The Aphids

Eichinaphis pamirica Narzikulov Apterae are greenish brown to dark brown, with a pair of dark spots on the dorsal abdomen; BL 1.3-1.6 mm. In small colonies on stems and young growth of Krascheninnikovia spp. in Tajikistan (Eastern Pamirs) and south-east Kazakhstan (Kadyrbekov & Aoitzhanova 2005), and on Krascheninnikovia sp. in Iran (BMNH collection, leg. G. Remaudière) and Mongolia. A single alata has been trapped in Pakistan (Naumann-Etienne & Remaudière 1995). B. Zhang (2015) reported its occurrence in Mongolia and provided a detailed redescription of apterae and alatae. Other morphs and life cycle are unknown. E. turanica Kadyrbekov, described from Krascheninnikovia spp. in the desert region of Kazakhstan, is differentiated by having longer, thicker dorsal hairs on more tuberculate bases, and by higher siphunculus/cauda and ANT/BL ratios. However, Iranian specimens show intermediate characters, and it seems likely that only one variable species is involved.

Elatobium Mordvilko

Aphidinae: Macrosiphini

Of the eight species currently placed in Elatobium, five are conifer feeders, one lives on Salix, one on Trochodendron and one on Artemisia. The Salix feeder probably belongs in Cavariella subgenus Cavariellinepicauda (see under Cavariella hidaensis), and the other two non-conifer feeders are possibly either misplaced in Elatobium or not collected on their true host plants. The Japanese species were revised by Miyazaki (1971), although his concept of the genus is broader, including related aphids (Ericolophium, Neoacyrthosiphon) that live on Ericaceae.

Elatobium abietinum (Walker) Plate 14 a,b Apterae are green (see influentialpoints.com/Gallery); BL 1.0-2.0 mm. They feed on the needles of Picea spp. and, much less commonly Abies spp., records from which may be referable to E. blackmani. In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), Iran (Rezwani et al. 1994), Central Asia (Kadyrbekov 2017a), Australia (Tasmania), New Zealand (Zondag 1983), Chile (Carrillo 1977), Argentina, Falkland Islands and North America. Monoecious holocyclic (with alate males) in parts of continental Europe (von Scheller 1963), apparently anholocyclic elsewhere. Spring colonies on Picea spp., especially P. sitchensis, cause discoloration and loss of old needles, sometimes involving serious defoliation (Parry 1969). There is a large literature on this aphid and its impact on spruce in western Europe, reviewed by Day et al. (1998). Overwintering survival was studied by Powell & Parry (1976), and flight activity by Carter & Cole (1977). Variation in susceptibility to attack by E. abietinum has been studied between species of Picea (Nicholls 1987), and between provenances (Carter & Nicholls 1988, Day 1984) and individual trees (Day 1986) of P. sitchensis. Halldórsson et al. (2004) found genetic differences between anholocyclic populations in the countries of north-west Europe. Specialised parasitoids are Lysaphidus schimitsheki (Stary 1974) and Ephedrus koponeni (Halme 1992). 2n=18.

Elatobium blackmani Binazzi & Barbagallo Apterae are greenish, similar in colour to needles of host; BL 1.2-1.8 mm (Binazzi & Barbagallo 1997). On Abies spp. in southern Europe (Italy) and the Middle East (Turkey; the specimens referred to under E . momii in Blackman & Eastop 1994), but possibly more widespread. Sexuales and life cycle are unknown.

Elatobium chomoense Zhang in Zhang & Zhong Colour of aptera in life is unknown; BL 2.2-2.5 mm. On Artemisia sp. (from original description; host identified as A. arenaria on label of slide with holotype) in Tibet. The host is unusual for this genus and requires confirmation.

Elatobium laricis (Rupais) Apterae are broadly oval, green, shining, without wax, similar in colour to larch needles; BL 1.5-1.8 mm. On Larix sibirica in east Siberia (Rupais 1974). Other morphs and life cycle are unknown.

Elatobium momii (Shinji) Apterae are green; BL 0.9-1.6 mm. On needles of Abies firma in Japan. Shinji (1941) also records Taxus cuspidata as a host, but the species involved requires confirmation. Sexuales and life cycle are unknown.

Elatobium piceanum (Inouye) Apterae are green; BL 1.0 mm. On needles of Picea jezoensis in Japan. This species has been synonymised with E. momii, but Binazzi & Barbagallo (1997) pointed out differences in PT/ANT BASE VI ratio (1.67-2.50 in piceanum, 2.30-3.60 in momii) and other characters. Sexuales and life cycle are unknown.

Elatobium trochodendri Takahashi Apterae are pale yellowish green, with a dark green thorax, a large dark green dorsal abdominal patch, and siphunculi that are black on distal two-thirds; BL 1.3-2.2 mm. On Trochodendron aralioides in Japan, feeding on upper sides of leaves along mid-ribs (Takahashi 1960). Sexuales and life cycle are unknown.

Eomacrosiphum Hille Ris Lambers

Aphidinae: Macrosiphini

A genus for one Macrosiphum-like species with 5 hairs on first tarsal segments and other distinctive features.

Eomacrosiphum nigromaculosum (MacDougall) ( = Bipersona hottesi Knowlton & Smith) Apterae are bright red with black bands or patches on dorsal abdomen, black siphunculi held at right-angles to body, and black and yellow legs; BL 2.0-2.8 mm. On stems of wild and cultivated Rosa spp. in western North America. Monoecious holocyclic with alate males (Palmer 1952).

Eonaphis Essig

Greenideinae: Schoutedeniini

A genus for four species in Africa differing from Schoutedenia in the presence of spinal processes on the head (two pairs) and all segments to abdominal tergite 7, and marginal processes at least from abdominal tergites 4 and 7. Remaudière (1988) revised the genus and provided a key to species.

Eonaphis crotonis Quednau Apterae are broadly oval, somewhat flattened, dirty brownish yellow, shining, with a diffuse brown patch on dorsal abdomen, pale finger-like dorsal processes and dark brown conical siphunculi; BL 1.1-1.7 mm. Alatae have forewing veins dark-bordered, the media being once-branched. In large colonies on leaves and young branches of Croton subgratissimus, especially along the mid-ribs on the lower leaf surfaces, attended by ants (Quednau 1962). In South Africa. Sexuales and life cycle are unknown.

Eonaphis euphorbiae Quednau Apterae are dirty brownish to blackish green, shiny; BL 1.1-1.6 mm. Alatae have secondary rhinaria distributed ANT III 8-16, IV 0-2 (Remaudière 1988). On Euphorbia spp. in Madagascar. Oviparae were found in November.

Eonaphis pauliani Essig Colour of apterae in life is unrecorded; BL 1.0-1.6 mm. Alatae have secondary rhinaria distributed ANT III 39-44, IV 3-4 on IV (Remaudière 1988). On Euphorbia spp. in Madagascar. The original description apparently confused the host plant with that of Paulianaphis madagascariensis (see Remaudière 1988).

Eonaphis phyllanthi Remaudière Apterae mainly pale, with antennae dark beyond the midpoint of ANT III, and dark conical siphunculi; BL c. 1.1-1.2 mm. On Phyllanthus sp. in Madagascar (Remaudière 1988). Other morphs and life cycle are unknown.

Eotrama Hille Ris Lambers

Lachninae: Tramini

Four or five species in south-west and central Asia differing from Trama in having a longer rostrum and shorter hind tarsi with relatively well-developed HT I. Czylok (1990) reviewed the genus, transferring some species from other genera. The host of one Eotrama species is unknown and another, E. orientalis (Narzikulov), was described from apterae collected in Tajikistan on the trunk of the non-native tree Robinia pseudoacacia, which is unlikely to be the true host.

Eotrama bazarovi (Narzikulov) Apterae are pale, BL 5.0-5.5 mm. On roots of Myricaria alopecuroides (= bracteata) in Tajikistan. Possibly these are large specimens of E. tamaricis.

Eotrama moerickei Hille Ris Lambers Apterae are pale yellowish; BL 3.4-4.2 mm. Apterae have secondary rhinaria distributed III 0-15, IV 4-10, V 3-6, VI 0-2, and alatae have them distributed III 35-36, IV 7-8, V 5-7, VI 1-2. On roots of thistles (Cirsium sp.?) in Lebanon.

Eotrama tamaricis (Nevsky) Creamy white aphids, BL 3.9-4.7 mm, attended by ants. Originally described from a single alata on roots of Tamarix ramosissima in Uzbekistan (Nevsky 1951, as Eulachnus tamaricis), but Trama xerophilaphica, apterae and alatae of which were described from the same host in Kazakhstan (Juchnevitch & Kan 1971), was placed as a synonym by Mamontova (1991), who provided a redescription. Czylok (1990) placed T. xerophilaphica in Eotrama (see also Remaudière & Remaudière 1997, p. 301).

Epameibaphis Oestlund

Aphidinae: Macrosiphini

Three North American species on Artemisia and Seriphidium, with the typical stilletto-shaped R IV+V of aphids feeding on Anthemideae, and related to Pseudoepameibaphis, Flabellomicrosiphum and the palaearctic genus Coloradoa, but with unusual knobbed siphunculi. Knowlton & Smith (1936) reviewed the genus.

Epameibaphis atricornis Gillette & Palmer Apterae are frosted brown to greenish with black appendages; BL 1.2-1.4 mm. On Artemisia longifolia and Seriphidium tridentata in western USA. Sexual morphs are not recorded.

Epameibaphis frigidae (Oestlund) (Fig.12e) Apterae are pale glaucous green, with frosted appearance due to numerous pale dorsal hairs (Gillette & Palmer 1932b); BL 1.0-1.3 mm. On leaves and stems of Artemisia frigida in western USA and Canada (Saskatchewan, Manitoba). Monoecious holocyclic; oviparae were collected in November in Colorado (BMNH collection, leg. L.C. Bragg).

Epameibaphis utahensis Knowlton & Smith (Fig.12f) Apterae are greyish-green with numerous pale dorsal hairs giving them a whitish to grey appearance; BL 0.9-1.3 mm. On Seriphidium tridentata in western USA. Monoecious holocyclic; oviparae and alate males found in October in Utah (original description, and BMNH collection, leg. G.F. Knowlton, 1969).

Ephedraphis Hille Ris Lambers

Aphidinae: Aphidini

Four palaearctic species with the general characters of Aphis, but with a distinctive R IV+V, the microsensilla being placed laterally on a short, acute, beak-like process (Nieto Nafrìa & Mier Durante 1985). B. Zhang & Qiao (2013) reviewed the genus.

Ephedraphis ephedrae (Nevsky) Apterae are blackish green with black eyes, antennae yellowish except at base, legs with femora brown and tibia yellow, and dark siphunculi; BL 1.4-2.0 mm. Colonising stems of various Ephedra spp. in Europe (Spain, France, Ukraine) and Asia (Turkey, Iran, Turkmenistan, Uzbekistan, Kazakhstan, Tajikistan, Pakistan, northern India). Monoecious holocyclic, with oviparae and alate males in late September to early November in Central Asia (Narzikulov et al. 1973). Mamontova-Solukha (1963) described a subspecies from Ukraine, E. ephedrae ssp. taurica, and this subspecies is also reported to occur in Kazakhstan, but the specimens in the BMNH collection come within the range of variation shown by other populations of E. ephedrae.

Ephedraphis gobica Szelegiewicz Apterae are dark green, almost black; BL c. 2.3 mm. On Ephedra spp. in Mongolia and China, and also recorded from east Siberia and western Kazakhstan (Kadyrbekov 2017a). Other morphs are undescribed. Jia et al. (2004) studied development in the laboratory. A population in western China was described as a subspecies, E. gobica ssp. xinjiangica (Kadyrbekov et al. 2002).

Ephedraphis haloxylon Qiao Apterae are dark brown, BL 1.8-2.2 mm. On leaves and stems of Haloxylon ammodendron, causing yellowing and distortion of stems. In Qinghai Province, China (Qiao et al. 2006d).

Ephedraphis mongolica B. Zhang & Qiao Apterae are dark brown to blackish brown, with transverse white wax dorsal stripes; BL c.1.5-1.7 mm. On stems of Ephedra sp. in an arid habitat at 1800m altitude in Umnogovi Province, Mongolia (B. Zhang & Qiao 2013).

Epipemphigus Hille Ris Lambers

Eriosomatinae: Pemphigini

A small genus of poplar gall-forming species in east Asia, resembling Pachypappa in that the fundatrices have no wax pore-plates and the alate emigrants and sexuparae have secondary rhinaria with ciliated rims, but also with some characters of Pemphigus, e.g. alatae have forewings with unbranched media and the first instar exules have short empodial hairs. Chakrabarti & Banerjee (1993b) reviewed the genus. The Chinese species were reviewed by G. Zhang et al. (1999c). G. Zhang et al. (1995) erected a new subgenus of Thecabius, subgenus Oothecabius Zhang 1995, for some species with unbranched media and ciliated secondary rhinaria from China. Oothecabius is here treated as a synonym of Epipemphigus.

Epipemphigus imaicus (Cholodkovsky) Galls on Populus ciliata and P. cathayana are elongate, caterpillar-like or cockscomb-like (figs 131 A,B), reddish-green, 4-7 cm long, on upper surface of leaf usually alongside the mid-rib, with an opening on underside (A.K. Ghosh et al. 1981, Maity & Chakrabarti 1981. The second generation are alatae (BL 2.0-2.6 mm), emerging from galls in May-July and migrating to form colonies on roots of Polygonum alatum (Chakrabarti & Banerjee 1991b, 1993a; Chakrabarti 2021). Apterae on Polygonum roots are wax-secreting and have BL 1.8-2.2 mm; they live inside small cocoon-like chambers on fine rootlets (Chakrabarti & Banerjee 1993b). Sexuparae return to poplar in November-December (Chakrabarti et al. 1985; Chakrabarti 2021). A record of galls of E. imaicus on Ailanthus glandulosa (Chakrabarti et al. 1971) must be in error, and a record from the roots of Pinus wallichiana (Habib & Ghani 1970) was possibly a misidentified Prociphilus. In Pakistan, northern India and China. Pemphigus chomoensis Chang and P. yunnanensis Chang, described from Populus spp. in China (Chang & Zhong 1979), are very similar and possibly both synonyms of E. imaicus. 2n=18.

Epipemphigus marginalis Chakrabarti & Banerjee Galls on Populus ciliata in northern India are greenish yellow to yellowish, 2.5-3.0 cm long, produced on the marginal area of the leaf lamina. Second generation are all alatae, BL 1.7-2.2 mm, migrating in May-June to found colonies on roots of Impatiens falcifer (Chakrabarti & Banerjee 1993a; subject to experimental confirmation, as there is possible confusion with E. niisimae, a species not yet recorded from India). Also recorded from Bhutan on P. nigra (Chakrabarti & Das 2014). Alate sexuparae return to P. ciliata in September-October. Chakrabarti & Banerjee (1993b) described all morphs.

Epipemphigus nanjingensis (Zhang) Galls and appearance in life are undescribed. BL of emigrant alata (one specimen) c. 1.9 mm. On Populus cathayana in Jiangxi, China (Zhang et al. 1995, in Thecabius subg. Oothecabius). The life cycle is unknown.

