallery and Quednau 1999); BL c. 1.5 mm. Immature alatae are yellow with dark spots at bases of dorsal abdominal hairs. On Quercus spp. (alba, macrocarpa, muehlenbergii)in eastern North America (erroneously reported to occur in western North America by Blackman & Eastop 1994). Quednau (1999) illustrated an ovipara collected on Q. macrocarpa in Quebec in late August, and an (alate) fundatrix collected in May in Minnesota.

Myzocallis (Lineomyzocallis) walshii (Monell) (= occulta Richards) Alatae in summer are bright yellow and have antennae ringed with black, broad bands of black pigment running down sides of thorax and along leading edges of wings, and black fore tibiae (see influentialpoints.com/Gallery); BL 1.6-2.0 mm. In spring they may be much more weakly pigmented (see Quednau 1999). Immatures are very pale yellow with variably developed dorsal sclerites. On Quercus spp. of the red oak group, common throughout eastern North America (Boudreaux & Tissot 1962), and introduced to the west coast; it has been collected in California on the native live oak, Q. agrifolia, and on planted red oaks in Oregon, Washington (aphidtrek.org) and British Columbia (as occulta). More recently it has been found on Q. rubra planted in France (Remaudière 1989a), and has since spread to England, Wales (Baker 2009b), Belgium, Netherlands (Piron 2009), Germany, Andorra, Poland, Switzerland, Hungary, Serbia, Czech Republic, Slovakia, Slovenia, Italy, Spain (Nieto Nafría & Mier Durante 1998), Portugal (Pérez Hidalgo et al. 2009b) and Turkey (Kök & Özdemir 2021). In Spain and Czech Republic it is parasitised by native Trioxys and Praon spp. (Pons et al. 2006, Havelka & Starý 2007). Oviparae and alate males occurred in Poland in September-October (Osiadacz & Wieczorek 2006), and Quednau (1999) illustrated an ovipara collected on Q. rubra in California in September. The male genitalia were described and illustrated by Wieczorek et al. (2011).

Myzodium Börner

Aphidinae: Macrosiphini

Three species with a nodulose head with rounded antennal and medial tubercles, and the characteristically-shaped antennal terminal process and cauda of moss-feeding aphids, although two were originally described from other hosts. K.S.Pike et al. (2010) provided a key to apterae and alatae.

Myzodium lutescens (Zhang & Qiao) Apterae are greyish yellow; BL c.1.6-1.8 mm. Alatae have a black dorsal abdominal patch and secondary rhinaria distributed III 44-54, V 18-19, V 3-4. On Carissa spinarum in Fujian Province, China (Zhang & Qiao 1998d, as Evallocotaphis), and subsequently collected from a species of Compositae (G. Qiao, pers. comm.) . The true host is more likely to be a moss.

Myzodium mimulicola (Drews & Sampson) Apterae are reddish green or brownish green with some orange colour between the siphunculi (see aphidtrek.org); BL 0.9-1.9 mm. Alatae have secondary rhinaria distributed III 17-35, IV 6-14, V 0-6, and abdomen with broad dark segmental cross-bands, sometimes partially fused into a more solid patch. It was originally described from populations occurring on lower, older leaves of Mimulus guttatus at two sites in California, USA. K.S.Pike et al. (2010) reported its wider occurrence in western USA including Alaska, on M. guttatus, on angiosperms growing in wet situations in two other families (Nasturtium, Veronica), and also on certain mosses (Aulacomnium, Brachythecium, Calliergon, Philonotis, Saniona). They provided a redescription and placed M. knowltoni as a junior synonym, based on morphological comparison and DNA sequence identity. No sexual morphs are known. It is now reported online (aphidtrek.org) that this species is probably heteroecious, the primary host in Idaho being a native North American species of Crataegus (C. douglasii).

Myzodium modestum (Hottes) Plate 16g (Fig.44e,g) Apterae are reddish brown to dark brown or olive, sometimes greenish posteriorly, dorsum shiny, antennae and legs brown, siphunculi blackish; BL 1.2-1.9 mm. Alatae have secondary rhinaria distributed III 21-45, IV 7-13, V 0-4, and a large dark dorsal abdominal patch. On various mosses, e.g. Catharinaea and Polytrichum (Polytrichaceae); Pohlia (Bryaceae); and Rhacomitrium (Grimmiaceae). It is also the only aphid recorded from Sphagnum (Sphagnaceae). It is not attended by ants. In Europe including Iceland, Jan Mayen Island, Greenland, in North America, and in Costa Rica (Villalobos Muller et al. 2012). K.S.Pike et al. (2010) provided host plant and morphometric data for collections made in western USA. It is possibly anholocyclic in Germany (Müller 1973b), but oviparae and alate males have now been found in Switzerland (Pérez Hidalgo et al. 2017), the first evidence of an aphid being able to complete its life cycle on mosses.

Myzosiphum Tao

Aphidinae: Macrosiphini

Two species, one with an unknown host in China, and one known only from the Ruykyu Islands, related to Myzus,but the siphunculi have subapical reticulation.

Myzosiphum ruykyuense Tao Colour of apterae in life is unrecorded, BL c.2.3 mm. Alatae have secondary rhinaria distributed III 26-28, IV 9-10, V 2-3. On Polygonum sp., Okinawa.

Myzotoxoptera Theobald

Aphidinae: Macrosiphini

One species similar to Rhopalosiphoninus but with less swollen siphunculi and a short pentagonal cauda.

Myzotoxoptera wimshurstae Theobald Plate 20h Apterae are shining dark olive green with blackish head, siphunculi and cauda; BL 1.5-1.9 mm. On Cardamine spp., feeding in damp situations at bases of stems and on roots, so they are difficult to collect. M. wimshurstae is only known from UK, and from Czech Republic (BMNH collection, leg. J. Holman), but presumably has a wider distribution in Europe. It is monoecious holocyclic, with oviparae and apterous males in September-October (Prior 1971).

