The Aphids

A genus for two or three species related to Iziphya, but these are longer-bodied aphids with siphunculi placed further back. Dorsal hairs are mostly fan-shaped, but on abdominal tergite 8 there is consistently a pair of backwardly pointing processes bearing long rod-shaped hairs. One species (S. tuberculata) is known only from alatae trapped in Korea (Quednau & Lee 2001).

Saltusaphis scirpus Theobald Plate 4g (Fig.18i,k; Fig.19a,b) Apterae are long-bodied, greyish yellow to greenish yellow, with dark markings tending to form longitudinal bands, the dorsum being powdered with a very thin layer of greyish white wax; BL 2.3-2.5 mm. Alatae have broad dark dorsal abdominal cross bars and dark-bordered wing veins with spots at their apices, and antennae with 10-21 rhinaria on III. On various Cyperaceae (Carex, Cyperus, Scirpus). In Europe (not UK), across Asia to east Siberia, widely distributed in Africa, also now in Argentina (Ortego et al. 2006), and with a single record from USA (Ohio, Smith 1940). Monoecious holocyclic; oviparae and apterous males occur in September-October. [Saltusaphis lasiocarpae (Ossiannilsson) on Carex lasiocarpa in Sweden (Heie 1982), is a seasonal variant of S. scirpus according to Quednau (2010), and Saltusaphis kienshuensis Shinji , described from Carex sp. in South Manchuria, cannot be separated from S. scirpus on the basis of the published description.] 2n=10.

Sanbornia Baker

Aphidinae: Aphidini

One North American species on Juniperus with uniquely specialized morphology and therefore of uncertain phylogenetic position, but perhaps closest to Siphonatrophia. In addition to features mentioned in the key to juniper aphids, marginal abdominal tubercles are absent and the reduced tarsi have hairless first segments.

Sanbornia juniperi Pergande ex Baker Plate 13e, f, fig. 53C Apterae are green, with a dorsal coat of flaky, transparent wax, an unusual humpback shape (Boudreaux, 1949, and see influential points/Gallery); BL 1.2-1.4 mm. Alatae have pale brownish-green head and thorax. On Juniperus spp. (scopularum, virginiana) in USA, and also recorded from Brazil on Cupressus lusitanica (Sousa-Silva & Ilharco 2001b) and J. chinensis (Bortolotto et al. 2013). Monoecious holocyclic; oviparae and alate males occur in USA in October-November. Trioxys cupressicola may be a specialized parasitoid.

Sappaphis Matsumura

Aphidinae: Macrosiphini

A small oriental genus with Pyrus as primary hosts, related to Dysaphis but with long hairs on body and antennae. Accounts are available for Japan (Miyazaki 1971) and China (G. Zhang & Zhong 1983b). The name Sappaphis was used in a broader sense from 1952 to 1965 to include species now in Dysaphis.

Sappaphis albinae Zhang, Chen, Zhong & Li Apterae are white, BL c.1.6 mm. On roots of Artemisia scoparia in Gansu Province, China (G. Zhang 1999). Possibly these are early-summer exules of S. sinipiricola.

Sappaphis angelicograstis Zhang, Chen, Zhong & Li Apterae are golden yellow, BL c.1.5 mm. On roots of Artemisia argyi in Gansu Province, China (G. Zhang 1999). Also recorded from Angelica sinensis, but this is unlikely to be a true host. These are possibly late summer apterae and gynoparae of S. sinipiricola.