Epipemphigus niisimae (Matsumura) Galls on Populus spp. (koreana, suaveolens) are elongate, caterpillar-like or cockscomb-like, yellow to crimson-red, along mid-rib of leaf on upper surface with opening underneath (Aoki 1975). Second generation are all alatae (BL 2.1-2.4 mm), emerging from galls in June-July. The secondary host generations have not yet been described, but Aoki et al. (1996a) tranferred alate emigrants from galls on P. maximowiczii (= suaveolens) and established colonies on roots of Impatiens nolitangere. Records from Compositae/Asteraceae (Agrimonia pilosa, Bidens tripartita) are based on uncertain identifications (Aoki 1975), and the identity of exules from Impatiens in India described as those of E. marginalis (see above) also needs confirmation. Alatae sexuparae are produced in September. Aoki & Makino (1982) studied gall usurpation and lethal fighting between the heavily-armoured 1st instar fundatrices, and Aoki et al. (1996a) studied defensive behaviour of immatures in the gall. The species is known on its primary host from east Siberia, Japan and Korea. It is probably also in China, as Prociphilus populi Tao 1970 and Epipemphigus sequelus Zhang (in Zhang et al. 1995, as Thecabius (Oothecabius) sequelus) are likely to be synonyms. 2n=20.

Epipemphigus sanpupopuli (Chang) Galls on Populus cathayana in China (Beijing) are smooth and pillow-like, on the upper side of the leaf, with an opening on the underside. Emigrant alatae, BL c. 2 mm, were collected in May (Chang & Zhong 1979c, as Sanpupemphigus). The life cycle is unknown. [Synonymised with E. imaicus in Remaudière & Remaudière (1997), but the hairy, club-shaped antennal terminal process coupled with the presence of siphunculi with partly sclerotised rims is distinctive.]

Ericaphis Börner

Aphidinae: Macrosiphini

A holarctic genus of about 12 Myzus-like North American aphids associated with Rosaceae, Ericaceae, and Liliaceae, and three European species on Ericaceae. The rims of the secondary rhinaria have a fine cilia-like fimbriation or striation, although this can sometimes be difficult to resolve. The nearctic species were previously placed in Fimbriaphis. Taxonomic problems with this genus in western USA are discussed online at aphidtrek.org. Blackman (2010) reviewed and keyed the four species that occur in UK, which include two introduced from North America. Pike & Starý (2003) provided a key to species, and also summarised their associations with hymenopterous parasitoids.

Ericaphis avariolosa (David, Rajasingh & Narayanan) Apterae are green; BL c.2 mm. On Rosa macrophylla in Himachal Pradesh, India. Other morphs and life cycle are unknown. This species was transferred from Myzaphis by Kanturski et al. (2018b).

Ericaphis ericae (Börner) Apterae are green or brownish green, with tips of antennae and legs black (see influential points/gallery); BL 1.1-1.7 mm. Alatae have a dark dorsal abdominal patch with clear windows. On Erica spp., especially E. tetralix, feeding on flowers and in shoot apices, where it is often overlooked because of small size and cryptic colouration. It is also recorded from Calluna vulgaris and Daboecia cantabrica. In northern and western Europe, east to Poland and south to the Iberian peninsula. Monoecious holocyclic, with oviparae and apterous males in September-October.

Ericaphis fimbriata (Richards) Apterae are pink or green: BL 1.1-1.9 mm. Alatae have a dark dorsal abdominal patch. On cultivated strawberries (Fragaria sp.) and Rosa nutkana (BMNH collection, leg. C.-k. Chan) in western North America (Oregon, British Columbia) and Hawaii. It is also recorded from blueberry, Vaccinium corymbosum (Forbes 1962), and the ecology of populations overwintering as eggs on this host has been studied by Raworth (2004). However it would probably be more correct to apply the earlier name E. scammelli (Mason) to blueberry-feeding populations (see below). In fact in British Columbia aphids identified as E. fimbriata are apparently monoecious holocyclic on both rosaceous and ericaceous hosts (C.-k. Chan, pers. comm.), indicating that the name is probably being applied to a complex of morphologically similar species, which includes E. scammelli (q.v.). 2n=14.

Ericaphis gentneri (Mason) Apterae are yellow-green to green,broadly spindle-shaped; BL 1.3-2.4 mm. Alatae have a broken dark dorsal patch. On leaves and young shoots of various woody Rosaceae in western North America (Amelanchier, Crataegus, Photinia, Sorbus, and also on the non-native Pyrus communis and Mespilus germanica). Monoecious holocyclic, with oviparae and alate males appearing in September-October (BMNH collection, leg. C.-k. Chan). All morphs have been described, but the biology is little known. There is considerable karyotype variation both within and between samples; 2n (female) = 18, 19, 20, 21 and 23, and 2n (male) = 17 and 19, have all been found in samples from British Columbia, Canada (R.L. Blackman and C.-k. Chan, unpublished data). The biological and taxonomic significance of this variation are not yet understood.

Ericaphis harmstoni (Knowlton) Alatae only are described, BL 2.3-2.5 mm, on Symphoricarpos sp. in Washington, USA. True host and generic placement need to be confirmed.

Ericaphis latifrons (Börner) Apterae are yellow-green, with tips of antennae and legs black; BL 1.3-1.9 mm. Alatae have broad dark cross-bands more-or-less fused into a patch. On various Ericaceae, most usually Empetrum nigrum but also recorded from Calluna vulgaris, Erica umbellatum and Vaccinium uliginosum. In northern and western Europe, including Iceland, east to north-west Russia, and south to the Iberian peninsula. Oviparae and apterous males occur in August-October (earlier at more northerly latitudes). Heikinheimo (1990) gave an account of this species.

Ericaphis leclanti Remaudière Apterae are white with dark apices to antennal segments and legs; BL1.5-2.2 mm. Alatae have antennae mainly dark and siphunculi dark distally, but the dorsal abdomen has only lightly pigmented marginal and intersegmental sclerites. Colonies occur on young shoots of Arctostaphylos uva-ursi in the French Alps (Remaudière 1971). Alate viviparae, apterous males and oviparae are included in the original description, the sexual morphs appearing in early August. Possibly this species would be better placed in Wahlgreniella.

Ericaphis lilii (Mason) Colour of apterae in life is unrecorded; BL 1.8-2.3 mm. Described from Lilium candidum in Oregon, and also recorded from lilies in California, Washington and British Columbia (BMNH collection). This species has previously been confused with E. scopioli (q.v.).

Ericaphis louisae Pike Apterae are green with legs pale except for tarsi and distal parts of tibiae; BL 1.2-1.8 mm. Alatae are undescribed. Monoecious holocyclic on Luetkea pectinata in mountainous regions of north-west USA (Washington) and western Canada (British Columbia). Oviparae and apterous males were collected in mid-August to October (Pike & Starý 2003).

Ericaphis scammelli (Mason) Plate 22f Apterae are pale yellow-green; BL 1.5-2.4 mm. Alatae have dark brown dorsal abdominal markings partly fused into a central patch, with a conspicuous pale window between the siphunculi. On young shoots of various Ericaceae (Vaccinium, Gaultheria, Andromeda, Empetrum). In North America, and introduced to Europe (UK, France, Sweden, Netherlands, northern Italy, Serbia). A record from roses in Iran (Jalalizand et al. 2012) was based on identification by mitochondrial DNA and needs further confirmation. British populations were described as a subspecies of E. fimbriata, (pernettyae), and were shown by Prior (1971) to have a holocycle on Pernettya (= Gaultheria) mucronata, with oviparae and alate males in October-November. In Italy it is monoecious holocyclic on Vaccinium spp. (Barbagallo et al. 1999). 2n=14. [Note: Figure 294 in Blackman (2010) is labelled in error as E. scammelli; it is in fact a repeat of Figure 295 (E. wakibae).]

Ericaphis scoliopi (Essig) Apterae are pale yellow or whitish; BL c.2 mm. Alatae have a whitish to pale yellow abdomen with a large black dorsal patch. Described from leaves of lilies (Lilium spp. and Scoliopus bigelovi). In western USA (California, Oregon). Synonymy with E. lilii (Mason) has been previously suggested, but after examination of specimens from the Essig collection it is clear that the two species are distinct.

Ericaphis voegtlini Barjadze, Jensen & Kanturski Apterae in life are dark green; BL 1.3-1.9 mm. Alatae have 4-9 secondary rhinaria on ANT III and an extensive dark dorsal abdominal patch. On Chamaebatia foliolosa in California, USA (Barjadze et al. 2018b). Sexual morphs and life cycle are unknown.

Ericaphis wakibae (Hottes) (= Placoaphis siphunculata Richards) Plate 22g Apterae are pale brownish green or greenish yellow, with dorsal abdomen somewhat sclerotic and variably tanned, often with darker brown spinal and marginal regions, and siphunculi pale brown with pale areas around their bases; BL 1.6-2.8 mm. Alatae usually have an extensive dark dorsal abdominal patch with a clear window between the siphunculi. It was originally described from Pedicularus, but has since been recorded mainly from Rosaceae (Fragaria, Rosa, Rubus). This species is widely distributed in North America, has been introduced to the United Kingdom, and is now also reported from north-east Siberia (Chukotka), where fundatrices were collected on Dryas octopetala var. asiatica (Stekolshchikov & Khruleva 2015). It is monoecious holocyclic on Fragaria in England (Prior 1971). The life cycle in North America has not been studied, but possibly there is at least a partial migration to Ericaceae, as apterae have been collected on Menziesia ferruginea, Vaccinium alaskaense and Gaultheria shallon in British Columbia (BMNH collection, leg. C.-k. Chan). Secondary hosts in other families are reported from western USA (see aphidtrek.org). A similar species occurs on Chamaebatia foliosa in California (BMNH collection, leg. D. Hille Ris Lambers). 2n=12.

Ericolophium Tao

Aphidinae: Macrosiphini

About 13 east Asian species mostly associated with Rhododendron. One Indian species lives on Salix. Ericolophium is still poorly known and needs biological study as well as further systematic revision. Host alternation is suspected (Chakrabarti et al. 1983), as in the closely related genus Indomasonaphis, but no secondary hosts have been identified. Some nominal species differ from one another only in dorsal abdominal pigmentation, which may vary with temperature or season. Chaetomyzus and Indiaphis are also closely related, and may not be worthy of separate generic status.

Ericolophium alpigeniae Zhang, Zhong & Zhang Apterae are undescribed, alatae are green with a dark dorsal abdominal patch; BL c.3.8 mm. Alatae have secondary rhinaria distributed III 134-158, IV 1-6. On Rhododendron simsii in Yunnan Province, China. But for the slightly clavate siphunculi, which put its generic position in some doubt, the first tarsal chaetotaxy (5:5:5) and hairy R IV and cauda would put this species close to Ericolophium holsti.

Ericolophium dubium (Chakrabarti & Maity) Apterae are pale green; BL 2.3-2.8 mm. On undersides of leaves and young growth of Salix denticulata (= elegans), S. tetrasperma and Salix sp. in north-west India (Chakrabarti & Maity 1984, Medda et al. 1997, as Neoacyrthosiphon dubium). Living all-year-round on Salix, with oviparae and alate males occurring in October.

Ericolophium euryae (Takahashi) Apterae (according to the original description, from Eurya sp. in Taiwan) are yellow with black dorsal markings consisting of bands on head and pronotum, a large abdominal patch, a band behind and confluent with the siphunculi, and a patch reaching the end of the abdomen (Takahashi 1937b); BL 1.2-1.5 mm. There are also records from Eurya japonica in Java (BMNH collection, leg. P. van der Goot) and north-east India (A.K. Ghosh 1974, and BMNH collection, leg. D. Raychaudhuri) The BNMH specimens from Java lack pigmentation (possibly lost during slide preparation), and the single BMNH specimen from north-east India (West Bengal) differs from the original description by lacking the black posterior abdominal patch.

Ericolophium holsti (Takahashi) ( = Neomasonaphis rhododendris Chakrabarti, Mandal & Raha) Plate 23f Apterae are green with pale brownish siphunculi and a pale green cauda (original description); BL 2.4-3.5 mm. Alatae have secondary rhinaria distributed III 60-110, IV 0-12. On undersides of young leaves of Rhododendron spp. in Taiwan, India (Himachal Pradesh, Uttar Pradesh, West Bengal, Assam) and Pakistan (Naumann-Etienne & Remaudière 1995, as Indumasonaphis inulae; identification corrected by G. Remaudière, pers. comm., 27.ix.2004). Now in Europe, but so far only found in Britain, where alatae have been trapped several times in different parts of England from 2011-2014, and in 2017 a colony was found on Rhododendron at Kew (Eastop et al. 2012, Wieczorek et al. 2019). 2n=22 (Dutta & Gautam 1993).

Ericolophium itoe (Takahashi) Apterae are reddish brown with a greenish tint; BL 1.8-2.0 mm. Alatae have secondary rhinaria distributed III 30-38, IV 4-7, V 0(-2). On Rhododendron spp. in Taiwan, and Japan (Miyazaki 1971, as Elatobium itoe). 2n=18.

Ericolophium nigripunctatum (Zhang, Zhong & Zhang) Apterae are green with reddish head, thorax and cauda, and a dark dorsal abdominal patch; BL c.2.7-2.8 mm. Alatae have secondary rhinaria distributed III 34-41, IV 0. On Rhododendron simsii in Yunnan Province, China. Very similar to E. rectisiphon from the same locality and host plant .

Ericolophium ninguidum (Zhang, Chen, Zhong & Li) Apterae are green; BL c.2.3 mm. On undersides of leaves of Rhododendron przewalskii in Qinghai Province, China (Zhang 1999, as Neoacyrthosiphon).

Ericolophium ovalifolii (M.R. Ghosh, A.K. Ghosh & Raychaudhuri) Apterae are green; BL c.3.2 mm. Alatae are undescribed. On apical leaves of Lyonia (= Pieris) ovalifolia in West Bengal, India (M.R. Ghosh et al. 1971, as Neoacyrthosiphon taiheisanum ssp. ovalifolii), and also recorded from Rhododendron sp. in Meghalaya (Chakrabarti et al. 1983). We have given this taxon full species status.

Ericolophium rectisiphon (Zhang, Zhong & Zhang) Apterae are yellowish green or pale green, with darker green dorsal markings; BL c. 2.6 mm. Alatae are undescribed. On Rhododendron simsii in Yunnan Province, China. Very similar to E. nigripunctatum.

Ericolophium rhododendri (M.R. Ghosh, A.K. Ghosh & Raychaudhuri) (Fig. 46i,j) Apterae are dark green; BL c. 2.2 mm. Alatae have secondary rhinaria distributed c.70-72, IV 30-40 V c.12. On undersides of leaves of Rhododendron arboreum in West Bengal, and there is also a record from Nagaland on Leucosceptrum canum (Lamiaceae), a possible secondary host (Raha et al. 1977) .

Ericolophium taiheisanum (Takahashi) Apterae are blackish brown; BL c. 2.5 mm. Alate are undescribed. On upper sides of leaves, along midribs, of Rhododendron formosanum in Taiwan.

Ericolophium takahashii (A.K.Ghosh) Apterae are probably pale green; BL 2.0-2.5 mm. Described from one aptera and one immature alata collected on Anemone rivularis, but the true hosts are Rhododendron spp. (Chakrabarti et al. 1983, as Neoacyrthosiphon). In India (Uttar Pradesh, West Bengal) and Nepal. Monoecious holocyclic, with oviparae and alate males on Rh. campylocarpum in October (Chakrabarti & Raychaudhuri 1975).

Ericolophium tianchiense (Zhang, Chen, Zhong & Li) Apterae are green; BL 2.8-2.9 mm. On leaves of Rhododendron sp. in Xinjiang, China (Zhang 1999, as Neoacyrthosiphon).