Myzus Passerini

Aphidinae: Macrosiphini

About 55 species with gibbous antennal tubercles probably all palaearctic, and mostly Asian, in origin. Alatae typically have a dark central dorsal abdominal patch. Primary hosts of heteroecious species are Rosaceae, usually Prunus. Biology and secondary host associations are diverse. Some very polyphagous species (ornatus, persicae, antirrhinii) include trees among their numerous host plants. Accounts are available for central Europe (Heinze 1961a), Fennoscandia and Denmark (Heie 1994), UK and Ireland (Blackman 2010), Japan (Miyazaki 1971), China (Tao 1963), Korea (Choi et al. 2019), Java (Noordam 2004) and India (R.C. Basu & Raychaudhuri 1976a, Raychaudhuri et al. 1980b). Moritsu & Tokumoto (1972) keyed fundatrices on cherry trees in Japan.

Myzus (Nectarosiphon) ajugae Schouteden Apterae are small, brownish to dirty yellowish green, BL 1.4-2.0 mm, in tightly curled leaves of Ajuga spp. (genevensis, orientalis, reptans) usually in deep shade. Alatae have a black dorsal abdominal patch and secondary rhinaria distributed III 7-12, IV 0(-3). Monoecious holocyclic, with apterous males in October. Widely distributed in Europe; UK, Netherlands, Belgium, Germany, Switzerland, Italy (Sicily), Hungary and former Yugoslavia. For a detailed account see Meier (1954). 2n=12.

Myzus alectorolophi Heinze Apterae are reddish brown or yellowish brown, with siphunculi pale or dusky, occasionally with dark apices; BL 1.3-1.9 mm. On Rhinanthus alectorolophus in Germany, and also recorded from Finland (Heie 1994). Closely related to M. cerasi, and possibly a synonym. The life cycle is unknown.

Myzus amygdalinus (Nevsky) Apterae are pale green, covered with delicate pruinose secretion, with pale, tapering siphunculi; BL 1.5-1.9 mm. Alatae have brown head, black pterothorax and pale green abdomen with marginal sclerites (Nevsky 1929a). In colonies on undersides of leaves of Prunus amygdalus and P. mahaleb in Spring (April-July), causing leaves to curl spirally, but not to change colour (Vereshchagin & Narzikulov 1961, Narzikulov 1965c). In Central Asia (Turkmenistan, Tadzhikistan, Uzbekistan). The life cycle is apparently unknown.

Myzus (Nectarosiphon) antirrhinii (Macchiati) Apterae are mid grey-green to dark green, occasionally dark red; BL 1.4-2.2 mm. On leaves and young growth of numerous plants, on which it may be confused with M. persicae (Blackman & Paterson 1986).In Europe, east Asia, Australia (Hales et al. 2000) and western North America. Unlike M. persicae, it often forms large, dense colonies, and only produces alatae rather sporadically. Anholocyclic almost everywhere, and only producing alatae rather sporadically, so it is most often found on perennial plants. However, there is now evidence of a possible sexual phase in Japan (Shigehara et al. 2006). Separation from M. persicae is difficult except using enzyme or molecular analysis (Hales et al. 2000). R IV+V length is in the range 0.102-0.128 mm (persicae 0.090-0.122 mm), siphunculi are rather uniformly darker and more swollen distally, and the function (138 × ANT PT) + (708 × R IV+V) – (53 × ANT III) – (500 × ANT BASE VI) is usually greater than 58 (usually less than 58 in persicae). 2n=11, 12 (heterozygous), 13 or 14 (see Spence et al. 1998).

Myzus asamensis Takahashi Apterae are whitish yellow (original description), pale green or green, with black siphunculi; BL c. 1.7 mm. On foliage of “a plant resembling Saxifraga” in Japan, and subsequently collected from Saxifraga punctata in east Siberia (Pashchenko 1988a).

Myzus (Sciamyzus) ascalonicus Doncaster Shallot Aphid Plate 19a Apterae are shining pale brownish green, straw-coloured or dirty yellow, with tips of antennae and legs black (see influentialpoints.com/Gallery); BL 1.1-2.3 mm. Alatae have a dark dorsal abdominal patch and secondary rhinaria distributed either III 7-21, IV 0-6, V 0-3, or III 24-56, IV 12-39, V 2-18 (strangely, both types of antenna can occur in one individual). Extremely polyphagous, with hosts in more than 20 families, but particularly Alliaceae, Caryophyllaceae, Compositae, Brassicaceae, Liliaceae and Rosaceae (see also Blackman & Eastop 2000). Almost worldwide, including sub-Antarctic islands (Hullé et al. 2003, Hullé 2012), and apparently completely anholocyclic everywhere. Unknown before 1940, and possibly of hybrid origin, although the parentage is unclear. Often placed in subgenus Nectarosiphon, but it clearly does not belong there, and is closer to Myzus cymbalariae. 2n=12.

Myzus asiaticus Szelegiewicz Apterae are yellow, with brownish antennae and siphunculi; BL 1.7-2.2 mm. Alatae have shiny dark brown head and thorax and yellow abdomen with dark brown dorsal patch. Rolling and yellowing the leaves of Padus asiatica Kom. (=Prunus padus var. pubescens Regel & Tiling) in Mongolia in August (Szelegiewicz 1969a), and also reported from Padus racemosa (= Prunus padus?) in eastern Kazakhstan (Kadyrbekov 2009a).. The life cycle is unknown.

Myzus asterale Choi, Kim, Lee, Lee, Shin & Lee Colour of apterae in life is unknown; BL 1.6-2.1 mm. Other morphs and life cycle unknown. On an unidentified species of Aster in South Korea (Choi et al. 2019).