Sappaphis piri Matsumura Plate 10f (Fig.12q) Apterae in spring on Pyrus are broadly oval, brownish or yellowish-brown, covered with white woolly wax; BL 1.8-2.5 mm. Alatae have dark dorsal abdominal cross-bars, more weakly developed on anterior tergites; secondary rhinaria are distributed III 21-28, IV 4-9, V 0-1. On undersides of leaves of Pyrus spp. in east Asia (Japan, Korea, China, Taiwan, eastern Siberia). Heteroecious holocyclic, migrating in May-June to roots of Artemisia spp. Apterae on Artemisia are pale yellow to orange-red; BL 1.7-2.1 mm. The return migration to pear occurs in October. Miyazaki (1971) provided a full redescription. Kim et al. (1986) studied seasonal population fluctuations on pears in Korea, and Takada & Hashimoto (1985) studied relations with parasitoids and ants on Artemisia. [G. Zhang et al. (in G. Zhang 1999) described two new species from north-west China, S. albinae (q.v.) and S. angelicograstis (q.v.), and also listed records from Artemisia of two species previously described from Pyrus, S. dipirivora Zhang and S. sinipiricola Zhang (q.v.). These will all run to S. piri in the key to aphids on Artemisia. Further work is needed to confirm whether all or any of them are distinct from S. piri.] 2n = 12.

Sappaphis sinipiricola Zhang Described from apterae and alatae collected on Pyrus in Henan Province, China, in May, and subsequently recorded from Artemisia (G. Zhang et al. 1999a). S. albinae and S. angelicograstis (q.v.) may be seasonal forms of this species, which is itself very close to S. piri. [Gynoparae of S. sinipiricola from China have ANT PT/BASE 1.5-2.7, whereas gynoparae of S. piri from Japan and Korea have ANT PT/BASE 1.4-2.1. Another sample of alatae from pear in October, described as S. montana Zhang, in G. Zhang (1999) may also be gynoparae of S. sinipiricola.] 2n=12.

Sarucallis Shinji

Calaphidinae: Therioaphidini

One species of east Asian origin related to Tinocallis but with a squat body, a very different pattern of dorsal sclerotisation, wings that are folded almost flat over the body in repose, and a host association with Lythraceae. It was placed as a synonym of Tinocallis by Remaudière & Remaudière (1997) but revalidated as a full genus by Quednau (2003).

Sarucallis kahawaluokalani (Kirkaldy) Alatae are broad-bodied, pale yellow or yellow-green with dark brown markings (dark longitudinal stripes on head and prothorax, dark brown pterothorax, transverse marks on abdominal tergites 1 and 2 incorporating the paired tubercles and distinctively marked forewings – see aphids of Karnataka website); BL 1.2-1.8 mm. Immatures are greenish-yellow with pale brown dorsal spots. On the undersides of leaves of Lagerstroemia spp. and also recorded from Lawsonia alba (Agarwala et al. 1989a). Widely distributed in east and south-east Asia; introduced to Europe (Netherlands, France, Spain, Italy, Croatia, Greece, Bulgaria, Turkey), Iran (Gholamzadeh-Chitgar 2017), Africa (Congo; Remaudière & Le Ru 1995), USA (where it is widely distributed, including Hawaii), Central and South America (Argentina, La Rossa et al. 1997; Columbia, Kondo & Cortés 2014) Venezuela, Honduras, Panama, Costa Rica, Puerto Rico, San Salvador, Dominican Republic (Villalobos Muller et al, 2010), Guadaloupe (Étienne et al. 2018, and now reported from Australia (New South Wales; Hales & Gillespie 2020). Oviparae and alate males occur in the northern hemisphere in September-October, and in April-May in Australia (Hales 2020). Patti (1984) gave a concise general account of this species with coloured photographs. The ecology was studied in Italy by Patti et al. (1984), and in North America by Alverson & Allen (1992). Mizell & Knox (1993) studied susceptibility of numerous Lagerstroemia cultivars in Florida. An extensive search failed to reveal any specific parasitoids of this aphid (Mizell et al. 2002). 2n = 6.