Eriosoma Leach

Eriosomatinae: Eriosomatini

About 30 species, most of which have host alternation between galls on Ulmus and secondary hosts in Pyroideae, Grossulariaceae or Compositae/Asteraceae. Feeding by the immature fundatrix of a growing elm shoot in spring induces a galling reaction in one or more leaves distal to the feeding site, the fundatrix moving into the leaf-roll after it is formed (Akimoto 1981). Alatae and apterous exules of Eriosoma typically have rather conspicuous siphuncular pores with partially chitinized rims and surrounding hairs. Alatae usually have a once-branched media in the forewing, but specimens of E. yangi can have both forewings with unbranched media.

The limits of the genus are difficult to define; the east Asian Colophina and the North American Georgiaphis are often included as subgenera (Smith 1985, Eastop 1987), but are treated separately in this account. Sano & Akimoto (2011) made a cladistic study of relationships with other genera of the tribe Eriosomatini. Eriosoma s.str. comprises 4-5 North American species forming rosette leaf galls on Ulmus spp. of the americana group, with summer generations in woolly masses on Pyroideae, plus one species that is monoecious and free-living on Ulmus. Among the palaearctic species, Eastop (1987) distinguished two groups or subgenera; Schizoneura that typically form open, curled-leaf galls on Ulmus and then migrate to roots of Grossulariaceae and other plants, and Mimaphidus that typically form bloated-leaf galls on Ulmus before migrating to roots of Pyroideae, or in one case, Senecio. These two groups are approximately equivalent to the ulmi group and lanuginosum group discussed by Akimoto (1983), but there are some discrepancies. The life cycles of several species are still unknown. Areopraon lepelleyi is a specialised parasitoid of the elm-feeding generations.

Krikoanoecia circula Qiao & Zhang in G. Zhang et al. (1996), described from Quercus sp. in China, is based on a vagrant sexupara of E. ulmi or a related species, so that Krikoanoecia Zhang & Qiao 1996 becomes a new synonym of Eriosoma. Siciunguis Zhang & Qiao should also probably be treated as a synonym.

There are accounts or partial revisions of Eriosoma from Sweden (Danielsson 1979), Fennoscandia and Denmark (Heie 1980b), the UK (Blackman et al. 2019b), the Iberian peninsula (Nieto Nafría et al. 2002a), India (A.K. Ghosh 1984), east Siberia (Pashchenko 1988a), Japan (Akimoto 1983), China (G. Zhang & Zhang 1993, G. Zhang & Qiao 1997a,b, G. Zhang et al. 1999c)) and North America (Smith 1985). The species having Ribes as secondary host were reviewed by Danielsson (1982).

Eriosoma americanum (Riley) Spring colonies on Ulmus americana roll the edges of the leaves downwards (see influentialpoints.com/Gallery). Emigrant alatae (fig.118C) emerging from galls in June are dark green to blue-black with some wax wool, and rather dark wing veins; BL 2.0-2.5 mm. They migrate to form colonies of pale pink, woolly apterae on roots of Amelanchier. Return migrants to elm in August to October produce dwarf sexuales that lay overwintering eggs on bark (Patch 1915a, Palmer 1952). It occurs throughout the natural range of U. americana in eastern North America, and in the west on planted elms.

Eriosoma (Schizoneura) anncharlotteae Danielsson Galls formed on Ulmus (minor, procera) by inward curling of both edges of leaves, which become blistered, yellow or light green,, or sometimes bright red on younger trees. and often considerably distorted (Danielsson 1979). Fundatrices dark green to bluish grey covered in wax meal. Alatae mainly produced in third generation are dark bluish grey, BL 1.8-2.3 mm, migrating in June-July to roots of Ribes spp. (alpinum, sanguineum). Apterous exules are yellow to pale red in white wax wool; BL 1.1-1.9 mm. Sexuparae, with secondary rhinaria distributed III 25-33, IV 4-6, fly back to elm bark from mid-September. In Europe (Czech Republic, England, Germany, Poland, Sweden, Switzerland), and also recorded from China (G. Zhang et al. 1985).

Eriosoma antennieurfum Zhang Described from one alata (sexupara?) collected from roots of Helianthus annuus in China (Zhang & Qiao 1997b).

Eriosoma (Schizoneura) auratum Akimoto Forms a leaf-roll gall on Ulmus japonica; one edge of the leaf is rolled tightly downward, and becomes yellowish green and stiff (Akimoto 1983). The fundatrix is greyish green. Almost all second generation are alatae, brown, BL 1.5-1.9 mm, migrating in June to found colonies on roots of Anthriscus sylvestris (Akimoto 1985). In Japan and Korea, and also now recorded from east Siberia and Kazakhstan (on U. pumila; Kadyrbekov 2017a). Gall invasion and competition with other Eriosoma on Japanese elm were studied by Akimoto (1988, 1989). Apterous exules and sexuparae are undescribed, although Tong et al. (2021) studied a mermithid nematode that parasitises alate sexuparae. E. eligulatum Pashchenko (1988), described from U. japonica and U. pumila in east Siberia, is probably a synonym, and neither E. togrogum Zhang nor E. usuense Zhang, both described from alatae fundatrigeniae from U. pumila in China (G. Zhang & Qiao 1997a), are clearly distinct from E. auratum on the basis of the published descriptions. 2n=12.

Eriosoma crataegi (Oestlund) Apterae on bark of twigs and stems of Crataegus spp. are bluish-black covered in white mealy wax, and often have two long wax filaments extending from the end of the abdomen; BL 2.0-3.0 mm. Widely distributed in North America. Crataegus is the usual host, but aphids apparently of this species are also sometimes collected on bark of Fagus, in colonies of Grylloprociphilus imbricator (Smith 1985), and it is also recorded from Pyracantha sp. Smith distinguished several forms of apterous exules on Crataegus, differing in dimensions of wax glands. Sexuparae fly to Ulmus (americana, fulva) and produce sexuales on elm bark in September-October. Heriot (1938) described differences in the sexuparae, males, oviparae and eggs between E. americana, E. lanigerum and E. crataegi. Leaf rosette galls similar to those of E. lanigerum are presumably formed on Ulmus in spring, but strangely, despite the biological studies of Heriot (1938) and Cox (1939), the spring generations of crataegi on elm have not been specifically identified; separation of crataegi in the Ulmus key is based on data provided by Cox for spring migrant alatae collected on Crataegus in May-June. 2n=12.

Eriosoma (Mimaphidus) flavum Jancke (= E. gomboriense) Forming leaf-roll galls on Ulmus spp. (minor, glabra, procera); the edge of the leaf is curled and swollen near its base (fig. 134N and Dzhidladze 1965a). Alatae, BL 1.4-1.7 mm, emerge in mid July-August. Summer generations are on the roots of Pyrus communis and Cydonia oblonga, often deep underground, the apterous exules being very small (BL 1.0-1.2 mm) and yellowish (cf. pyricola), with filamentous wax. In Europe (England, Netherlands), Israel (exules on Cydonia), Iran (BMNH collection, leg. S.H. Hodjat) and Georgia (as gomboriense; Dzhibladze 1965a).

Eriosoma (Schizoneura) grossulariae (Schüle) Forming open, green to yellow curled-leaf galls on Ulmus spp. (fig. 134L), similar to those of E. ulmi, with which this species was confused for many years (see Danielsson 1982). Immature alatae in galls are light green (cf. ulmi), maturing in June; dark green when adult, migrating to found colonies on roots of Ribes sanguineum and R. uvacrispa. Apterous exules are pale red to yellowish white in white wax wool; BL 1.0-2.3 mm. Alate sexuparae in autumn bluish grey to black, with secondary rhinaria distributed III 15-24, IV 2-5. Throughout Europe, Japan, and also introduced on European elms into North America. 2n=10.

Eriosoma (Schizoneura) harunire Akimoto Leaf-roll galls on Ulmus japonica are formed by rolling one margin of the leaf near its tip firmly towards the underside, and remain green and unswollen even when mature (Akimoto 1981). Fundatrix is brownish or greenish grey, and produces both apterous and alate fundatrigeniae. Alatae (fig.118G) mature in the galls in July, later than E. auratum (Akimoto 1989), and migrate to found colonies on roots of Plantago asiatica (Akimoto 1985b). Apterous exules and sexuparae are undescribed. Only known from Japan. Gall invasion and competition with other Eriosoma were studied by Akimoto (1988, 1989). 2n=10.

Eriosoma (Schizoneura) japonicum Matsumura (= Schizoneurella gei Bozhko; synonym by Eastop & Blackman 2005) Leaf-roll galls on Ulmus japonica are indistinguishable from those of E. harunire (see above). Fundatrix reddish brown. Alatae emerge in June and migrate to found colonies on roots of Rosaceae (Agrimonia pilosa, Geum japonicum, Fragaria ananassa; Akimoto 1983, 1985). Apterous exules are oval, with dark head and appendages, secreting flocculent wax; BL 1.5-1.7 mm. In Japan, China, east Siberia and Taiwan (Tao 1970; as E. ulmosedens). 2n=10.

Eriosoma (Schizoneura) kashmiricum L.K. Ghosh, Verma & Raychaudhuri Forming leaf-roll galls on Ulmus villosa and U. wallichiana in India (Kashmir, Uttar Pradesh) and Pakistan (Murree hills); the leaf margin is folded downwards and twisted into a spiral (A.K. Ghosh et al. 1981). A record from ?Alnus sp. can presumably be discounted. The fundatrix is dark brown (Bhattacharya et al. 1980). Alatae are produced in June and fly to an unknown secondary host. 2n=12 (Pal & Khuda-Bukhsh 1983).

Eriosoma (?Schizoneura) laciniatae Pashchenko Forming curled-leaf galls on Ulmus laciniata in Siberia (Pashchenko 1988a). Alatae are produced in July, secondary host (if any) unknown. Appears closely related to grossulariae or longicornutum, but lacks a ciliary fringe to the primary rhinarium on ANT V. Some fundatrices collected from yellow curled leaves of U. laciniata in China were possibly of this species (BMNH collection, leg. VFE, No. 18,182); these specimens had 2n=16*, different from any other known karyotypes of Schizoneura.

Eriosoma lanigerum (Hausmann) Woolly Apple Aphid Apterae on Pyroideae are purple, red or brown, covered with thick white flocculent wax (see influentialpoints.com/Gallery); BL 1.2-2.6 mm. On roots, trunk or branches, often causing deformation and cancer-like swellings of bark. It is a severe pest of apple (eg. Weber & Brown 1988), and may also be found on Cotoneaster and Pyracantha, and more rarely on Cydonia, Crataegus, Pyrus and Sorbus, but is not so injurious to these plants. Anholocyclic populations, sometimes with an apparently abortive sexual phase, occur on apple throughout the world, overwintering on the roots (see Blackman & Eastop 2000, p. 271). Alate sexuparae producing oviparae and males on apple are reported from various parts of the world, and eggs may be laid on apple leaves (e.g. Fotedar & Kapur 1943, Asante 1994, Sandanayaka & Bus 2005), but there is no firm evidence that these are capable of hatching, and it unlikely that fundatrices would be able to develop on apple and complete the holocycle.

Siciunguis decima Zhang & Qiao, described from apterae and alatae (sexuparae?) and causing heavy damage to apple in China (G. Zhang et al. 1999b) seems likely to be a synonym.

In North America, aphids of the E. lanigerum group induce leaf rosette galls on Ulmus americana, but there is still doubt about the relationship of these to apple-feeding populations. Smith (1985) successfully transferred alatae from such galls to apple. However, alatae indistinguishable from those of E. lanigerum were obtained from reddish pink leaf curls on U. alata, and these would not transfer to apple but successfully colonised Crataegus uniflora. Similar aphids occur in leaf-cluster galls on sterile elms in Australia (Fisk et al. 1992).

The ecology and natural enemies of E, lanigerum on apple were studied by Bouchard et al. (1984; Quebec, Canada), Gontijo et al. (2012; Washington, USA), Thakur et al. (1988; India) and von Kogler (1989; Germany). Aphelinus mali is a widespread specialised parasitoid (but Aphelinus gossypii, which parasitises a wide range of hosts including Aphis and Rhopalosiphum spp., have often been misidentified as A. mali). Lavandero et al. (2011) studied genetic variability in the aphid and its parasitoid in Chile, Timm et al. (2005) studied its genetic diversity in South Africa, and Zhou et al. (2015) analysed the genetic diversity and structure of the populations introduced into China. Ruiz-Montoya et al. (2015) studied morphometric and allozyme variation in populations collected on Pyracantha koidzumii in Mexico. A study of predation by earwigs was made by Mueller et al. (1988). Costa et al. (2014) compared performance parameters of four clones of E. lanigerum in Australia. Davidson (1913) made a detailed study of morphology and anatomy. M.Su et al. (2016) studied the distribution and functioning of the wax gland pores. The complete mitochondrial genome has been published (Y. Wang et al. 2016a). See Molinari (1986) for a general review. 2n=12.

Eriosoma (Mimaphidus) lanuginosum (Hartig) Forms clusters of large, closed, bloated-leaf galls near ends of branches of various Ulmus spp. The gall is produced by extreme hypertrophy of the leaf parenchyma on one side of the mid-rib near its base (fig. 134M); the hypertrophied tissue is light green at first and covered with fine white hairs, becoming brown as the gall matures (Marchal 1933). Fundatrices are blackish, wax-powdered, apterous fundatrigeniae are dark green with black legs. Alatae produced in the second and third generations on elm are dark green to black, wax powdered; BL 2.1-3.1 mm (Plate 5e). They emerge from the galls in late June-July and migrate to found colonies on fibrous rootlets of Pyrus communis or Cydonia. Apterae on pear roots are variable in colour, pale yellow to reddish, BL 2.0-2.7 mm (Plate 1b). Return migration to elm bark occurs in September. Occurs throughout Europe, the Mediterranean area and Asia east to Pakistan, introduced to South Africa. Often confused with E. pyricola, a smaller species with more caudal hairs which has a much wider geographical distribution (de Fluiter 1933, Blackman & Eastop 2000, p. 272). 2n=10.

Eriosoma (Schizoneura) lishanense Akimoto Leaf-roll galls are formed on Ulmus uyematsui in mountains of Taiwan. The fundatrix has 5-segmented antennae. Emigrant alatae are similar to those of E. auratum, with secondary rhinaria distributed III 23-26, IV 5, V 0, VI 0, and BL 1.4-1.5 mm (Akimoto 1983). It was subsequently also recorded from northern China (G. Zhang & W. Zhang 1993). The life cycle and secondary host are unknown.

Eriosoma (Schizoneura) longicornutum Akimoto Forms open leaf-roll galls on Ulmus japonica and U. laciniata in Japan; the leaf edge on one side is rolled downwards and becomes inflated and yellowish-green. The fundatrix is yellowish-green with dark brown legs and antennae. Alatae produced in June-July migrate to an unknown host. Closely related to E. grossulariae, but with a difference in the wax glands of first instar exules (Akimoto 1983). 2n=10.

Eriosoma mimicum Hottes & Frison On Ulmus spp. (americana, rubra) in eastern North America, probably forming leaf-curl galls. Alatae were collected in June, migrating to an unknown host (Hottes & Frison 1931, Smith 1985). Georgiaphis maxsoni is probably a synonym.