Myzus (Nectarosiphon) asteriae Shinji (Fig.15f) Apterae are yellow, BL c. 1.8 mm, on upper sides of leaves of Aster and related genera (Boltonia, Kalimeris) in Japan, Korea and east Siberia. Alatae have a dark head and thorax, and a large dark dorsal abdominal patch (Takahashi 1965c). This is probably the species listed by Essig & Kuwana (1918) as a Rhopalosiphum on Boltonia indica. 2n=12.

Myzus beybienkoi Narzikulov Apterae are yellowish green with a light dusting of wax, antennae brownish, legs and siphunculi dark-tipped; BL 1.7-2.1 mm. The cauda is triangular, only about as long as R IV+V. Alatae have an extensive dark dorsal abdominal patch on abdominal tergites 3-6, often broken into separate transverse bars on abdominal tergites 5-6. On leaves and shoot tips of Fraxinus spp. in spring, causing slight twisting of leaves. In south-west and central Asia (Turkey, Iran, Afghanistan, Kirghizia, Tajikistan, Kazakhstan). Apparently heteroecious holocyclic, migrating in July to an unknown secondary host. Return migrants and oviparae were found on Fraxinus in October-November (Narzikulov 1957).

Myzus biennis Sanborn Apterae have not been described; alatae are green with a dark dorsal abdominal patch, and BL c.2.6 mm. In colonies on terminal stems of Oenothera biennis in Kansas, USA. M. oenotherae could be a synonym.

Myzus boehmeriae Takahashi Apterae are white or yellowish white with pale appendages, BL 1.6-2 mm. On undersides of leaves of Boehmeria spp. in Japan, Taiwan and South Korea. Monoecious holocyclic in Japan; alatae males and oviparae occur in November (Miyazaki 1971).

Myzus borealis Ossiannilsson Apterae are brownish yellow with variably developed dark brown dorsal spots, to almost black; BL 1.3-1.6 mm. On Galium boreale in northern Europe (Sweden, Finland) and on G. rubioides in Ukraine (Bozhko 1976a, as M. rubioidis). It is reported also to occur in Iran (Hodjat & Rezwani 1987). Oviparae and alatae males were reared in Sweden (original description). Mitochondrial DNA sequence analysis indicates a close relationship with M. cerasi (Rakauskas et al. 2014a; İnal & Kandemir 2020).

Myzus brevisiphon A.N.Basu (Fig.42 l,m) Apterae are dark brown; BL 1.2-1.7 mm. Alatae have secondary rhinaria distributed III 14-23, IV 0-5. On undersides of leaves of Polygonum alatum, Polygonum sp.and Persicaria capitatum,sometimes curling and yellowing leaves. In India (West Bengal, Meghalaya, Uttar Pradesh). The life cycle is unknown; alate males were found in late September-October (A.K. Ghosh 1974c). To judge from a paratype slide in the BMNH collection, the synonymy with M. formosanus Takahashi proposed by Eastop & Hille Ris Lambers (1976) is incorrect.

Myzus cerasi (Fabricius) Cherry Blackfly (fig. 89D) Apterae on Prunus are shiny, very dark brown to black, with antennae and legs bicoloured yellow and black, and siphunculi and cauda wholly black (see influentialpoints.com/Gallery); BL 1.5-2.6 mm. Alatae have a yellow-brown abdomen with an extensive dorsal patch. In dense, often ant-attended colonies at shoot tips of Prunus spp., especially P. cerasus and P. avium, in spring, curling leaves and often doing severe damage. Heteroecious holocyclic, migrating to secondary hosts in Rubiaceae (Asperula, Galium), Orobanchaceae (Euphrasia, Rhinanthus), Plantaginaceae (Veronica), and (especially in North America) certain Brassicaceae (Capsella, Cardamine, Coronopus, Lepidium). Apterae on secondary hosts are shining dark brown to yellowish brown or olive green, with black siphunculi and brown cauda; BL 1.1-1.7 mm. In Europe, across Asia to Pakistan and India, and introduced into Australia, New Zealand and North America. Rakauskas (1984) studied phenology and natural enemies of M. cerasi in Lithuania. Cichoka (2007) described its life cycle in Poland. The male genitalia were described and illustrated by Wieczorek et al. (2011). Apparently a complex of species or subspecies with different life cycles and/or secondary host associations is involved (Gruppe 1988a); electrophoretic differences were found between populations on P. avium and P. cerasus (Gruppe 1988b), and mitochondrial DNA data coupled with multivariate morphometrics have now confirmed that M. cerasi pruniavium Börner should be regarded as a subspecies with P. avium as its only winter host (Rakauskas et al. 2014a – but see also İnal & Kandemir 2020). Some populations in northern Europe are monoecious holocyclic on Galium and Veronica, and differences from cerasi in morphology of the fundatrix (Dahl 1968) indicate that they may be specifically distinct, or should at least be regarded as a subspecies, for which the available name is M. cerasi veronicae (Walker). In Japan, M. cerasi umefoliae (Shinji) occurs on Prunus (Armeniaca) mume and is morphologically like European M. cerasi, but does not colonise cherries and migrates to Artemisia capillaris (Takahashi 1965c); Myzus prunisuctus, described from Prunus sp. in China (Zhang & Zhong 1980a), could perhaps be this aphid. M. alectorolophi, M. borealis and M. erythraeae are also very closely related members of the cerasi group. See also Blackman & Eastop 2000, p.306. 2n=10 (European and North American populations) or 2n=12 (India, possibly umefoliae; Khuda-Bukhsh & Pal 1986a, Kurl & Chauhan 1988).