Schizaphis Börner

Aphidinae: Aphidini

About 36 palaearctic and six nearctic species resembling Rhopalosiphum but with tapering siphunculi and media of forewing only once-branched. Probably about half of the species live all year on Poaceae, and most of the rest (subgenus Paraschizaphis) are on Cyperaceae and Typhaceae, but a few mainly Asian species overwinter as eggs on Pyrus or Malus. More than half the species are European, and the others occur in Middle East, Central Asia, east Asia, Africa and North America. The S. graminum group comprises several closely-related and morphologically very similar species or subspecies, some of which can only be reliably identified by their associations with different grass genera or species, or by other differences in biology. The taxonomy and life cycles of the pear-feeding species also require further study. Eastop (1961b) provided a key to the world fauna then known. Pear-feeding species were keyed by Shaposhnikov (1952), and included in regional accounts for India (A.K. Ghosh & L.K. Ghosh 2006); eastern Siberia (Pashchenko 1988a) and China (Tao 1962). Ilharco (2002) reviewed the world fauna of Paraschizaphis (treating it as a full genus), and Leclant (1967) gave an account of the “S. cyperi group”. Regional accounts are available for UK (Stroyan 1984), north-west Europe (Heie 1986), Iberian peninsula (Nieto Nafría et al. 2005a), China (Zhang et al. 1998), Japan (Miyazaki 1988) and east Siberia (Pashchenko 1988a).

Schizaphis (Paraschizaphis) acori (Shinji) Apterae are very dark green-black, with reddish tinge around bases of siphunculi; BL 1.8-2.4 mm. Alatae have secondary rhinaria distributed III 11-15, IV 2-8, V 0-1. On Acorus calamus in Japan; specimens from Cyperus difformis in Japan may also be this species. Monoecious holocyclic, with apterous and brachypterous males (Miyazaki 1988). This species has been erroneously synonymised in catalogues with S. rotundiventris. 2n=8.

Schizaphis agrostis Hille Ris Lambers Apterae are grass-green to yellowish green with darker green spinal stripe; BL 1.4-2.1 mm. Alatae have secondary rhinaria distributed ANT III 4-10, IV 0-2. On uppersides of leaves of Agrostis spp., especially canina and stolonifera, causing leaves to turn red or yellow-brown, and eventually to die off (see influential points/Gallery); sometimes on Poa annua, particularly on dry, sandy soil, sometimes ant-attended. In northern and central Europe, and aphids identified as this species have been collected on an Elymus sp. in Kurgan city (south-west Siberia; Novgorodova & Stekolshchikov 2013). Perhaps introduced to USA, as molecular and morphometric studies (Kati et al. 2013) indicate that US “biotype H” of S. graminum may be this species. Monoecious holocyclic, with oviparae and alate males in October. It is a member of the S. graminum group, all of which are very closely related. Morphometric discrimination of alatae of this species from S. holci and S. graminum were discussed by Kati et al. (2013).

Schizaphis aurea Miyazaki Apterae are bright yellow with antennal flagellum and siphunculi black, legs banded with black, and cauda pale; BL 1.6-1.9 mm. On Pennisetum alopeculoides in Japan. Monoecious holocyclic with oviparae and alate males in mid-November (Miyazaki 1988).

Schizaphis borealis Tambs-Lyche Apterae are pale, colour not recorded; BL 1.3-1.9 mm. Alatae have secondary rhinaria distributed ANT III 4-6, IV 0-1. On Phleum pratense in Fennoscandia.

Schizaphis (Paraschizaphis) brachytarsus (Takahashi) Apterae are brown with rather long black siphunculi; BL 1.4-2.3 mm. Alatae are unknown. Described from a plant of family Cyperaceae in Japan, and since recorded from Carex in Japan (neotype, described by Ilharco 2002) and recorded from Scirpus in east Siberia (Pashchenko 1988a).

Schizaphis (Paraschizaphis) caricis (Schouteden) (Fig.18f) Apterae are shiny blackish brown to blackish green; BL 1.3-2.4 mm. Alatae have secondary rhinaria distributed III 7-20, IV 2-10, V 0-5. On Carex and Scirpus, living in small compact colonies on basal parts of leaves, and often in ant “shelters”. In Europe, eastwards to Ukraine and Iran (Hodjat 1984b), and in east Siberia (Pashchenko 1988a, as caricicola) and USA (Ilharco 2002). Monoecious holocyclic, with apterous males (Heie 1986).