Eriosoma (Schizoneura) moriokense Akimoto Leaf-curl galls are formed on Ulmus japonica; often whole leaves are strongly twisted, becoming reddish or brownish and dusted with wax powder when the gall is mature (Akimoto 1983 and see http://www.agr.hokudai.ac.jp/systent/akimoto/gall/E.%20moriokense%20gall.jpeg). The fundatrix is reddish brown, and produces both apterous and alate fundatrigeniae. Alatae (BL 1.6-2.2 mm) (fig.118F) leave galls in late June and found colonies on roots of Sedum spp. (Akimoto 1985). Apterous exules, presumably secreting wax, have BL c.1.3 mm. Sexuparae return to the trunks of elm in September. Pashchenko (1988) records it from U. pumila in east Siberia, and G. Zhang & W. Zhang (1993) record it from U. davidiana and U. macrocarpa in northern China. In Japan, China, east Siberia, and probably Korea, as according to Akimoto (1983), Eriosoma sp. B of Paik (1972) is probably E. moriokense. Akimoto (1983) recorded evidence of aggressive behaviour towards predators by second and third instar larvae in galls. 2n=10.

Eriosoma nigra Akimoto Apterae are oval, dark-pigmented, secreting wax; BL 1.5-1.8 mm. On roots of Epilobium sp. in Japan (Akimoto 1983). Apparently anholocyclic on Epilobium; alatae found in September contained embryos with mouthparts (i.e., were not sexuparae).

Eriosoma (Mimaphidus) patchiae (Börner & Blunck) Galls consist of curled, twisted and blistered leaves on shoots of Ulmus spp. (glabra, minor, procera, pumila), stunting and twisting young growth, often on suckers arising from the trunk (Marchal 1933). Galls green or yellowish, fundatrices yellowish green, apterous fundatrigeniae whitish green, according to Marchal (1933); but the fundatrix is dark bluish grey according to Danielsson (1979). Alatae produced in the second and subsequent generations are green or brownish, with narrow brown dorsal abdominal cross-bands; BL 2.0-2.5 mm. Migration occurs in June-July to roots of Senecio, or Cineraria, with return to elm in September-October. Apterous exules, secreting wax, have BL 2.0-2.3 mm. However, the host alternation seems to be facultative, because alatae emerging from galls later in July-August are often sexuparae. Possibly a distinct, monecious species in involved, for which the name ulmosedens Marchal 1919 is available, but according to Marchal (1933) some of the alatae from galls can give birth to both sexuales and apterous exules. In north-west and central Europe; Sicily (Barbagallo & Stroyan, 1982); Madeira, where no Ulmus spp. occur and it is presumably anholocyclic on roots of Senecio (Aguiar et al. 2013); western Siberia, and Kazakhstan. Parker (1984) studied the association of E. patchiae with Anthocoris gallarum-ulmi, and Dessart & Gardenfors (1985) studied parasitism and hyperparasitism. 2n=10.

Eriosoma (Schizoneura) phaenax (Mordvilko) Leaf galls on Ulmus (glabra, minor, wallichiana) appear to be similar to those of patchiae; the illustration of a gall-cluster on U. campestris (= minor) by Mordvilko (1924, Pl. IV) ascribed to patchiae and copied by Shaposhnikov (1964) and Heie (1980), applies to phaenax. The simple marginal leaf-roll described for E. phaenax by A.K. Ghosh et al. (1981) was perhaps a young gall. Apterous fundatrigeniae in galls are light brown (A.K. Ghosh 1984). Emigrant alatae emerging from galls in June-July are brownish with narrow darker brown dorsal abdominal cross-bands; BL 2.1-2.5 mm. Apterous exules have not been described, but sexuparae have been collected in October on Cotoneaster in Pakistan (BMNH collection, leg. R.H. Ghani) and Kashmir, India (BMNH collection, leg. Rishi), and this is probably the secondary host; sexuparae collected on Androchne cordifolia (Euphorbiaceae) in Pakistan (A.K. Ghosh 1984) were probably vagrants. Mordvilko (1935), however, noted that E. phaenax was monoecious, so possibly host alternation is facultative, as in E. patchiae. In Transcaucasus, Turkmenistan, Tajikistan, Afghanistan, Pakistan and India. This species was at one time erroneously synonymised with patchiae (see Marchal 1933, Narzikulov 1965a).

Eriosoma (Mimaphidus) pyricola Baker & Davidson Galls on Ulmus spp. (americana, procera) are clusters of bloated leaf tissue like those of E. lanuginosum (see Alfieri 1920; as inopinatum). The fundatrix and apterous fundatrigeniae are grey, emigrant alatae produced in second and third generations are brown with pale yellow abdomen (cf. lanuginosum); BL 1.3-2.4 mm. Migration occurs in June-July to fibrous rootlets of Pyrus communis or Cydonia. Apterous exules are yellowish pink when immature, pink to red when adult, rather sparsely clothed with filamentous wax (Baker & Davidson 1916); BL 1.3-2.0 mm. Sexuparae are shiny dark green to brown, and return to elm bark in September-October. In southern Europe (Italy, Yugoslavia), Turkey, and introduced into south-east Australia, Tasmania, New Zealand, USA (western and some north-eastern states), Canada (British Columbia), South America (Argentina, Chile) and (probably) South Africa. Swenson (1971) studied the influence of the secondary host on sexupara production. This aphid is often confused with E. lanuginosum in the literature. E. alabastrum Pashchenko (1988), described from U. japonica in Siberia, is very similar and possibly a synonym. It also seems likely that E. dilanuginosum Zhang (1980b), described from U. pumila in China, and with all morphs including the apterous exules on Pyrus now described (Qiao et al. 1999a; and again by Qiao et al. 2001b under the name Siciunguis novena), is this species. If this is so, then a record of E. dilanuginosum from Kazakhstan (Kadyrbekov 2018a) may also be ascribable to E. pyricola.

Eriosoma rileyi Thomas On bark of trunk and branches of small elms (Ulmus americana, rubra) in North America, forming small colonies covered with wax wool very like those of E. lanigerum on apple, and causing similar deformation of woody tissue. It lives all-year-round on elm, without forming leaf-galls. Apterae under the wax are pinkish, yellow or brown; BL 1.8-2.3 mm. Alatae produced in June to August have ANT III 0.8-1.0 × head width across (and including) eyes, with secondary rhinaria rather widely spaced. Sexuparae are apterous, produced in September-October, depositing dwarf sexuales directly onto bark (Patch 1913, Palmer 1952).

Eriosoma (Schizoneura) sorbiradicis Danielsson Apterae are yellowish white, with sparse wax; BL 0.7-1.4 mm. They live singly or in small colonies on rootlets of Sorbus aucuparia (Danielsson 1979). Recorded from Scotland (Stroyan 1991), Sweden (Danielsson) and Oregon USA (BMNH collection; leg. R.L. Furniss). Other morphs and life cycle are unknown.

Eriosoma (Schizoneura) ulmi (Linnaeus) Galls on Ulmus spp. are formed by downward curling, twisting and blistering of one lateral edge of a leaf, the deformed part becoming yellowish or whitish green. Fundatrix and apterous fundatrigeniae are wax-covered, dark green, immature alatae are brownish or dull green (cf. grossulariae; and see influentialpoints.com/Gallery). Adult alatae are dark green to bluish grey, migrating in June-July to found colonies on roots of Ribes spp. (especially nigrum, rubrum, aureum). Marchal (1933) recorded it from Sedum sp. in France, and it is also reported to occur sporadically on roots of Vitis (Börner 1952), but these could possibly be other hitherto unrecognised species. Apterous exules are pale red to brownish red, in wax wool; BL 0.8-2.0 mm. Return migration to elm bark is in September-November; alate sexuparae are dark bluish grey to black, with secondary rhinaria distributed III 14-23, IV 1-3. In Europe (including Iceland and Faroes), south-west and central Asia, Pakistan, India, eastward to Mongolia and China according to Heie (1980b), and more recently introduced to British Columbia, Canada (Foottit et al. 2006). E. mediocornutum Pashchenko (1988), described from Ulmus japonica and U. pumila in Siberia, is very similar and possibly a synonym. Parker (1984) studied predation by Anthocoris gallarum-ulmi, and Dessart & Gardenfors (1985) studied parasitism and hyperparasitism. A detailed study was made of its biology and natural enemies on elm in the Czech Republic by Urban (2003b), and the histology of the gall was studied by Álvarez et al. (2013). 2n=10 (Europe), but Chen & Zhang (1985a) recorded 2n=16 from China, possibly having another, closely-related species.

Eriosoma (Schizoneura) ulmipumilae Ivanoskaya In curled-leaf galls on Ulmus pumila in the Altai Region of Siberia (Ivanoskaya 1976), and also recorded from Kazakhstan (Kadyrbekov 2005c) and Uzbekistan (Kadyrbekov 2013c). The life cycle is unknown. Very similar to E. japonicum or E. yangi. The alate fundatrigenia described as E. cerum Zhang (in G. Zhang & Qiao 1997a) is also not clearly distinct.

Eriosoma ulmipumilicola Zhang Only the apterous fundatrigenia is described, from an undescribed gall on Ulmus pumila in China (Hebei; Zhang & Zhang 1993). Wax glands are apparently very well developed on head and abdominal segments. Two other Eriosoma species were subsequently described from apterous fundatrigeniae on U. pumila (E. fukangense Zhang and E. spirifolium Zhang in G. Zhang & Qiao 1997a), also with well developed wax glands, and neither clearly distinct from E. ulmipumilicola.

Eriosoma (Schizoneura) yangi Takahashi Forms yellowish green or green rolled-leaf galls on Ulmus parvifolia in Japan, China and Korea (Akimoto 1983, and also see http://www.agr.hokudai.ac.jp/systent/akimoto/gall/yadori.jpg). Alatae from galls in May-June have BL 1.4-1.7 mm, and migrate to found colonies on roots of Salix sp. and Fragaria ananassa (Akimoto 1985). Apterous exules are undescribed. A closely-related form regarded as a subspecies, E. yangi parasiticum, occurs on Ulmus japonica in Japan and is an obligatory “cuckoo” parasite of other Eriosoma species, invading and usurping their galls; Akimoto (1988a,b) made a detailed study of this phenomenon and compared morphology and behaviour of fundatrices of the two forms. E. multilocularis Zhang, described from an unidentified Ulmus sp. in China (G. Zhang & W. Zhang 1993), is said to be similar to E. yangi s. str., but has multilocular galls. 2n=10.

Essigella Del Guercio

Lachninae: Eulachnini

18 nearctic species, all living on Pinus needles except one species (wilsoni) specific to Pseudotsuga. They are long-bodied aphids, as befitting their needle-feeding habit, and are the New World equivalent of Eulachnus, but smaller and with 5-segmented antennae. Apterae are spindle-shaped, with tapered abdomen, but the “spindle” is truncated at the head end, which is relatively broad. They are often overlooked, as they feed near the bases of the needles and move quickly when disturbed; their biology is consequently little known. Sorenson (1988, 1992, 1994) revised Essigella, synonymising many of the species described by Hottes (1957c,1958f). Théry et al. (2018a) provided a review of the genus, a simplified key to species and an updated checklist, differentiating three new species in the E. californica group specific to particular pine species, largely on the basis of molecular differences. Théry et al. (2018b) provided a molecular phylogenetic analysis combining data from mitochondrial, nuclear and endosymbiont DNA and discussed the problems of species delimitation in this genus. Diaretus essigellae is a specialised parasitoid.

Essigella alyeska Sorenson nsp; Apterae are spindle-shaped, grey-green with yellow-orange head and mainly pale appendages; BL 1.4-1.7 mm. Feeding singly on needles of Pinus banksiana in Canada (Ontario, Quebec), and also collected on Picea glauca in Alaska (Sorenson 1988). ABD TERG 2-4 have 8 spinopleural and 2 marginal hairs, and ABD TERG 8 has 6-8 hairs. Alatae have 0-2 secondary rhinaria on ANT III. Other morphs unknown.

Essigella braggi Hottes Apterae are spindle-shaped, grey-green, or grey to light brown; BL 2.2-2.5 mm. On needles of Pinus contorta (var. murrayana) in Sierra Nevada and Cascades of California USA (Hottes 1957c, Sorenson 1994). Closely related to E. knowltoni, and treated as a subspecies of knowltoni by Sorenson (1994).

Essigella californica (Essig) Plate 16c Apterae are spindle-shaped, with grey-green thorax and lime green abdomen, with or without brown dorsal spots (see influentialpoints.com/Gallery); BL 1.5-2.0 mm. Legs variably pigmented, often mainly pale, tibiae sometimes darker. Alatae have 2-4 secondary rhinaria on ANT III. Feeding singly on needles of Pinus spp. of subsections Cembrae, Strobi, Ponderosae, Sabinianae, Oocarpae, and several introduced Palaearctic pines, but not on Pinus albicaulis, P. monticola, or P. muricata, which have cryptic species of the californica group specific to them (respectively E. domenechi, E. gagnonae and E. sorenseni; see separate accounts of these species below, and Théry et al. 2018a,b). Rarely found on pines of subsections Contortae or Cembroides, and records from such plants are quite likely to be misidentifications of other species. It also sometimes occurs on Pseudotsuga (menziesii, macrocarpa). In western North America from southern British Columbia and Alberta, south to Mexico, and introduced into Brazil (de Carvalho & Lazzari 2000); Argentina (Ortego & Mier Durante 2012); Europe (France,Turpeau & Remaudière 1990; Spain, Seco Fernandez & Mier Durante 1992; Madeira, Aguiar & Ilharco 2001; Italy, Barbagallo et al. 2005b; and most recently found in southern England (Reid et al. 2015, and see influentialpoints.com/Gallery); China (Théry et al. 2017); Australia (Carver & Kent 2000); New Zealand (Flynn et al. 2003); and probably also in North Africa (Tunisia; Boukhris-Bouhachem et al. 2007, as Essigella sp.). A record from Malta (Mifsud et al. 2009) was based on a misidentification (Théry et al. 2017). DNA sequence studies have shown that introduced populations of E. californica have originated from at least four source populations (Théry et al. 2017), and shown that the name has been applied to a group of cryptic species in North America living on different Pinus species (Théry et al. 2018a,b). Outbreaks in Pinus radiata plantations in Australia have caused needle yellowing, premature senescence and defoliation (May & Carlyle 2003), leading to research programmes for biological control (Kimber et al. 2010) and development of resistant varieties of this host plant (Sasse et al. 2009). Oviparae and apterous males have been recorded in California (Sorensen 1994), but not elsewhere, even in colder climates. Wharton et al. (2004) studied development and showed that there are only three larval instars. The natural enemy complex in California was examined by Triapitsyn et al. (2015). 2n=8.

Essigella critchfieldi Sorensen Apterae are spindle-shaped,black, or rarely green with black dorsal spots; BL 1.6-1.9 mm. Feeding on needles of Pinus contorta s. str. (with one record from var. latifolia) in coastal north-western Canada and USA as far south as northern California (Sorensen 1994). Other morphs are unknown.

Essigella domenechi Théry, Kanturski & Favret Apterae are spindle-shaped, appearance in life unknown, probably very similar to E. californica; BL 1.6-2.1 mm. Living at a high elevation on Pinus albicaulis in California, USA, and probably in other montane localitions where this pine species occurs (Théry et al. 2018a). Other morphs are unknown.