Myzus (Nectarosiphon) certus (Walker) Apterae are pink to dark reddish‑brown (see influentialpoints.com/Gallery); BL 1.2-2.0 mm. Alatae have secondary rhinaria distributed III 8-20, IV 0(-4). On Caryophyllaceae (Cerastium, Dianthus, Stellaria)causing spotting and curling of leaves, and also on Violaceae. Monoecious holocyclic, with apterous males; anholocyclic populations also occur. In Europe, Iran, and North America. Closely related to and difficult to distinguish morphologically from M. persicae; distinguishing characters for alatae of the two species in trap catches are given by Hille Ris Lambers (1959). For a detailed account see Meier (1954). 2n=12.

Myzus cornutus Medda & Chakrabarti Apterae are pale, colour in life not recorded; BL c. 2.0 mm. Alatae have a dark central dorsal abdominal patch on abdominal tergites 4-6, and separate cross-bands on other tergites. Spring populations occur on Prunus cornuta on India (Uttar Pradesh), feeding on undersides of leaves, which are rolled longitudinally and turn red or yellow (Medda & Chakrabarti 1986b). It is also reported as common in the north-eastern hills of Pakistan (Naumann-Etienne & Remaudière 1995). There is also a record (on P. persica) in Turkey (Akyürek et al. 2019) which is doubtful and needs confirmation. Apparently heteroecious holocyclic; migration occurs during June to an unknown secondary host (Medda et al. 1986). Das & Chakrabarti (1990) described a new parasitoid species (Aphidius staryi) from this aphid.

Myzus (Sciamyzus) cymbalariae Stroyan Plate 19b Apterae are dull yellowish green or yellowish brown, to dark brown or crimson red (more pigmented in cold conditions); BL 1.2-2.0 mm. Alatae, unlike other Myzus, have more or less separate transverse dark bands on the dorsal abdomen. Polyphagous, colonizing a similar range of hosts to M. ascalonicus (especially species in Alliaceae, Caryophyllaceae, Compositae, Iridaceae, Liliaceae, Plantaginaceae, and Violaceae). Apparently M. cymbalariae is entirely anholocyclic, although males have been obtained in trap catches and reared in laboratory cultures (Brown 1983). Its origin is unknown; the present distribution indicates recent dispersal by man (England, Scotland, Switzerland, Italy, Spain, Rwanda, South Africa, Iran, India, Pakistan, Australia, New Zealand, Bolivia and Chile). It is now also recorded from and may be established in USA (Skvarla et al. 2017). 2n=12.

Myzus debregeasiae Noordam Apterae are yellowish white; BL 1.3-1.7 mm. They live singly on undersides of older leaves of Debregeasia longifolia (Urticaceae)in Java , and have also been collected from Wendlandia densiflora (Rubiaceae) (Noordam 2004).

Myzus (Nectarosiphon) dianthicola Hille Ris Lambers Apterae are deep yellow‑green; BL 1.3-2.0 mm. On Dianthus caryophyllus, usually in glasshouses, causing white or yellow spots or blotches on the leaves. Closely resembling M. persicae, but distinguishable from that species by the reaction of the host plant, and from M. certus by its colour in life. In North America, western Europe, Iran (Rezwani 1998) and New Zealand. This species is apparently completely anholocyclic. 2n=14 (heterozygous).

Myzus duriatae Noordam Apterae are brownish black; BL 1.3-1.5 mm On flower-buds of Aeschynanthus radicans in Java (Noordam 2004).

Myzus dycei Carver (Fig.55b) Apterae are green or yellow, with pale appendages; BL 1.4-2.0 mm. Alatae have secondary rhinaria distributed III 13-23, IV 1-2. On Urticaceae (Girardinia, Laportea, Urtica) and Ulmaceae (Hemiptelea, Ulmus)in Australia and eastern Asia (Japan, China, Korea, east Siberia, Nepal and India). The life cycle is unclear; M. dycei was found on Ulmus pumila and Hemiptelea davidii in South Korea by Lee et al. (2002c), who suggested host alternation, but these are probably alternative secondary hosts (Ulmaceae being allied to Urticaceae). 2n=12.

Myzus erythraeae Mamontova Apterae are shining brown, with dark siphunculi and cauda; BL c. 1.3 mm. On Centaurium erythreae (= Erythraea centaurium) in Ukraine. This species is not clearly distinguishable from M. cerasi, which has been recorded from Centaurium.

Myzus fataunae Shinji Apterae are pale green with darker head and thorax, and pale appendages, BL 1.0-1.6 mm. Alatae have 7-11 secondary rhinaria on ANT III, and without a dark abdominal patch. Described from Fatoua (as Fatauna) sp. (Moraceae; Shinji 1924), but since recorded only from Urticaceae (Boehmeria, Parietaria, Pilea; Takahashi 1965c, Miyazaki 1971). In Japan and Korea, and recently introduced to Florida, USA (S. Halbert, pers. comm.). Alate males have been collected in Japan in October (BMNH collection, leg. M. Sorin). 2n=8.

Myzus filicis A.N. Basu (Fig.43n) Apterae are creamy yellow; BL 1.5-1.7 mm. On underside of fronds of an unidentified fern in West Bengal, India, and also collected on unidentified ferns in Nepal (BMNH collection, leg. K.C. Sharma and R. van den Bosch). It was recently reported from Tibet (China), colonising Dennstaedtia appendiculata, by Su et al. (2014), who noted some differences from the original description, and tabulated morphometric data. The life cycle is unknown.

Myzus formosanus Takahashi (Fig.42n) Apterae are dark brown to black, with pale antennae and legs, and blackish siphunculi and cauda; BL 1.6-1.8 mm. Alatae are apparently undescribed. On Polygonum spp., rolling leaves. In Taiwan and India (West Bengal; David et al. 1972, as M. leptotrichus). 2n=12.

Myzus godetiae Shinji Apterae are pale green; BL c. 1.6 mm. Alatae have dark dorsal abdominal cross-bands. On Clarkia amoena in California, USA. Not recorded since the original description, which is not adequate for inclusion in a key. It could possibly be M. hemerocallis, or Abstrusomyzus valuliae.