Schizaphis chaenometicola Zhang Apterae are yellowish green, BL c.1.5 mm. On leaves of Chaenomeles speciosa in China (Gansu Province). Other morphs have not been described (Zhang et al. 1998).

Schizaphis cuprea Miyazaki Apterae are dull brown with black antennae and siphunculi, and yellow-brown legs ringed with black; BL 1.5-1.8 mm. On Calamagrostis sp. in Japan. Monoecious holocyclic with oviparae and alate males in late September-October (Miyazaki 1988).

Schizaphis dubia Huculak Apterae are green or greenish yellow; BL 1.6-2.1mm. Described from Calamagrostis canescens, and there are also records from Agrostis stolonifera and Bromus arvensis. Alatae have 5-8 secondary rhinaria on ANT III. Pettersson (1971c) studied host preferences. In northern Europe (Finland, Poland, Sweden). A record from Turkey (Şenol et al. 2014b) requires further confirmation. Monoecious holocyclic with alate males (original description). Pettersson (1971b) provided morphometric data and illustrated four morphs.

Schizaphis (Paraschizaphis) eastopi (van Harten & Ilharco) Apterae are black, BL 1.4-2.0 mm. Alatae have secondary rhinaria distributed III 7-10, IV 1-4, V 0-1. On Typha capensis in southern Africa (Angola, Mozambique). Records of this species in the literature from other countries in Africa, and from Italy (Sicily) and Pakistan, should be referred to S. rosazevedoi (Ilharco 2002).

Schizaphis gracilis Richards Apterae are undescribed, and appearance of alatae in life is unknown; BL of alata 1.4-1.8 mm. On Carex maritima in Northwest Territories, Canada.

Schizaphis graminum (Rondani) ‘Greenbug’ Plate 9b Apterae have head and prothorax yellowish or greenish straw‑coloured, rest of thorax and abdomen yellowish green to bluish green with a darker spinal stripe (see influential points/Gallery); BL 1.3‑2.1 mm. Alatae have secondary rhinaria distributed ANT III 4-10, IV 0-4, V 0-1. On leaves of very many genera and species of Poaceae, often causing yellowing and other phytotoxic effects. It is sometimes attended by ants (Orlob 1963). Records from grasses in western Europe are now thought to apply to other closely-related species or subspecies with more specific host associations (agrostis, borealis, dubia, holci, jaroslavi, phlei, thunebergi), all of which are very difficult to tell apart (e.g. Pettersson 1971b). S. graminum is of palaearctic origin, but now widely distributed; southern Europe, Middle East, Central Asia, Africa, India, Nepal, Pakistan, Thailand, Korea, China, Taiwan, Japan, and North, Central and South America. Two alatae trapped in UK had identical mitochondrial and nuclear DNA sequence to the sorghum-adapted form in the USA (Kati et al. 2013), but there are no records from field crops in northern Europe. Records from Australia and Philippines are referable to S. hypersiphonata. A population without marginal tubercles on an unidentified grass (possibly Agrostis) in Iceland was described as a subspecies, S. graminum ssp. gigjai Stroyan, and specimens collected by sweeping in north-west Russia have also been assigned to this subspecies (Stekolshchikov & Buga 2018). The North American “biotypes” have been shown by mtDNA analysis to fit into three clades with different host relationships, the divergence of which predates modern agriculture (Shufran et al. 2000), and this is supported by analysis of nuclear sequences (Shufran 2011). This indicates that these were three separate introductions, and that palaearctic S. graminum may be a complex of still-unrecognised incipient species or subspecies (Blackman & Eastop 2007). S. graminum is monoecious holocyclic, with alate males, in cold temperate climates, e.g. in northern USA, where overwintering in the egg stage occurs predominantly on Poa pratensis (although the sorghum-adapted form is able to produce sexuales at more southerly latitudes, indicating that it probably originated from further south in Europe or Asia). S. graminum is anholocyclic wherever winter conditions permit. Transcriptome sequencing and analysis in relation to insecticide resistance has been conducted by J. Lui et al. (2019). 2n=8.