Essigella eastopi Sorensen Apterae are spindle-shaped, varying in colour from wholly yellow to dark brown with yellow on front of head and a yellow spinal stripe on thorax and abdomen; BL 1.6-2.0 mm. On needles of Pinus coulteri in coastal ranges of California, south of San Francisco Bay, to Mexico (Sorensen 1994). Other morphs are unknown. A member of the E. fusca group, and difficult to distinguish from that species (Théry et al. 2018b).

Essigella essigi Hottes Apterae are spindle-shaped, black or dark green throughout, or with yellow-green head and green thorax and abdomen, often with dark dorsal spots; BL 1.6-1.8 mm. Alatae have 2-3 secondary rhinaria on ANT III and 0-1 on IV. On needles of Pinus spp. of subsection Oocarpae (attenuata, radiata), and rarely on subsection Ponderosae. In south-west Oregon and California. Sexual morphs are unknown.

Essigella fusca Gillette & Palmer Apterae spindle-shaped, head and thorax dusky yellow-brown, abdomen green with brown spots; BL 2.0-2.6 mm. Feeding singly on needles of Pinus spp. of subsection Ponderosae, especially P. ponderosa; also recorded from P. coulteri and P. leiophylla, and collected once from Callitris drummondii (BMNH collection, leg. H.G. Walker). In southern British Columbia, Canada, and south through Rocky Mountains to Mexico. Apterous males and oviparae occur in September-October (Palmer 1952). Populations in California and south-west Oregon are regarded as a distinct subspecies (E. fusca voegtlini) by Sorenson (1994). E. eastopi and E. hillerislambersi are closely related (Théry et al. 2018b).

Essigella gagnonae Théry, Kanturski & Favret Apterae are spindle-shaped, appearance in life unknown, probably very similar to E. californica; BL 1.6-2.0 mm. Living at higher elevations on Pinus monticola in western USA (Washington, California, Nevada; Théry et al. 2018a). Other morphs are unknown.

Essigella hillerislambersi Sorensen Apterae are spindle-shaped, straw yellow, often with dark spots, with head yellow to orange-brown; BL 2.0-2.7 mm. Alatae have 0-2 secondary rhinaria on ANT III, 0-1 on IV. On needles of Pinus jeffreyi in south-west Oregon and California USA (Sorensen 1994), and also recorded from Pinus sp. in New Mexico (Miller et al. 2016). Closely related to E. fusca. Oviparae occur in September, males are undescribed.

Essigella hoerneri Gillette & Palmer Apterae spindle-shaped, head and thorax grey-green, abdomen lime green, legs yellow-brown; BL 1.4-2.0 mm. On needles of Pinus spp. of subsection Cembroides (pinyon pines) in south-western USA; a common species, and the only Essigella to preferentially colonise pinyon pines (Sorenson 1994). Oviparae occur in September-November. Closely related to, and difficult to distinguish from, E. californica, with which it has frequently been confused in the literature.

Essigella kathleenae Sorenson Apterae narrowly spindle-shaped, wholly pale yellow; BL 1.3-2.0 mm. On needles of Pinus lambertiana in south-west Oregon and California USA (Sorenson 1988). Oviparae have been collected in September. Other morphs are unknown.

Essigella kirki Sorenson Apterae narrowly spindle-shaped, grey-green or occasionally pale yellow; BL 1.7-2.1 mm. On needles of Pinus flexilis and P. strobifomis in the Rocky Mountains from Montana to Arizona and New Mexico, and in the southern Sierra Nevada and White Mountains of California USA (Sorenson 1988). Oviparae collected in September, other morphs unknown.

Essigella knowltoni Hottes Apterae spindle-shaped, usually dark brown to black with front of head yellowish, sometimes grey-green; BL 1.8-2.1 mm. Alatae have 0-5 secondary rhinaria on ANT III and 0 on IV. On needles of Pinus contorta (var. latifoliae and var. murrayanae) in western Canada (British Columbia, Alberta), and in interior parts of Washington and Oregon, south through the Rocky Mountains to Utah and S Colorado (Hottes 1957c, Sorenson 1992). Oviparae were collected in August. E. braggi on P. contorta contorta in California is closely related, and was regarded as a subspecies of knowltoni by Sorenson (1994).

Essigella patchae Hottes Apterae are spindle-shaped, green; BL c. 1.8 mm. Alatae have 3-4 small secondary rhinaria on ANT III only. On needles of Pinus strobus in eastern USA (Maine) and eastern Canada (Quebec). Oviparae collected on P. strobus in North Carolina in October (Sorensen 1994, as E. pini) should perhaps also be assigned to this species, which was for a long time synonymised with E. pini (see Théry et al. 2018a,b).

Essigella pini (Wilson) Apterae spindle-shaped, mainly green to greenish yellow with yellow-orange to reddish orange head, and with rows of brown dorsal spots on abdomen (Wilson 1919); BL 1.5-2.0 mm. Alatae have 2-3 secondary rhinaria on ANT III, none on IV. On needles of Pinus spp. of subsection Australes (taeda, virginiana) in eastern USA, with one record from Canada (southern Quebec). Records from other hosts and from western North America (e.g. Palmer 1952) are based on misidentifications. Oviparae collected on P. strobus in North Carolina in October (Sorensen 1994) should possibly be assigned to E. patchae (q.v.). Males are unknown. Patti & Fox (1981) studied the seasonal occurrence of E. pini on P. taeda in South Carolina, where viviparae are present during the winter months.

Essigella sorenseni Théry, Kanturski & Favret Apterae are spindle-shaped, appearance in life unknown, probably very similar to E. californica; BL 1.9-2.3 mm. Living on Pinus muricata in coastal locations in California USA (Théry et al. 2018). Other morphs are unknown.

Essigella wilsoni Hottes Apterae are spindle-shaped, wholly lime green; BL 1.1-1.8 mm. Alata (1 specimen) has 1 secondary rhinarium on ANT III only. On needles of Pseudotsuga spp. (menziesii, macrocarpa) in western Canada (British Columbia, Alberta) and in western USA south to Mexico. Not recorded from Pinus. Oviparae were found in Oregon in August (Sorensen 1994), males are unknown.

Eucallipterus Schouteden

Calaphidinae: Panaphidini

Active, delicate insects on Tiliaceae, with all viviparae alate. The anal plate is deeply cleft. Parasitoids specialising on Eucallipterus and related genera such as Chromaphis, Myzocallis, Tinocallis and Tuberculatus are: Trioxys pallidus (emerging from pale mummies), Praon flavinode (forming “tents” under the dead aphids), and Aphelinus subflavescens (emerging from black mummies).

Eucallipterus tiliae (Linnaeus) Plate 9f Alatae are pale yellow with black markings, including lateral stripes on head and prothorax and two rows of black dorsal abdominal spots (see influentialpoints.com/Gallery); BL 1.8-3.0 mm. On undersides of leaves of Tilia spp. in Europe, south-west and central Asia, north Africa, and introduced to North America and New Zealand. Monoecious holocyclic; apterous oviparae and alate males occur in late August to October. Numerous aspects of the biology of E, tiliae have been studied including population dynamics (Dixon 1971, Barlow & Dixon 1980, Heimbach 1986); aggregation behaviour (Kidd 1976); flight behaviour (Kidd 1977); sexual morph production (Dixon 1972); effect on tree growth (Dixon 1971, Llewellyn 1972); susceptibility of different Tilia species (Carter & Nichols 1986); interactions with predators in UK (Glen & Barlow 1980, Wratten 1973); and parasitoids in USA (Hajek 1986). Biocontrol has been used in California USA (Zuparko 1983). 2n=10 (female), 2n=8 (Male); the female has 2 pairs of X chromosomes, and sex determination is X₁X₁X₂X₂/ X₁X₂0 (Blackman & Eastop 1994).

[DNA barcoding has revealed that under the name E. tiliae there may in fact be at least two cryptic species, differing in relative lengths of antennal segments and HT II (Y. Lee et al. 2017). There may also be at least two other species in this genus: (1) Specimens with a slightly longer ultimate rostral segment from Tilia tomentosa in Greece, Turkey and Iran may represent an undescribed species. (2) Shinji (1933) described Therioaphis tilicola from Tilia japonica and T. miqueliana in Japan. This species has not since been recognized (Higuchi 1972, Quednau 2003); it seems more likely to belong in Eucallipterus, rather than in the east Asian Tilia-feeding genus Tiliaphis. Qiao et al. (2005b) described an ovipara under this name from T. mongolica in China.]

Eucarazzia del Guercio

Aphidinae: Macrosiphini

One distinctive species with markedly clavate siphunculi associated with Labiatae/Lamiaceae, alatae having distinctive markings on wings and body.

Eucarazzia elegans (Ferrari) Plate 21c (Fig.48a) Apterae are pale green; BL 1.4-2.1 mm. On undersides of leaves, shoots, and flowers of Mentha spp. and various other Labiatae/Lamiaceae (Salvia, Coleus, Lavandula, Melissa, Nepeta, Origanum, etc.). Alatae have extensive and distinctive black dorsal abdominal markings, siphunculi with swollen part dark and the cylindrical basal part paler, and wings with dark triangular spots at the ends of all the veins. In the Mediterranean area, Madeira, Middle East, Central Asia, Pakistan, northern India, and also now in southern Poland (Wieczorek & Chłond 2019), Australia (see Hales et al. 2009), Africa south of the Sahara (Burundi, Kenya, S Africa, Zimbabwe), western USA (California, Oregon) and South America (Argentina, Ortego 1994; Bolivia, Remaudière et al. 1992; Brazil, Peronti & Sousa-Silva 2002). Also now recorded from Chile (Nieto Nafría et al. 2016b), from some surprising host plants; Phrodus microphyllus (Solanaceae) and Pitraea cuneato-ovata (Verbenaceae). Single alatae have also been trapped in the UK. Monoecious holocyclic in Iran (Naumann-Etienne & Remaudière 1995); sexual morphs were found there on Nepeta in November, the males being apterous, and fundatrices were found on the same host in April (G. Remaudière, pers. comm.). Sexual morphs from Iran and Pakistan were described by Wieczorek & Chłond (2019). 2n = 12.

Euceraphis Walker

Calaphidinae: Calaphidini

A genus of 6 or more species living on Betula and Alnus. They are rather large aphids with a relatively short antennal terminal process, knobbed cauda and rounded anal plate. The alder-feeding aphids are somewhat distinct from the birch-feeders. All adults apart from oviparae are winged, active insects which secrete bluish white wax, often in the form of tufts on their legs. For photographs of North American species see aphidtrek.org. The birch-feeding species often have black dorsal abdominal markings, but these vary greatly according to season and are often absent in midsummer. They seem very host-specific, although adults fly a lot and may be found on many plants besides their true hosts (but see Pashchenko 1984b). The oviparae are large, apterous, yellowish to dark brown with dark dorsal abdominal markings, and with the posterior abdomen prolonged into a conical “ovipositor”. Blackman (1988) reported on the unusual karyotypic variation in this genus. Blackman & De Boise (2002) reviewed the genus and keyed the species.

Euceraphis betulae Koch Plate 10a Adult alatae have a pale green to pale yellow abdomen with bluish white wax and a very variable degree of black pigmentation (see influentialpoints.com/Gallery); BL 3.0-4.2 mm. The head and thorax are normally black, both dorsally and ventrally, the legs and antennae variably pigmented, sometimes quite dark, and the dorsal abdomen may be unmarked or have transverse black bands (in spring or autumn), or black patches on abdominal tergites 4 and 5 only (in autumn). Oviparae and alate males occur in September-November (see “A year in the life of a birch aphid”). E. betulae occurs on Betula pendula throughout Europe, in south-west and Central Asia, and on introduced B. pendula in North America, Australia and New Zealand. A morphological comparison with E. punctipennis in Europe is given by Blackman (1977). Hajek & Dahlsten (1988) studied ecology of introduced populations in California, Neuvonen & Lindgren (1987) studied effects of simulated acid rain in Scandinavia and Holopainen et al. (2009) demonstrated its marked preference for yellowing birch leaves in autumn. The name has been applied to populations on native birches in North America and east Asia, now shown by karyotypic and morphometric studies to be separate species (Blackman & De Boise 2002). 2n=9 or 10 (difference is due to number of “B” chromosomes; Blackman 1988).

Euceraphis betulijaponicae (Matsumura) Adult alatae pale green or greenish yellow, covered in bluish wax; BL 3.4-4.2 mm. Described from Japan, and found on various Betula spp. and on introduced Alnus glutinosa in east Siberia (Pashchenko 1988a), but sexual reproduction is apparently limited to Betula platyphylla var. japonica and its closest relatives (B. dahurica, B. kamschatica, B. mandshurica). Oviparae and males collected in October were described by Pashchenko (1984b, 1988b). 2n= 9 (with one small “B” chromosome).

Euceraphis borealis Blackman Colour of adults in life unknown, probably pale green with variably developed dark dorsal abdominal markings, and bluish-white wax; BL 3.1-3.8 mm. On Betula glandulosa and B. nana in northern Canada, Alaska and Finland (Blackman & De Boise 2002), and now also recorded from north-east Siberia (Chukotka; Stekolshchikov & Khruleva 2015). Oviparae and males in late July-August. 2n=8 (differs from other members of E. betulae group by having only one pair of X-chromosomes).

Euceraphis caerulescens Pashchenko Adult alatae pale green, covered in bluish wax; BL 3.5-4.4 mm. Recorded from various Betula spp., in east Siberia, especially B. dahurica and B. ermanii, and also recorded from Alnus hirsuta (Pashchenko 1988a). Also collected in Japan on B. ermanii (Blackman 1986, wrongly identified as E. ontakensis), and recently reported from Kazakhstan (Kadyrbekov 2017a). Sexual morphs have been collected in October from B. dahurica. 2n=22.

Euceraphis gillettei Davidson Adult viviparae are active, pale green alate aphids with pale brown thorax, dorsal abdomen never with dark markings, and only a small amount of wax (see influentialpoints.com/Gallery); BL 2.5-3.5 mm. On Alnus spp. (rhombifolia, rubra, rugosa, tenuifolia) in North America. Oviparae and alate males occur in September-November. The name may be being applied to more than one species, as indicated by the variation in karyotype. 2n=15, 16, 18 or 19 (Blackman 1988).

Euceraphis lineata Baker Adults are pale yellow-green, with distinctive brown and black dorsal markings; BL 3.3-4.0 mm. The head and prothorax have a pale to dark brown longitudinal band, the mesothoracic dorsal lobes are brown, and in the most pigmented specimens the scutellum may be black, but the thorax never has any pigment ventrally. Sexuparae in late summer-autumn usually have a large black patch on abdominal tergite 3, and a smaller one on abdominal tergite 4. Femora are pale except for black “knees”, tibiae have a dark dorsal longitudinal stripe and black apices, and tarsi are black. Immatures have banded antennae and dark tarsi contrasting with their pale tibiae. This species is specific to B. populifola in north-eastern USA and eastern Canada. Sexuales occur in September-October. 2n=16.

Euceraphis mucida (Fitch) Adult viviparae are pale green to yellow-green with an orange-brown thorax and dark tibiae coated with blue-grey wax; BL 3.4-3.8 mm. Dorsal abdomen usually unmarked in spring, but alate sexuparae in late summer-autumn may have transverse bars on abdominal tergites 4 and 5. On Betula lenta in northern USA east of the Rocky Mountains (not in western USA, contrary to Blackman & Eastop 1994). Oviparae and alate males occur in September-October. 2n=20, 21 or 22, the differences are due to variation in the number of accessory (“B”) chromosomes (Blackman 1988).