Myzus hemerocallis Takahashi Apterae are pale yellowish green or greenish white (see influential points/Gallery); BL 1.6-2.4 mm. They attack basal parts of young leaves of Hemerocallis spp. and Agapanthus umbellatus. On plants grown in shade they may form large colonies in the flower-heads. Presumably of east Asian origin, this species is now widely distributed on Hemerocallis. Present distribution is India, Pakistan, China, Taiwan, Korea, Japan, Indonesia, Australia, New Zealand, Hawaii, Kenya (BMNH collection, leg. V.F. Eastop), South Africa, UK (since 2000; R.M. Smith et al. 2007), (France (Remaudière & Muños-Viveros 1992a), Madeira (on Scilla madeirensis; Aguiar & Ilharco 1997), and North, South and Central America. It does not really belong in the genus Myzus, as now confirmed by molecular data (Nieto Nafría et al. 2013b), and is possibly closer to the genus Hyalomyzus. The life cycle is not known. 2n=12 (for specimens from China, Kenya and Brazil, R.L. Blackman, unpublished data), but Chen & Zhang (1985a) recorded 2n=8 from China.

Myzus (Nectarosiphon) icelandicus Blackman Apterae are red-brown; BL 1.5-1.9 mm. Alatae have secondary rhinaria distributed III 13-17, IV 0-3. It feeds mainly on Caryophyllaceae (Cerastium, Silene, Stellaria), but also in summer it is common on various other plants in Crassulaceae, Brassicaceae, Gentianaceae and Polygonaceae. Only known from Iceland. Monoecious holocyclic with an abbreviated life cycle, oviparae and apterous males being produced in July-August on Cerastium and Sedum (Prior & Stroyan 1960, as M. polaris). 2n=10.

Myzus indicus R.C. Basu & Raychaudhuri Apterae are pale, BL 1.2-1.5 mm. On Boehmeria sp. (Urticaceae) in Meghalaya, India, and also recorded from Aneilema sp. (Commelinaceae) in Uttar Pradesh (Chakrabarti & Sarkar 2001). In most respects the description of this aphid agrees with the polyphagous species Myzus ornatus, differing only in the characters given in the key to aphids on Boehmeria. Other morphs are unknown.

Myzus inuzakurae Shinji The fundatrix and spring migrant alata of this species on Prunus buergeriana in Japan have recently been redescribed by Sorin (2011). Other morphs and life cycle are unknown.

Myzus isodonis (Takahashi) Apterae are yellow, with pale yellow appendages; BL c. 1.2 mm. On Isodon sp. in Japan and Korea. Probably this is not a Myzus; it could be the secondary host form of a Tuberocephalus. Lehrius papillicaudus Gredina, described from Elscholtzia in east Siberia, may be a synonym.

Myzus japonensis Miyazaki Apterae are pale yellow to yellowish green or green; BL 1.4-1.9 mm. Alatae have secondary rhinaria distributed III 17-24, IV 7-12, V 3-7. On undersides of leaves of Rosa rugosa in Japan and Korea. Oviparae and alate males were found in late October (Miyazaki 1968a).

Myzus kalimpongensis (M.R. Ghosh, R.C. Basu & Raychaudhuri) Colur of apterae in life is unrecorded, probably pale; BL 1.5-1.7 mm. On Artemisia sp. and Leonurus sibiricus (only one of which is likely to be the true host). In north-east India (West Bengal).

Myzus kawatabiensis Miyazaki (Fig.42j,k) Apterae are orange-red to brick-red with black head, siphunculi and cauda; BL 1.1-1.3 mm. Alatae have secondary rhinaria distributed III 12-17, IV 1-5, V 0-1. On Polygonum thunbergii, curling margins of leaves.In Japan and possibly east Siberia (as M. formosanus; Pashchenko 1988a). To judge from paratypes in the BMNH collection, the synonymy of this species with M. formosanus Takahashi (Eastop & Hille Ris Lambers 1976) is incorrect. This species might be a synonym of M. polygoniyonai Shinji, described from P. viscosum.

Myzus (Nectarosiphon) lactucicola Takahashi Apterae have dorsum shining brown-black, and appendages pale yellow; BL 1.5-1.7 mm. Alatae have rhinaria on ANT III-IV or III-V. On undersides of leaves of Lactuca spp., Crepidiastrum lanceolatum and Youngia japonica in Japan (Miyazaki 1971), and on Ixeris spp.in Korea (Lee et al. 2002c).

Myzus (Galiobium) langei (Börner) Plate 18i (Fig.29b,c) Apterae are somewhat flattened, dull yellowish to pale green, the abdomen tinged anteriorly with rosy red (see influentialpoints.com/Gallery); BL 1.3-1.9 mm. Alatae have a large dark dorsal abdominal patch. Immatures are bright rosy red. On Galium spp., stunting and deforming new growth so that the foliage becomes bunched like a “witches’ broom” (Stroyan 1950). Colonies are visited by ants. Widely distributed in Europe, and eastward to west Siberia and Kazakhstan. Narzikulov (1953) distinguished a population of smaller individuals on G. verum in Tajikistan as a separate species, M. (Galiobium) galinarium, subsequently relegated to the status of subspecies (M. langei ssp. galinarium)by Narzikulov & Daniyarova (1990), and also reported to occur in Kazakhstan (Kadyrbekov 2014d).

Myzus lefroyi R.C. Basu & Raychaudhuri Colour of apterae in life is unrecorded; BL c. 1.2-1.4 mm On an unidentified fern in West Bengal, India. The generic position of this species is uncertain; it should probably be in Micromyzus or Micromyzella.

Myzus ligustri (Mosley) Apterae are shiny yellow, with distal parts of siphunculi brownish black (see influentialpoints.com/Gallery); BL 1.0-1.5 mm. Sporadically common on privet hedges (Ligustrum ovalifolium, L. vulgare); leaves are rolled longitudinally into narrow tubes and spotted with yellow. M. ligustri was described from England (1841) before the introduction of the east Asian L. ovalifolium to Europe, and is not known from Asia, so presumably the native European L. vulgare is the type host. In Europe, and introduced into North America. Monoecious holocyclic, with oviparae and alate males in November. 2n=12.

Myzus (Nectarosiphon) linariae Holman Apterae are yellowish green to yellow with somewhat dusky head, antennae, legs and cauda, and siphunculi darker at apices (see aphidtrek.org); BL 1.6-2.3 mm. On upper parts and inflorescences of Linaria genistifolia in Czech and Slovak Republics, Serbia (Petrović 1998), Hungary, Italy and Turkey. A. Jensen (aphidtrek.org) has now reported its occurrence in western USA (see also Skvarla et al. 2017). Monoecious holocyclic in Europe, with oviparae and apterous males in September-October (original description).

Myzus lythri (Schrank) Apterae on Prunus are green to yellowish green with darker green longitudinal stripes; antennae, legs, siphunculi and cauda pale; BL 1.8-2.0 mm. Alatae are much darker and have a large quadrate dorsal abdominal black patch. On Prunus mahaleb in spring, in curled apical leaves. Heteroecious holocyclic, migrating to aerial parts of Lythrum spp., and occasionally to other plants in aquatic situations. There are also several records from Epilobium spp., and one from Rhamnus pushiana (in Oregon, USA; BMNH collection, leg. A Gutierrez). Throughout Europe, in North Africa (Tunisia), Middle East (Iran, Lebanon), central (Burundi) and southern Africa, central Asia, possibly in Korea (where males of this or a very similar species were attracted in large numbers to pheromone traps; Boo et al. 2000), and introduced into North America. 2n=12.

Myzus manoji R.C. Basu & Raychaudhuri Appearance of apterae in life unrecorded; BL c.1.2-1.4 mm. Described from an unidentified host in West Bengal, and subsequently recorded from Hedera nepalensis in Meghalaya (A.K. Ghosh 1976).

Myzus moriokae Shinji Apterae are dark green with black markings on thorax and sides of abdomen, dark siphunculi and a black cauda; BL c.1 mm. Alatae have black transverse bands on dorsal abdomen. On young shoots as well as older leaves of Boehmeria spp. in Japan. Not found since the original description, and the long hairs make its generic position uncertain (Takahashi 1965c). Oviparae were included in the original description.

Myzus (Nectarosiphon) mumecola (Matsumura) Apterae are very pale yellow-green to straw-coloured with pale antennae, legs, siphunculi and cauda; BL 2.0-2.4 mm. Alatae are much darker and have a broad dorsal abdominal patch with lateral extensions on abdominal tergites 3 and 4. Spring populations curl and discolour the leaves of Prunus armeniaca and P. mume in Japan; it occurs also in China, Taiwan and east Siberia, is also recorded from Prunus sp. in India (Uttar Pradesh; Medda et al. 1986), and has recently become established in apricot orchards in Italy (Panini et al. 2017), Hungary (Borbély et al. 2021) and Serbia (Petrović-Obradović 2021). It can be a serious pest of apricots (Pashchenko 1988a). Sexual morphs were described by Tomić et al. (2024) who also transferred the species to the subgenus Nectarosiphon. Some apterae collected on Lonicera quinquelocularis in Himachal Pradesh, India could perhaps be the secondary host form of this species (BMNH collection, leg. A.N. Chowdhuri), but this requires experimental confirmation. Bhagat (1981) recorded without comment that many apterae and some oviparae were collected on Rubia cordifolia in July in Kashmir (as umecola Shinji, which is considered to be a synonym of mumecola), but in view of the presence of oviparae (on a host other than Prunus) the identity of this sample also needs to be confirmed. 2n=12 (Chen & Zhang 1985b).

Myzus mushaensis Takahashi (fig. 89E) Apterae are bright yellow to yellowish green with pale antennae, legs and cauda; siphunculi sometimes pale brown but usually either distally or wholly black. BL 1.7-1.9 mm. Alatae have variable dorsal abdominal markings, often divided medially and/or intersegmentally and not forming a solid patch. On Prunus spp. (cherries) in Japan, Taiwan and Korea. Feeding by fundatrices and their progeny causes the leaf lamina to roll longitudinally upwards on both sides of the mid-rib (Moritsu & Tokumoto 1972; Moritsu 1983, p.109). Heteroecious holocyclic in Japan, the secondary host being Plectranthus (=Isodon) japonicus. Apterae on the secondary host are green, BL 1.0-1.1 mm. Specimens in the BMNH collection from Scutellaria sp. at Nainital, India (leg. K. Narayanan) are very similar and could also be this species. Sexuales are undescribed. Yano et al. (1984) studied predation by Elateridae.

Myzus (Nectarosiphon) myosotidis Börner Apterae are yellowish green; BL 1.5-1.9 mm. Alatae have secondary rhinaria distributed III 8-14, IV 0-5, V 0-2. In rolled leaves of Myosotis scorpioides (= palustris). In Europe (England, Netherlands, Sweden, Germany, Switzerland, Poland, Russia). Records from USA (California) are erroneous (Foottit et al. 2006). Monoecious holocyclic with oviparae and apterous males in September-October. For a detailed account see Meier (1954). 2n=12.

Myzus (Nectarosiphon) ninae Gredina Colour of apterae in life is unrecorded; BL c. 0.7 mm? (all measurements are possibly miscalculated). On Artemisia sp. in east Siberia (Gredina 1996).

Myzus obtusirostris David, Narayanan & Rajasingh Apterae are shining black with dark appendages; BL 1.2-1.7 mm. On grasses and bamboos in India, Pakistan and Sri Lanka; common on Zea in Pakistan (Hamid 1984). The life cycle is unknown; males are alate (BMNH collection, leg. C.I.B.C.). M. maculocorpus R.C. Basu & Raychaudhuri, described from an unidentified grass in West Bengal and also reported from Setaria in Pakistan (Naumann-Etienne & Remaudière 1995), may be a pale form of this species. 2n=12.