Euceraphis ontakensis Sorin Adult alate viviparae are pale green with dark legs, often with black bars on abdominal tergites 4 and 5, and secreting bluish white wax; BL 3.0-4.0 mm. On Betula ermanii in Japan; also recorded from Alnus fruticosa in Japan (Sorin 1970a), and from several other Betula and Alnus spp. in east Siberia. The occurrences on Alnus, and possibly those on Betula spp. other than ermanii, may only be casual. Sexuales are unknown.

Euceraphis papyrifericola Blackman Adult alatae are pale green with mid- to dark brown thorax, and (in autumn) variably developed dorsal abdominal cross-bands, often with bluish-white wax (see influentialpoints.com/Gallery); BL 3.4-4.1 mm. On Betula papyrifera, and occasionally on B. populifola and B. cordata, in north-east USA and across Canada (Blackman & De Boise 2002). Oviparae and males occur in late August to October. 2n=9 (including one small “B” chromosome).

Euceraphis punctipennis (Zetterstedt) Adult viviparae are pale green with a dark brown head and thorax (see influentialpoints.com/Gallery), often somewhat larger than E. betulae (BL 3.0-4.8 mm), and with dorsal black patches when present often confined to abdominal tergites 4 and 5, even in spring generations (Blackman 1977, and see “A year in the life of a birch aphid”). On B. pubescens throughout Europe (including Iceland and the Faroes), and also reported from Kazakhstan (Kadyrbekov 2017a). Records from east Siberia (Pashchenko 1984b) and China (Qiao et al. 2005b) may apply to another member of the E. betulae group. Oviparae and alate males occur in October-November. 2n=7 or 8, according to whether there are one or two (“B”) chromosomes (Blackman 1976).

Euceraphis quednaui Blackman Adult alatae with pale green to yellow head, pale yellow-brown thorax encrusted laterally with bluish-white wax, and abdomen pale green with dorsal cross-bands of bluish-white wax (there can also be copious wax on appendages giving them a feathery appearance); BL 3.0-4.0mm. On Betula occidentalis in western USA (Idaho, Utah). An ovipara was collected in late October, but the male is undescribed (Blackman & De Boise 2002). 2n=11 (including 3 “B” chromosomes).

Eulachnus Del Guercio

Lachninae: Eulachnini

About 15 species of long-bodied aphids living on needles of Pinus, the Old World equivalent of the nearctic genus Essigella. They are cryptic when feeding, but very active when disturbed. The best-known species show preferences for certain Pinus spp., but none are strictly monophagous. Accounts are available for Italy (Binazzi 1983, 1984, 1989), the Iberian peninsula (Nieto Nafría et al. 2002a), Britain (Carter & Maslen 1982), Fennoscandia and Denmark (Heie 1995), central Europe (Pintera 1968, Szelegiewicz 1978), former Czechoslovakia (Pasek 1954), Hungary (Szelegiewicz 1978), India (A.K. Ghosh 1982; Chakrabarti et al. 2020), east Siberia (Pashchenko 1988a), Korea (Lee et al. 1994) and Japan (Inouye 1970). For illustrations of and more detailed biological information on some European species see Binazzi & Scheurer (2009). Kanturski et al. (2015) provided stereoscan images of European species, in particular comparing chaetotaxy and male genitalia, and Kanturski et al. (2017c) reviewed and keyed the European species, with detailed photographic images of all available morphs. The genus needs further revision, including biosystematic and molecular studies, as many of the characters that have been used in species discrimination are undoubtedly subject to environmental influences. Lampel & Burgener (1987) studied phylogenetic relationships with other Cinarini based on enzyme electrophoretic data. Diaeretus leucopterus and Praon bicolor are specialised parasitoids.

Eulachnus agilis (Kaltenbach) Apterae are spindle-shaped, bright green with numerous reddish-brown spots, without any significant wax (see influentialpoints.com/Gallery); BL 1.6-2.5 mm. The hind legs often have mottled pigmentation. On many Pinus spp., but particularly common on P. sylvestris, usually feeding on old needles. It occurs throughout Europe, in Siberia and south-west and Central Asia (Kadyrbekov 2017a), and is also recorded from east Asia (China, Korea). It occurs on planted P.sylvestris in North America, where feeding damage was studied by Bliss et al. (1973); and is more recently report as a pest in Brazil (Iede et al. 2007). Monoecious holocyclic; oviparae and alate males are produced in October-November, and the oviparae lay eggs singly, mainly on leaf scars on the branches (Bliss & Kearby 1971). The male genitalia were described and illustrated by Wieczorek et al. (2012). Kearby & Bliss (1969) studied population trends and control, Kidd et al. (1985) studied its association with Schizolachnus pineti.

E. agilis is a member of a closely related group of species discussed and keyed by Binazzi (1983, 1989). E. alticola Börner, described from P. mugo and occurring at high altitudes in central and south-eastern Europe (Heinze 1962, Pintera 1968, Kanturski et al. 2017c), will key to E. agilis in our key, and may or may not be specifically distinct; the minor differences from agilis, involving hair lengths and pigmentation, could easily be due to geographical variation, or environmentally induced. Gabrid (1989) recorded aphids identified as E. alticola from numerous native and introduced Pinus spp. in Kirghizia, and G. Zhang et al. (1992a) recorded it from China. Kanturski et al. (2017c) described oviparae and males of E. alticola and designated a lectotype. E. cretaceus Mamontova, from P. (sylvestris var.) cretacea in Ukraine (Mamontova 1968), is also very close to agilis, and was synonymised with it by Kanturski et al. (2017c). 2n=8.

Eulachnus brevipilosus Börner Apterae are spindle-shaped, pale to bright green with or without faint brown dorsal spots, and without wax (see influentialpoints.com/Gallery); BL 1.4-2.2 mm. An orange form has been found in Poland and Czech Republic (Kanturski et al. 2017c).They feed on the needles of Pinus spp. (mostly on mugo, nigra, sylvestris) throughout Europe, especially in montane areas, and have been introduced to New Zealand (Zondag 1983) and north-western USA (Washington; Hottes & Essig 1955) and British Columbia, Canada (BMNH collection , leg. C.K. Chan). Monoecious holocyclic; fundatrices, oviparae and alate males have been described and illustrated by Kanturski et al. (2017c). However, viviparae have been collected in winter months in England, indicating that anholocycly can occur. Ponsen (1981) studied the digestive system. 2n=30.

Eulachnus cembrae Börner Apterae are spindle-shaped, bright green to yellowish green, dusted with wax powder (Pasek 1954); BL 2.3-3.0 mm. On needles of Pinus cembra in montane Europe (Austria, Italy, Switzerland, Poland, Slovakia). Possibly also in Turkey (Görür et al. 2017). There are also records from P. strobus and P. peuce. Oviparae and alate males occur in Late August to mid-October and were fully described by Kanturski & Wieczorek (2014) and Kanturski et al. (2017c). This species was treated as a synonym of the east Asian species E. pumilae by authors including Blackman & Eastop (1994), but Remaudière & Remaudière (1997, p.302) listed several differences.

Eulachnus intermedius Binazzi Apterae are pale brown, covered with a grey waxy secretion: BL 1.8-2.5 mm. Described from P. mugo in a montane locality in central Italy. Kamturski et al. (2017c) provided a full redescription, including oviparae and alate males collected in early September. As the name suggests, this species has many characters intermediate between the much more common and widely distributed E. agilis and E. rileyi (q.v.). Apterae are most likely to run to agilis in our key to pine aphids.

Eulachnus isensis Sorin Apterae are reddish brown, faintly dusted with white powder, with a pattern of dark brown dorsal spots similar to that of Eu. thunbergii, and dusky to dark legs; BL c. 1.9 mm. On Pinus densiflora in Japan (Mie Prefecture, Honshû). Oviparae and small apterous reddish-brown males were collected in October-November (Sorin 2012b).

Eulachnus mediterraneus Binazzi Apterae are pale to mid green, without wax; BL 1.8-2.7 mm. On needles of Pinus pinaster, and occasionally on the P. nigra group, in the western Mediterranean region (Italy, Corsica, Madeira and the Iberian peninsula). Apparently entirely anholocyclic, with apterae collected from February to October. Large populations can occur causing stunting of growth of young plants, and premature leaf-fall of older ones (Nieto Nafría et al. 2002a). Binazzi & Mier Durante (1997) described another anholocyclic form with darker hind tibiae on P. pinaster and P. sylvestris and apparently native to the Iberian peninsula as a subspecies, E. mediterraneus ssp. ibericus, which has now been given full species status by Kanturski et al. (2017c).

Eulachnus nigricola (Pasek) Apterae are spindle-shaped, pale green, with brownish legs and antennae and slight wax dusting on head, sides of thorax and dorsal abdomen; BL 1.6-2.3 mm. On needles of Pinus nigra and other pines of subsect. Sylvestres, mainly in montane or coastal areas. Widely distributed in Europe (see Danilov 2019b), east to Turkey (Canakçioğlu 1975 and BMNH collection) and Iran (Rezwani et al. 1994). A record from Tunisia requires further confirmation. Holocyclic, with oviparae and very small alate males in October (Nieto Nafría et al. 2002a).

Eulachnus piniarmandifoliae Zhang Apterae are green, with black antennae and legs, without wax; BL c. 2.0 mm. Other morphs are undescribed. On Pinus armandii in China (Hunan and Yunnan; G. Zhang et al. 1992a).

Eulachnus pumilae Inouye Apterae are spindle-shaped, bright green to yellowish green; BL c. 2.4-2.5 mm. On needles of Pinus spp. of subsection Cembrae (parviflora, pumila, koraiensis), in northern India, east Siberia, Korea and Japan. An ovipara and alate male collected in South Korea in early November were described and compared with those of P. cembrae by Kanturski &Wieczorek (2014).

Eulachnus rileyi (Williams) Plate 16d Apterae are spindle-shaped, varying in colour from dark olive green to orange-brown or grey, with a dusting of bluish-grey wax (see influentialpoints.com/Gallery); BL 1.8-3.0 mm. At least the hind pair of legs are usually dark. On needles of a taxonomically wide range of Pinus, including many North American species, but on Old World pines of subsection Sylvestres it occurs much more commonly on P. nigra and P. mugo than on P. sylvestris. Heavily attacked needles turn yellow and are lost prematurely (Felt & Bromley 1936). In Europe, Mediterranean area, south-west and Central Asia, and introduced into Africa south of the equator (Mills 1990) and North, South and Central America. Monoecious holocyclic in Europe and North America, with oviparae and alate males appearing in September-November, but apparently anholocyclic in warmer regions (Marchant 1981). Katerere (1983, 1984) studied its population ecology in Zimbabwe, and reported on the incidence of a fungal infection. Tremblay & Micieli de Biase (1970) recorded parasitism in southern Italy. 2n=8.

Although most populations of E. rileyi are readily distinguished from the related species E. agilis, forms with apparently intermediate or combined characters of the two species occur in south and central Europe, and apparently also in China. E. intermedius Binazzi (q.v.) has the appearance in life of E. rileyi but hairs more like those of agilis. Two species described from China, E. pinitabulaeformis Zhang from P. tabuliformis and P. massoniana (G. Zhang et al. 1992a), and E. pinisuctus Zhang, Chen, Zhong & Li from P. armandii (in G. Zhang 1999) also have features of both E. agilis and E. rileyi. E. tauricus Bozhko, described from Pinus nigra var. pallasiana in Crimea (Bozhko 1957, 1961) also differs from typical E. rileyi only in the characters of the hairs on the antennae and tibiae; it was given subspecies status by Hille Ris Lambers (1966), and populations on P. halepensis in Italy (Binazzi 1983) and on P. pinea in Israel (Halperin 1986) have subsequently been identified as E. rileyi tauricus. In more recent publications tauricus was again given separate species status, and Nieto Nafría et al. (2002) provide several key characters separating apterae and alatae of the two taxa. However, it has now been synonymised with E. rileyi by Kanturski et al. (2017c). Without evidence of specific host associations, and until the molecular systematics of the group has been fully investigated, it might be wise to regard all these forms as variant anholocyclic populations of E. rileyi, or possibly permanently parthenogenetic lineages resulting from natural hybridisation between agilis and rileyi, reflecting the patterns of variation that one might expect to find in populations that have ceased to undergo sexual reproduction.

Eulachnus similialticola Zhang Appearance in life is unrecorded; BL of aptera c.1.9 mm. Alatae are undescribed. On Pinus massoniana in Guangxi, China (Qiao et al. 2002).

Eulachnus stekolshchikovi Kanturski Appearance in life is unknown; BL of aptera 1.9-2.0 mm. Described from specimens collected on Pinus sylvestris and P. nigra in Scotland (including one specimen collected in June and identified as a fundatrix), and from P. sylvestris in Russia (Murmansk). Very similar to E. mediterraneus, but in our key to pine aphids it is most likely to run to agilis.

Eulachnus tauricus Bozhko See under E rileyi above.

Eulachnus thunbergii (Wilson) Apterae are spindle-shaped, dirty bluish green to yellowish green or yellowish brown, dusted with bluish white wax (A.K. Ghosh 1982, Moritsu 1983); BL 2.3-3.2 mm. On needles of various native Pinus spp. in east and south-east Asia (India, Vietnam, China, Japan, Korea, Siberia, Taiwan, Java, Philippines), and also in Australia (Carver & Kent 2000). A record from Turkey (Şenol et al. 2014b) requires further confirmation. Oviparae and alate males occur in Japan in October-November (Inouye 1970), and oviparae are also recorded from India (Raychaudhuri et al. 1983). Mao (1986) studied population trends in Japan, and A.K. Ghosh (1982) noted predators and parasitoids in India. E. drakontos Zhang & Qiao (in Qiao & Zhang 1999b) is very similar except for having shorter hairs, and seems likely to be a synonym. 2n=8 (Khuda-Bukhsh & Kar 1990).

Eulachnus tuberculostemmatus (Theobald) Apterae are narrowly spindle-shaped, pale green to greenish yellow, with small brown dorsal spots; BL 1.4-2.7 mm. On needles of Pinus spp., especially halepensis; also brutia, canariensis, eldarica, pinaster, pinea and nigra (incl. vars laricio, salzmannii). In southern Europe, Mediterranean area including North Africa, and Asia eastward to Pakistan and China. Its population ecology in Iran was studied by Hosseini & Poorjavad (2020). No sexual morphs are recorded. 2n=8.

[E. tuberculostemmatus ssp. garganicus Binazzi (1983b) was described from one sample from P. halepensis in southern Italy, but has now been given full species status by Kanturski et al. (2017c).]

Eumyzus Shinji

Aphidinae: Macrosiphini

Eleven east Asian species with various host associations, and in none is the life cycle completely known. Three species on woody Rosaceae in spring in India migrate to unknown secondary hosts. Three species on Impatiens and two on Hydrangea induce galling responses in their hosts. The host of one species is unrecorded. Eumyzus have a spinulose head like Myzus but the antennal tubercles are divergent and the dorsal hairs are rather long and stiff, usually with tuberculate bases. Chakrabarti & Bhattacharya (1985) reviewed the genus. The biological diversity within the genus may indicate that some species are misplaced and that the concept of the genus needs to be revised.