Myzus oenotherae Williams Apterae are pea green, BL not recorded. On stem and leaves on Oenothera biennis in Nebraska, USA. Possibly this is a synonym of M. biennis.

Myzus ornatus Laing Violet Aphid Plate 18f Apterae are somewhat dorso-ventrally flattened, pale yellow or green, marked dorsally with a pattern of dark green or brownish dots and transverse flecks (see influentialpoints.com/Gallery); BL 1.0-1.7 mm. They live singly on the leaves of host plants in many different plant families including especially Bignonaceae, Compositae, Lamiaceae, Polygonaceae, Primulaceae, Rosaceae, and Violaceae. M. certus can be found on both upper and undersides of leaves, often feeding away from the main veins. World-wide on cultivated ornamental plants, but although common in India since 1956, there is still only one record from south-east Asia (New Guinea). Anholocyclic populations occur throughout the world, in colder climates probably overwintering in glasshouses, on pot plants, or in sheltered situations. Sexual morphs are reported from northern India; the alate male was described by Chakrabarti & Raychaudhuri (1975), and Maity & Chakrabarti (1981) described the oviparae. Specimens from Spiraea lindleyana in the Himalayas (BMNH collection, leg. S. Sekhon) are possibly the primary host forms. See also Blackman & Eastop (2000, p.308). 2n = 12.

Myzus (Prunomyzus) padellus Hille Ris Lambers & Rogerson Apterae on Prunus are bright yellow with blackish brown head; antennae, femora and tibiae bicoloured yellow and blackish brown, and siphunculi black. BL 2.0-2.3 mm. Alatae have a yellow abdomen with a relatively small and weakly pigmented dorsal patch. Red and yellow blister leaf-galls are formed on Prunus padus in spring (Hille Ris Lambers & Rogerson 1946). In Europe (England, Norway, Sweden, Finland, Austria, Switzerland, Russia, Turkey), Iran, Georgia (Barjadze 2008), and across Russia and Central Asia to east Siberia (Pashchenko 1988a) and Korea (Choi et al. 2019). The record from Turkey refers to blackish apterae (Şenol et al. 2014b) and therefore needs further confirmation. Aphids collected on Brassica in Fujian province, China may also be this species (Qiao & Zhang 1999c, as Tuberocephalus longqishanensis). Apparently heteroecious holocyclic, migrating to Lamiaceae (Galeopsis spp. – records from Norway and Sweden; Tambs-Lyche 1957) and Orobanchaceae (Pedicularis verticillata, Rhinanthus sp. – specimens from Austria; BMNH collection, leg. H.L.G. Stroyan & R.N.B. Prior). Apterae on secondary hosts are smaller (BL 1.3-1.9 mm), bright yellow to yellowish-green and look very different from the bird cherry aphids. However, host alternation has not been fully confirmed by experimental transfers, and sexuales are unknown.

Myzus (Nectarosiphon) persicae (Sulzer) Peach-Potato Aphid, Green Peach Aphid Plate 18h Apterae vary from whitish or pale yellowish green to mid-green, rose-pink or red, rather uniformly coloured, not shiny, often darker in cold conditions (see influentialpoints.com/Gallery); BL 1.2-2.3 mm. The fundatrix and her progeny have tapering, unswollen siphunculi. Alatae have a dark dorsal abdominal patch and 7-14 secondary rhinaria on ANT III only. Immature alatae are often pink or red, especially in autumn populations (and immature males are yellow). Spring colonies curl young leaves of the primary host, Prunus persica or, in north-eastern North America, P. nigra (Shands et al. 1969). Heteroecious holocyclic, migrating to secondary hosts in over 40 different plant families, and an important vector of numerous plant viruses (Mackauer & Way 1976; Blackman & Eastop 2000, p.308-9). Secondary colonisation of young growth of various Prunus spp. also commonly occurs. M. persicae is probably of east Asian origin, but is now world-wide. Margaritopoulos et al. (2009) have used microsatellite markers to study the global distribution patterns of various genotypes, and Hlaoui et al. (2019) also used microsatellites to study its genetic diversity in Tunisia. Populations are partially anholocyclic on secondary hosts in milder climates, and exclusively so in tropics or in the absence of the primary host. The genetic basis of its ability to colonise numerous hosts without specialisation has been studied by Mathers et al. (2017). Mandrioli et al. (2019a) compared the genomes of two clones of M. persicae and discussed the differences. Also see Y. Li et al. (2020) and Mathers et al. (2021) for recent research involving chromosome-level genome assembly and comparisons with Acyrthosiphon pisum and Rhopalosiphum maidis). Populations colonising tobacco are distinguished as a subspecies, M. persicae ssp. nicotianaeBlackman, and this form also has a sexual phase on peach in some areas, e.g. Greece (Margaritopoulos et al. 2000, 2007). An analysis of the complete mitochondrial genome of a M. persicae clone reared on tobacco (H. Yang et al. 2017) should presumably be applied to this subspecies. A population in Germany with holocyclic overwintering on Lycium halimifolium (= L. barbarum), and distorting young growth of that plant in spring, was given subspecies status as M. persicae ssp. dyslycialis by Müller (1955b), but should perhaps be regarded as a distinct sibling species. The taxonomy of M. persicae and its close relatives was discussed by Blackman & Eastop (2007, 2017). 2n=12 (incl. ssp. nicotianae).