Eumyzus clinopodii Takahashi Apterae are oval, shiny dark reddish black with dark siphunculi and cauda and paler brownish antennae and legs; BL 1.1-1.5 mm. It was described from Clinopodium gracile in Japan, and aphids producing reddish bag- or boat-shaped galls on Isodon spp. have also been identified as this species (M. Miyazaki, pers. comm.). Probably the same species has also been recorded forming upwardly-rolled leaf margin pocket galls on Isodon rugosus in Pakistan (Naumann-Etienne & Remaudière 1995, as Eu. plectranthi). Its generic position is uncertain. [The earlier name Eumyzus plectranthi (Shinji) is applied to this aphid by some authors, including Moritsu (1983), but others regard this as a nomen dubium.]

Eumyzus darjeelingensis R.C. Basu & Raychaudhuri Appearance in life is unknown; BL of apterae 1.3-1.9 mm. On Hydrangea paniculata in West Bengal, India, presumably in leaf-roll galls. Alatae have 4-9 secondary rhinaria on ANT III.

Eumyzus eastopi Maity & Chakrabarti Colour in life is unrecorded, probably dark; BL of aptera 1.7-1.9 mm. On Pyrus vestita (= Sorbus cuspidata) in June in Uttar Pradesh, India, rolling the edges of the leaves (Maity et al. 1982, Chakrabarti & Bhattacharya 1985). Other morphs and life cycle are unknown. 2n=10 (Khuda-Bukhsh & Pal 1986b).

Eumyzus gallicola Takahashi Colour in life is unknown; BL of aptera c. 1.9 mm. In bag-shaped pseudogalls on leaves of Impatiens noli-tangere in Japan (Miyazaki 1971). 2n=12.

Eumyzus hydrangi Chakrabarti & Bhattacharya Colour in life is unknown; BL of aptera 1.2-1.4 mm. In closed marginal leaf-roll galls on Hydrangea scandens in Uttar Pradesh, India. Medda & Chakrabarti (1986a) described the alatae, which have secondary rhinaria distributed III 8-13, IV 0-2.

Eumyzus impatiensae (Shinji) Plate 17e (Fig.33f) Apterae are pink, red or reddish brown, with siphunculi and cauda, and apices of antennae and legs, black; BL 1.8-2.0 mm. Forming bag-shaped pseudogalls on veins of leaves of Impatiens spp. in Japan, Korea, east Siberia and India (Uttar Pradesh). Re-described by Miyazaki (1971). Monoecious holocyclic (Gredina 1996). 2n=12 (Blackman 1986) or 10 (Pal & Khuda-Bukhsh 1980).

Eumyzus indicus Medda & Chakrabarti Apterae are dark brownish; BL 1.5-2.1 mm. In bag-galls protruding from undersides of leaves of Impatiens spp. in Uttar Pradesh, India.

Eumyzus nokuli Raychaudhuri, Singh & Raychaudhuri Colour in life is unknown; BL of aptera c.1.5 mm. On Senecio sp. in Manipur, India. No effect on the host plant is reported, and the generic position of this aphid is uncertain.

Eumyzus pruni Chakrabarti & Bhattacharya Colour in life is unrecorded, probably dark, with dark siphunculi; BL of aptera 1.7-1.9 mm. On Prunus cornuta, living in tight reddish marginal leaf-rolls in spring (Medda et al. 1986). Heteroecious holocyclic, migrating to an unknown secondary host; oviparae and alate males are described by Medda & Chakrabarti (1989), collected on P. cornuta in November. In Uttar Pradesh, India and in the Murree Hills, Pakistan (Naumann-Etienne & Remaudière 1995).

Eumyzus prunicola Medda & Chakrabarti Colour in life is unrecorded, probably pale, with pale siphunculi; BL of aptera 1.8-2.1 mm. On Prunus padus in Uttar Pradesh, India, rolling the leaves in spring. According to Medda & Chakrabarti (1986b), the fundatrices induce two types of gall; either the edge of the leaf is curled ventrally and becomes somewhat shrunken, shortened and rugose, or a caterpillar-like pouch gall is produced on the upper side of the leaf lamina, with a slit-shaped aperture to the underside. (This gall difference suggests that two species are involved and needs further investigation). Alatae are produced in June and presumably migrate to an unknown secondary host. Sexual morphs are unknown.

Euthoracaphis Takahashi

Hormaphidinae: Nipponaphidini

Small, heavily sclerotised aphids related to Nipponaphis, with rather numerous fine hairs on dorsal and marginal areas of prosoma, and with sutures dividing the dorsum of the prosoma into a large quadrate central area, plus an anterior and two dorsolateral areas. Four species are known, three on Lauraceae, and one reportedly on Senecio which is unlikely to be the true host (A.K. Ghosh & Raychaudhuri 1973a, Noordam 1991). J. Chen et al. (2009) reviewed the genus with photographs and a key to apterae, and Qiao et al. (2018) reviewed the Chinese species..

Euthoracaphis heterotricha Ghosh & Raychaudhuri Apterae are aleyrodiform, dark brown to bluish black with dorsum covered by a thin floury secretion; BL 1.0-1.8 mm (Noordam 1991). On lower sides of leaves of Cinnamomum spp. (zeylandicum, iners) in Java, and on C. burmanni in China (Yunnan: J. Chen et al. 2009). Originally described from an unknown host in West Bengal (A.K. Ghosh & Raychaudhuri 1973a). Alatae were found in Java in May, and in China in January. Sexual morphs and life cycle are unknown.

Euthoracaphis longisetosa A.K. Ghosh & Raychaudhuri Apterae are presumably black in life; BL not recorded. Described from 4 apterae found on undersides of leaves of a Senecio sp. in Arunachal Pradesh, India. This is probably not the true host.

Euthoracaphis oligostricha Chen, Fang & Qiao Apterae are aleyrodiform, dark brown with pale antennae, legs, anal plate and cauda, the body covered with white wax; BL 1.1-1.4 mm. Alatae are undescribed. On undersides of leaves and shoot tips of Persea (= Machilus) yunnanensis in China (Yunnan: J. Chen et al. 2009).

Euthoracaphis umbellulariae (Essig) Apterae are aleyrodiform, sedentary, almost circular, pill-box-shaped, black, covered with white filamentous wax (Essig 1932); BL c. 0.8 mm. Alatae are jet black. On undersides of leaves of Cinnamomum spp. (japonicum, camphoratum) in Japan (Takahashi 1959b), and presumably introduced from east Asia into California USA, where it is very common on Umbellularia californica, and also recorded from Sassafras variifolium (= albidum) and Cinnamomum spp. (Essig 1932, Hille Ris Lambers 1966a). Sexuales and life cycle are unknown; presumably any holocycle would involve Distylium as primary host. Populations in California are anholocyclic. Hille Ris Lambers (1966a) queried Takahashi’s synonymy of cinnamoniae and umbellulariae, but specimens in the BMNH collection do not show any consistent differences between Japanese and Californian populations.

Eutrichosiphum Essi & Kuwana

Greenideinae: Greenideini

A genus of about 50 species, mostly on Fagaceae; the exceptions are described from Betulaceae, Elaeocarpaceae, Lauraceae, Apocynaceae, Sonneratiaceae, Euphorbiaceae, Sterculiaceae and unidentified plants. Eutrichosiphum have an elongate oval or pear-shaped body, a long, tapering and distinctly subdivided last rostral segment, hind tibiae without stridulatory ridges, siphunculi without any reticulation in apterae, and a cauda without a median process. Separate genera or subgenera have been erected for such aphids, grouping species according to the number of antennal segments, extent of dorsal spinulosity, and the nature of the siphuncular or dorsal body hairs. None of these characters are entirely satisfactory for subgeneric classification (A.K. Ghosh 1987), so here we adopt a conservative approach, regarding Brevitrichosiphon, Holotrichosiphon, Paratrichosiphum, Neoparatrichosiphum and Neotrichosiphum as all part of Eutrichosiphum and not recognizing any subgenera. Raychaudhuri (1956) revised the species then known, Takahashi (1962b) reviewed the Japanese species, Noordam (1994) reviewed the species in Java, and A.K. Ghosh & Agarwala (1993) provided an account of the Indian species. The genus is still in need of further revision, particularly in the case of the complex of species feeding on oaks in the eastern Himalayan region. Sugimoto (2001) discussed seasonal and geographical variation in length and form of hairs on body and siphunculi of E. heterotrichum in Japan – information which may be applicable to other Eutrichosiphum and one of the reasons for the many taxonomic problems in this genus.

Eutrichosiphum alnicola (Basu) Apterae are brown (Das & Raychaudhuri 1983); BL 1.8-2.7 mm, on tender shoots of Alnus nepalensis in north-west India (A.N. Basu 1968) and Nepal. Saha & Chakrabarti (1988a) described the alate male, collected in October in western Himalaya. The alate oviparae collected in association with greenish apterae on Quercus and ascribed to alnicola by Raychaudhuri et al. (1980a) are unlikely to be this species. It is not clear how alnicola differs from alnifoliae Tao (see below).

Eutrichosiphum alnifoliae (Tao) Apterae are mainly blackish brown; BL c.1.7-1.8 mm. Alatae are light yellowish brown. On tender shoots of Alnus cremastogyne in Szechuan, China (Tao 1958). Sexuales and life cycle unknown. E. alnicola (Basu) could be a synonym.

Eutrichosiphum alnisuctum Zhang Apterae are brown; BL c. 2.5 mm. On an unidentified Alnus sp. in Tibet (Xizang; G. Zhang & Zhong 1988). This species is probably misplaced in Eutrichosiphum and may be a synonym of Mollitrichosiphum nandii.

Eutrichosiphum apicifuscum Sugimoto Apterae are yellow to yellowish brown, with pale brown legs and siphunculi darker distally than at their bases; BL 1.6-2.0 mm. On undersides of young leaves of Lithocarpus edulis in spring, with an abbreviated life cycle, alate oviparae and males appearing in May-June (Sugimoto 2011b). Only known from central Japan.

Eutrichosiphum arunachali Basu, Ghosh & Raychaudhuri Apterae are dark brown; BL 1.3-1.6 mm according to original description, but 2.8 mm for one specimen from the type series measured by Raychaudhuri & Chatterjee (1980). Described from an unidentified Quercus sp. in Arunachal Pradesh, India (R.C. Basu et al. 1972), and also recorded from Sikkim (Mondal et al. 1979), and from Castanopsis sclerophylla in Fujian province, China (Zhang et al. 1999d; a sample of two apterae is described as a subspecies, Eu. arunachali jianglense). The alata is described by Chatterjee et al. (1981); it is longer-haired than that of the otherwise similar E. pasaniae.

Eutrichosiphum assamense Ghosh, Basu & Raychaudhuri Appearance in life is unrecorded, probably brown with pale antennae, legs and siphunculi; BL of aptera 1.7-2.3 mm (A.K. Ghosh et al. 1969a). On Quercus fenestrata (= Lithocarpus fenestratus) in Meghalaya, India, and on Quercus spp., Nepal (BMNH collection, leg. K.C. Sharma) and Tibet (Q. Liu et al. 2013). It is also tentatively recorded from Q. incana in Pakistan (Naumann-Etienne & Remaudière 1995). A.K. Ghosh & Agarwala (1993) redescribed apterae from unidentified Quercus sp(p.) in Meghalaya, Himachal Pradesh and Uttar Pradesh, and also described an alate vivipara, an alate ovipara and an alate male under this name.

Two species treated separately in Blackman & Eastop (1994), E. garhwalense Maity & Chakrabarti and E. taoi A.K. Ghosh, Basu & Raychaudhuri, were synonymised with E. assamense by A.K. Ghosh & Agarwala (1993). E. garhwalense was described from Q. incana in Uttar Pradesh (Maity & Chakrabarti 1980), and its sexual morphs, collected in June-July, were described by Saha & Chakrabarti (1988a). Aphids identified as E. taoi have been collected in Uttar Pradesh on Quercus sp. (Chakrabarti et al. 1972b), and in Manipur on Q. serrata (Agarwala et al. 1980). E. simlaense L.K. Ghosh, described from an unidentified Quercus sp. in Himachal Pradesh (L.K. Ghosh 1986), is another possible synonym.

There are problems with the application of the name assamense, as it is a secondary homonym of E. tattakanum ssp. assamense A. K. Ghosh & Raychaudhuri 1962, which was synonymised with E. khasyanum (Raychaudhuri & Chatterjee 1980; but see also under E . blackmani, below), and is therefore invalid. The two are confused in the check-list of Agarwala & A.K. Ghosh (1984), and most of the references they give refer to the latter species. We retain the name here in anticipation that future work may reveal further synonymies.

Eutrichosiphum betulae Mondal, Chatterjee & Raychaudhuri Appearance in life is not recorded, probably pale; BL of aptera 2.9-3.3 mm. On on unidentified Betula sp. in Sikkim, India (Mondal et al. 1979). Other morphs and biology unknown. To judge from the descriptions, E. nepalensis could be a synonym.

Eutrichosiphum blackmani A.K. Ghosh & Agarwala Appearance in life is not recorded. On Quercus sp. in Meghalaya, India (A.K. Ghosh & Agarwala 1993), and there are also records from Castanopsis and Lithocarpus (see Singh & Singh 2017c). This is part of the material described as E. tattakanum ssp. assamense A. K. Ghosh & Raychaudhuri 1962, the rest being synonymised with E. khasyanum (see under assamense above). It is apparently very similar to E tattakanum, but the antennal terminal process is longer (PT/BASE VI ratio 1.54-1.93) than that of other Quercus-feeding Eutrichosiphum with a long R IV+V and non-spinulose dorsal abdomen.

Eutrichosiphum davidi Raychaudhuri Apterae are probably dark brown; BL 1.7 (holotype)-3.1 mm. On Quercus serrata in southern India (Raychaudhuri 1956); also recorded from Quercus sp. Sikkim (Mondal et al. 1979), and alate viviparae have been trapped in Nepal (BMNH collection, leg. K.C. Sharma). Sexuales and life cycle are unknown.

Eutrichosiphum dubium van der Goot Apterae are pear-shaped, dark brown; BL 1.9-2.7 mm. On young growth of Quercus spp. (dealbata, glauca), Castanopsis sp. and Lithocarpus spp. (glaber, konishii); also collected on Litsea sebifera (= glutinosa) (Raychaudhuri et al. 1977). In India (West Bengal, Manipur), China (Sichuan) and Taiwan. Specimens from Hong Kong and Japan in the BMNH collection listed under this name in Blackman & Eastop (1994) were identified as E. heterotrichum by Sugimoto (2001). G. Zhang & Zhong (1985) distinguished a population from Yunnan, China (which included apterae with 5-segmented antennae) as spp. yulongshanense. Devi & Singh (1987) studied ant attendance.

Eutrichosiphum flavum Takahashi Apterae are orange-yellow, with rather short, pale siphunculi, pale legs, and ANT PT/BASE about 1; BL 1.6-1.9 mm. Some of the paratypes in the BMNH collection have 5-segmented antennae. On young leaves of ?Lithocarpus sp. in Thailand (Takahashi 1941); reported to occur in India (Manipur, Meghalaya) on Litsea khasyana, as well as on Castanopsis sp. and Quercus sp. (A.K. Ghosh & Agarwala 1993). Singh et al. (1980) described an alate vivipara purported to belong to this species. Sexuales and life cycle are unknown.

Eutrichosiphum glabrum Noordam Apterae are pale brownish yellow with a pair of orange spots on the pronotum and at siphuncular bases, orange cross-bands on other segments, legs almost colourless and siphunculi with distal part black; BL 1.4-2.1 mm. On Quercus spp. (glauca, gemelliflora) in Java, living mostly on undersides of leaves and young shoots (Noordam, 1994). Other morphs and life cycle are unknown.

Eutrichosiphum heterotrichum (Raychaudhuri) Apterae are pear-shaped, yellowish brown to almost black with orange-brown head, black siphunculi and legs mainly pale brown or yellow brown; BL 2.1-3.3 mm. Described from Quercus sp. in Java (Raychaudhuri 1956), and also collected on Lithocarpus spp. and Quercus spp. in China (Hong Kong, Fujian, Hainan, Guandong; Q. Liu et al. 2013) and Japan (Sugimoto 2001). The aphids live on young green stems and between scales of young shoots as well as on 2-year-old branches (Noordam 1994). Detailed descriptions were provided of the aptera and alata from Java by Noordam (1994) and of all morphs in Japan by Sugimoto (2001). Holocyclic in Japan with alate sexual morphs appearing from October onward, but also with part of the population overwintering parthenogenetically (Sugimoto 2001). 2n=20 (this chromosome number was erroneously applied to E dubium in Blackman & Eastop 1994).

Eutrichosiphum jugeshwari Singh Apterae are elongate pear-shaped, appearance in life not recorded but probably rather pale, with black-tipped siphunculi; BL 2.3-2.6 mm. On Quercus sp. in Manipur, India (T.K. Singh et al. 1979). Other morphs are unknown,

Eutrichosiphum khasyanum (Ghosh & Raychaudhuri) Apterae are pear-shaped, dorsal abdomen yellowish brown to dark brown, with paler head and thorax and pale antennae, legs and siphunculi (according to original description – but Raychaudhuri et al. (1980a) reported cream-coloured apterae); BL 1.8-2.1 mm. Originally collected from undersides of leaves of an unidentified Quercus sp. in Meghalaya, India; subsequent records are from Q. dealbata and Q. griffithii in northern India and from Q. semicarpifolia in Nepal (Miyazaki 1977). Raychaudhuri & Chatterjee (1980) provide measurements of an alate vivipara identified as this species. Life cycle and sexuales are unknown.

Eutrichosiphum kumaoni Chakrabarti & Debnath Apterae are yellowish with pale head and siphunculi; BL c.2.2-2.3 mm. Alatae are blackish and immatures green. On undersides of young leaves of a Castanopsis sp. in the Himalayan region (Uttarakhand) of northern India (Chakrabarti & Debnath 2009). The life cycle is unknown.

Eutrichosiphum litseae Raychaudhuri, Raha & Raychaudhuri Apterae are pear-shaped, colour in life unrecorded, probably mainly pale; BL 1.5-1.8 mm. Described from Litsea sebifera (= glutinosa) in Manipur, India. Other morphs and biology are unknown. In every respect except host plant this species seems to resemble E. flavum Takahashi, which has also been recorded from Litsea in India..

Eutrichosiphum makii Raychaudhuri & Chatterjee Appearance in life is not recorded, probably dark brown; BL 1.5-1.7 mm. Described from Glochidion sp. in Meghalaya, India (Raychaudhuri & Chatterjee 1974), and subsequently found on Litsea sp. A.K. Ghosh (1976) described the alate vivipara. Sexual morphs and biology are unknown.

Eutrichosiphum manaliense A. K. Ghosh & Agarwala Appearance in life is unrecorded, apterae on slides are brown to dark brown; BL c. 1.8-1.9 mm. On Betula utilis in Himachal Pradesh, India (A.K. Ghosh & Agarwala 1993).

Eutrichosiphum manipurense Singh, Raychaudhuri & Raychaudhuri Appearance in life is not recorded, probably rather pale except for a brownish central abdominal patch and dark-tipped siphunculi (T.K. Singh et al. 1979); BL 1.7-2.1 mm. On an unidentified Quercus sp. in Manipur, India. Other morphs and biology are unknown.

Eutrichosiphum mukerjii (Raychaudhuri, M.R. Ghosh, Banerjee & A.K. Ghosh Apterae are pear-shaped, dark brown with paler head, antennae and legs and stout blackish siphunculi; BL c.1.3 mm. Described from an unidentified host in West Bengal (Raychudhuri et al. 1973) and on Quercus sp. in Manipur, India (Singh et al. 1979, as Brevitrichosiphon nungsireiae). Other morphs and biology are unknown.

Eutrichosiphum nasensis Lee & Lee Apterae pear-shaped, dorsal abdomen with central part pale greenish yellow, encircled by a broad, contrastingly brown-black margin, and with shiny black siphunculi; BL 1.6-2.0 mm. On undersides of young leaves of a Quercus sp. in Laos (Y. Lee et al. 2016). Alatae and life cycle are unknown.

Eutrichosiphum nepalensis A.K. Ghosh & Agarwala Apterae are greenish; BL 3.2-3.5 mm. On Alnus nepalensis in Nepal (Kirtipur), feeding on undersides of leaves and young shoots, in a heavy infestation that appeared to cause slight curling of the leaves. The original description (Das & Raychaudhuri 1983, under the name E. alnifoliae) agrees remarkably well with that of E. betulae from Betula sp. in Sikkim, India.

Eutrichosiphum nigrisiphon Sugimoto Apterae are yellow to yellowish brown, with pale brown legs and black siphunculi; BL 1.4-1.8 mm. On undersides of young leaves of Lithocarpus edulis in spring, with an abbreviated life cycle, alate oviparae and males appearing in May-June (Sugimoto 2011b). Only known from central Japan.

Eutrichosiphum nigrum Noordam Apterae are pear-shaped, shiny black, sometimes “somewhat spotted”; BL 1.3-1.7 mm. On Lithocarpus bennettii and Castanopsis javanica in Java, feeding on upper and undersides of very young leaves along the main veins, and on young shoots (Noordam, 1994). Other morphs and life cycle are unknown.

Eutrichosiphum pallidum Noordam Apterae are elongate oval, yellowish white or sometimes pale brown, with pale brown siphunculi; BL 1.7-2.4 mm. Alatae have a yellow abdomen with greyish brown spinal spots. On undersides of leaves and young shoots of Castanea crenata in Java (Noordam 1994).

Eutrichosiphum parvulum Eastop & Hille Ris Lambers Plate 5i Apterae are shining black, with head yellowish brown; BL 1.2-1.6 mm. On stems and undersides of leaves of Trachelospermum spp. in China and Taiwan (Takahashi 1923, as E. minutum). 2n=26.

Eutrichosiphum pasaniae (Okajima) Plate 11c, d Apterae are variably pigmented, pale brown to dark greenish or blackish brown with black siphunculi; BL 1.4-1.8 mm. Alatae have an extensive black dorsal abdominal patch. On young shoots and undersides of leaves of Lithocarpus spp., Castanopsis spp., Castanea crenata and Elaeocarpus japonicus; it is likely that records from Quercus are erroneous. In east and south-east Asia. Alate males were recorded in Taiwan in October (Takahashi 1923). Szelegiewicz (1968) distinguished populations on Castanopsis sp. in Vietnam, with a pale body and longer R IV+V than pasaniae from Japan and Taiwan, as E. pseudopasaniae Szelegiewicz. However, there is little or no correlation between pigmentation and length of last rostral segment in populations in north-east India, where both light and dark-bodied apterae may occur on the same plant (Raychaudhuri & Chatterjee 1974). E. menglunense Zhang 1980, described from China, also seems to be intermediate between pasaniae and pseudopasaniae. E. izas Zhang, only known from alate viviparae in China, is also in this group. Comparison of the descriptions of pasaniae from Java by Noordam (1994) and pseudopasaniae from India by Ghosh & Agarwala (1993) reinforce the view that further work is needed to establish whether or not a single, variable species is involved.

Eutrichosiphum pullum Noordam Apterae are shiny black with brown or pale brown head and black siphunculi; BL 1.2-1.7 mm. On young leaves and shoots of Castanopsis spp. in Java (Noordam 1994). Other morphs and life cycle are unknown.

Eutrichosiphum pyri Chakrabarti, Ghosh & Raychaudhuri Apterae are pear-shaped, colour in life not recorded; BL 1.7-2.1 mm. Described from Pyrus communis (Chakrabarti et al. 1972b), but presumably the true host is Quercus; there are subsequent records from Q. semicarpifolia and Quercus sp. (Chakrabarti et al. 1988). In northern India (Meghalaya, Uttar Pradesh), Other morphs and biology are unknown.

Eutrichosiphum quercifoliae Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae are elongate pear-shaped, pale (Raychaudhuri et al. 1973); BL 2.2-2.6 mm. Alatae have a solid brown patch on abdominal tergites 2-4. Note that the alate morph included in the original description was an ovipara, not a vivipara (R. C. Basu & Raychaudhuri 1980). Collected on Quercus sp. in West Bengal in December. Other morphs and biology are unknown. E. manoji Chatterjee, Mandal & Raychaudhuri (1981), described from Quercus sp. in Sikkim, is a synonym (A. K. Ghosh & Agarwala 1993).

Eutrichosiphum querciphaga Chakrabarti & Maity Aptera are pale except for a dark brown patch of irregular shape on the dorsal abdomen; BL 1.8-2.1 mm. On an unidentified Quercus sp. in Uttar Pradesh, India (Chakrabarti & Maity 1980). Other morphs and biology are unknown.

Eutrichosiphum rameshi (Raychaudhuri, Chatterjee & Raychaudhuri) Apterae are pear-shaped, appearance in life unrecorded, probably pale with dark brown siphunculi; BL c. 1.9 mm. On Quercus dealbata in Meghalaya, India (Raychaudhuri et al. 1977). Other morphs and biology are unknown.

Eutrichosiphum raychaudhurii (A.K. Ghosh) Apterae are pear-shaped, colour in life unrecorded; BL 2.0-2.3 mm. Described from an unidentified Quercus sp. in West Bengal, India (A.K. Ghosh 1969). Specimens identified as this species have subsequently all been collected from Alnus sp(p)., in Pakistan (Naumann-Etienne & Remaudière 1995) and northern India, and include alate as well as apterous viviparae (Raychaudhuri & Chatterjee 1974), and an alate ovipara (collected in May; Singh et al. 1980).

Eutrichosiphum roepkei (van der Goot) Apterae are pale yellowish brown, with dark apices to siphunculi; BL c.1.6 mm. In small numbers on young shoots and leaves of ?Eurya sp. attended by ants. In Malaysia (Singapore).

Eutrichosiphum russellae (A.K. Ghosh, M.R. Ghosh & Raychaudhuri) Apterae are elongate oval, dark green to blackish; BL 2.5-2.6 mm. Alate viviparae have siphunculi about 0.33 × BL. Originally collected on young twigs of Quercus sp. in West Bengal, India, attended by ants (A.K. Ghosh et al. 1971e); subsequently recorded from Q. dealbata in Meghalaya (Raychaudhuri et al. 1977), and from Castanopsis sp. (as neotattakanum, see below). Sexuales and life cycle unknown. Very similar to E. rameshi. Zhang & Zhong (1985f) described apterous specimens from Quercus sp. in China as a subspecies, russellae ssp. lijiangense Zhang. E. neotattakanum Agarwala & Ghosh (in A.K. Ghosh & Agarwala 1993) is almost identical to E. russellae and seems to be a new synonym.

Eutrichosiphum sankari Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae are pear-shaped, probably rather dark; BL 1.3-1.9 mm. Described from an unidentified host in West Bengal, India (Raychaudhuri et al. 1973); subsequently recorded on Litsea cubeba in Meghalaya by A.K. Ghosh (1976), who described the alate vivipara. Sexuales and biology are unknown.

Eutrichosiphum shiicola Takahashi Aptera are elongate, yellowish, with siphunculi distally black; BL 2.3-2.5 mm. On Castanopsis cuspidata (principal host) and Quercus dentata in Japan. Green and yellow alate males and alate oviparae occur on Castanopsis in May-June in Osaka Prefecture, but not until autumn at Tokyo (Takahashi 1962b). E. sclerophyllum Zhang (1980), described from Castanopsis sclerophylla in China, looks like a synonym.

Eutrichosiphum sikkimense (Raychaudhuri, Ghosh, Banerjee & Ghosh) Aptera are elongate oval, colour in life not recorded, probably pale; BL c. 2.7 mm. Described from a single aptera plus immatures collected on Duabanga sonneratioides in Sikkim, India (Raychaudhuri et al. 1973). Possibly this is not the normal host plant.

Eutrichosiphum simlaense L.K. Ghosh Aptera are greenish with dark siphunculi; BL 1.7-2.0 mm. On undersides of leaves and apical shoots of an unidentified Quercus sp. in Himachal Pradesh, India (L.K. Ghosh 1986). From the description this species is very similar to E. garhwalense.

Eutrichosiphum sinense Raychaudhuri Aptera are elongate oval, yellow, with siphunculi black with pale bases; BL 1.5-1.8 mm. Alate viviparae have a brown dorsal abdominal patch, and black siphunculi a little more than 0.5 × BL (Takahashi 1962b). Described from an unknown host in China (Raychaudhuri 1956), and subsequently recorded from Castanopsis spp. (indica, ferox) in China (Q. Liu et al. 2013), C. cuspidata in Japan (Takahashi 1962b, Moritsu 1983), and from C. javanica and Lithocarpus ovalis in Java (Noordam 1994). Apparently not found on Quercus, although E. narafoliae (Shinji 1922b), described from Q. serrata in Japan and not since recognised, seems closely related. Sexuales and life cycle are undescribed, but the males are winged; their genitalia were described and illustrated by Wieczorek et al. (2012). Allotrichosiphum castanopse Qiao seems to be a Eutrichosiphum, and may be this species.

Eutrichosiphum subinoyi Raychaudhuri, Ghosh, Banerjee & Ghosh Aptera are pear-shaped, colour in life not recorded; BL c. 1.4 mm. Alate vivipara elongate, BL c. 2.0 mm, with siphunculi c. 1.0 mm. Described from Pterospermum sp. in Meghalaya, India (Raychaudhuri et al. 1973). According to Raychaudhuri & Chatterjee (1980), E. subinoyi has rather few dorsal abdominal hairs (10-14 per segment) compared with related species.

Eutrichosiphum tapatii Mondal, Chatterjee & Raychaudhuri Aptera are pear-shaped, pale green with dark markings on dorsal abdomen leaving a pale central area, and dark siphunculi; BL 1.7-2.0 mm. Described from an unidentified plant in Sikkim, India (Mondal et al. 1979); subsequently apterae and immatures identified as this species were found on Quercus sp. in Uttar Pradesh (Raychaudhuri et al. 1980a). Other morphs and biology are unknown.

Eutrichosiphum tattakanum (Takahashi) Aptera are pear-shaped, darkish but with a pale spinal area on thorax and anterior abdomen and pale siphunculi with dusky apices (colour in life apparently unrecorded); BL 1.8-2.2 mm. Alate viviparae have broad dark partially coalescent bars on abdominal tergites 3-5. On undersides of leaves of Quercus spp., Castanopsis formosana, and also Castanea formosana (Takahashi 1931). In Pakistan, northern India, Nepal, Thailand, China, Taiwan and Japan. Alate males and alate oviparae are produced in June-July in Honshu (Takahashi 1962). E. neoalnicola Raychaudhuri, Ghosh & Das (1980), described from Quercus sp. in India, is very similar and possibly a synonym. [DNA analysis (Liu et al. 2015) grouped this species in a clade with Mollitrichosiphum, although it lacks the ridges on the hind tibiae that characterise that genus.]