Myzus philadelphi Takahashi Apterae are orange-yellow in life with four conspicuous dark green-black dorsal transverse stripes, and a similar dark longitudinal stripe running down from abdominal tergite 4 between the siphunculi (the stripes all disappear in preserved material), dark antennae, legs and siphunculi, and a pale cauda; BL 2.2-2.6 mm. Collected on Philadelphus satsumi and Deutzia crenata in spring in Japan, and on Hydrangea macrophylla form otaksa in Korea (Paik 1972). Miyazaki (1971) described the fundatrix.

Myzus pileae Takahashi Apterae are pale yellowish green; BL c.1.4-1.5 mm. Alatae have secondary rhinaria distributed III 19-23, IV 2-5. On Pilea hamaoi in Japan.

Myzus (Nectarosiphon) polaris Hille Ris Lambers Colour of apterae in life is unrecorded, probably reddish; BL 1.8-2.1 mm. Alatae have secondary rhinaria distributed III 12-18, IV 1-4. On Cerastium alpinum in Greenland, and also found in north-eastern Canada (northern Quebec, Baffin Island) and on Wrangel Island in the Siberian arctic (Stekolshchikov & Khruleva 2014). Monoecious holocyclic with an abbreviated cycle, oviparae and apterous males being produced in July-August.

Myzus prunense Choi, Kim, Lee, Lee, Shin & Lee Colour of apterae in life is unknown; BL 1.7-2.2 mm. Other morphs and life cycle unknown. Collected in May from an unidentified species of Prunus in South Korea (Choi et al. 2019).

Myzus (Nectarosiphon) raphanense Choi, Kim, Lee, Lee, Shin & Lee Colour of apterae in life is unknown; BL 1.3-1.8 mm. Other morphs and life cycle unknown. On Raphanus sativus in South Korea (Choi et al. 2019).

Myzus siegesbeckiae Takahashi(= yamatonis Miyazaki) (Fig.48e, fig. 89C) Apterae on Prunus are black in life, with antennae and legs pale except at apices, and siphunculi and cauda black; BL 1.4-1.6 mm (Miyazaki 1971, as M. yamatonis). Alatae are black in life, but with variably developed abdominal sclerotisation, the central patch being irregular and with large perforations. On Prunus spp. (cherry trees), especially P. yedoensis, spring populations causing heavy leaf-curl, and also recorded from P. persica and P. (Armeniaca) mume. In Japan, Korea, China, Taiwan and India (Simla; BMNH collection, leg. S. Chakrabarti). Heteroecious holocyclic, migrating to Lamiaceae (Salvia nipponica, Plectranthus, Isodon). although populations on Prunus may persist throughout the growing season. Apterae on summer hosts look very different; they are yellow or whitish, with pale appendages, and have BL 0.8-1.1 mm. Yano et al. (1984) studied predation by Elateridae.

Myzus siegesbeckicola Strand (Fig.15d,e) Apterae are pale to dark green, BL 1.2-1.5 mm. Alatae have paired dark spino-pleural sclerites on abdominal tergites 4 and 5 (R.C. Basu & Raychaudhuri 1976a, and BMNH collection, leg. M. Miyazaki). On Sigesbeckia spp., and also recorded from other Compositae (Artemisia, Aster, Chromolaena, Montanoa), in Asia (Taiwan, Sumatra, Japan, Korea, east Siberia, India, Pakistan). Life cycle is unknown, but Maity & Chakrabarti (1981) described an alate male found on Sigesbeckia orientalis in October in north-west India. Possibly this species would be more correctly placed in Tuberocephalus.

Myzus sorbi Bhattacharya & Chakrabarti Apterae are yellowish, BL 1.6-1.9 mm . On undersides of leaves of Sorbaria tomentosa at above 2,000 m in north-west India and Pakistan. Feeding causes leaves to curl at margins, and heavy infestations cause twisting, shrinking and yellowing of leaves. Monoecious holocyclic, with oviparae and alate males in October-November (Medda & Chakrabarti 1987). 2n=12.

Myzus (Nectarosiphon) stellariae (Strand) Apterae are green, BL 1.6-2.0 mm. On Stellaria aquatica (= Myosoton aquaticum) and Stellaria sp. in Taiwan, Japan and Korea (Choi et al. 2019). Takahashi (1965c) provided a redescription.

Myzus terminalae Sath & Jahvad NOMEN NUDUM Apterae are dark greenish-black; BL c. 1.7 mm. On Terminalia tomentosa in India (Sathe & Jahvad 2008). The combination of Myzus-like head and siphunculi more than 0.25 of body length appear to be distinctive. However the failure to designate type specimens in accordance with the ICZN code renders the name and description invalid.

Myzus (Nectarosiphon) titschaki (Börner) Colour of apterae is unrecorded; BL c. 1.8 mm. Described from specimens sieved from grass, sedge or rush tussocks, but subsequently collected from Stellaria holostea (BMNH collection, leg. D. Hille Ris Lambers), which is a more probable host. Only known from Poland.

Myzus varians Davidson Plate 18g Apterae are pale green or yellow-green with banded antennae, and distal parts of siphunculi contrastingly black (see influentialpoints.com); BL 1.7-2.3 mm. Alatae are very dark, with a large dorsal abdominal black patch. Spring colonies tightly roll the young leaves of Prunus persica longitudinally. Heteroecious holocyclic, migrating to Clematis spp., but populations persist on peach throughout the growing season (Alma & Arzone 1983). Presumably it is native to eastern Asia, where it is recorded from India, China, Taiwan, Korea, Japan, Thailand and Vietnam. Introduced into Europe in about 1947, and subsequently to south-west Asia, where it has become a serious pest on peaches (Arzone & Alma 1984, Tkachuk 1988). It is a vector of plum pox virus (Sharka disease). M. varians has also been in North America since at least 1912, and spring populations on peach were reported by Essig (1917) although most subsequent records are from Clematis (Stoetzel & Miller 1998). A single alata was recently reported from Chile (Nieto Nafría et al. 2018). See also Blackman & Eastop 2000, p.309). 2n=12.