Schizaphis hierochlophaga Zhang & Chen Apterae are yellowish green; BL c. 1.6 mm. Alatae have secondary rhinaria distributed III 15-16, IV c.9, V 6-7. On Cymbopogon sp.(spp?) in Beijing, China (Zhang et al. 1998).

Schizaphis holci Hille Ris Lambers Apterae are greenish yellow or pale yellow, with dark-tipped siphunculi; BL 1.6-1.9 mm. Alatae have secondary rhinaria distributed ANT III 7-8, IV 0. On leaves of Holcus spp., usually attended by ants; infested leaves turn brown and die. In Europe (UK, Netherlands, France, Germany, Austria, Czech Republic). Monoecious holocyclic with apterous males. Kati et al. (2013) studied host specificity and sexual morph production in the laboratory, and also discussed morphological discriminants between this species, S. agrostis and S. graminum.

Schizaphis hypersiphonata A.N. Basu Apterae are yellowish‑green, BL 1.1-1.7 mm. On various Poaceae (Brachiaria, Chloris, Dactylotenia, Digitaria, Eleusine, Panicum, Phalaris, Phragmites, Triticum), and also recorded from Cyperus exaltata, but without having the phytotoxic effects of S. graminum. In West Bengal, Sri Lanka, Thailand, Philippines, Irian Jaya (Indonesia) and Australia (Eastop 1966, as a form of graminum). No sexual morphs have been recorded. Mordvilko’s (1921) description of S. graminum is possibly referable to this species.

Schizaphis jaroslavi (Mordvilko) Apterae are pale green to yellow greeen with a darker green median stripe and dark tips to siphunculi, slightly wax-powdered; BL 1.5-2.1 mm. Al. have 3-6 secondary rhinaria on ANT III only. Recorded from various grasses (Calamagrostis, Phalaris, Phleum, Secale) in Europe (Scotland, Netherlands, Denmark, Sweden, Finland, Poland, Ukraine) and Asia (Kazakhstan, Mongolia, Korea, Japan). However, the name is perhaps being applied to more than one species, as in Europe S. jaroslavi occurs on Calamagrostis epigeios in dry sandy habitats (Heie 1986), whereas Japanese populations have a shorter antennal terminal process than European ones and were only found on the wetland grass Phalaris arundinacea (Miyazaki 1988). In Korea it is recorded from a dicot, Cocculus trilobus (Lee et al. 2002c). The European aphid has apterous males, and is often ant-attended.

Schizaphis longicaudata Hille Ris Lambers Apterae are greenish, with black-tipped siphunculi; BL 1.8-2.2 mm. Alatae have secondary rhinaria distributed ANT III 6-10, IV 0-1. On grasses (Alopecurus, Apera, Bromus, Luziola, Phalaris); Pettersson (1971c) studied host preferences. Widely distributed in Europe. Monoecious holocyclic with alate (sometimes brachypterous) males (Heie 1986).

Schizaphis (Paraschizaphis) longisetosa (Higuchi) Apterae are dark green to nearly black; BL 1.9-2.3 mm. Alatae have secondary rhinaria distributed III 12-19, IV 2-7, V 0-3. On Carex spp. in Japan (Higuchi 1970). Ilharco (2002) provided a redescription, noting that the original description included specimens of S. brachytarsus.

Schizaphis mali Shaposhnikov Apterae are yellowish green with concolorous siphunculi and cauda; BL c. 2.8 mm. In spring colonies on Malus baccata in Russia (Urals), curling the leaf into a pod-shaped pseudogall, and also recorded from Kazakhstan (Kadyrbekov 2017a). Migration occurs in June to Carex spp. Appearance of apterae on secondary host is undescribed; BL c.1.8-1.9 mm. Alatae produced on Carex have secondary rhinaria distributed III 12-14, IV 2-6, V 0. 2n=8. Alate males were collected on Carex, and oviparae on Malus, in early October (Shaposhnikov 1979b). This species seems closely related to species alternating from Pyrus to Cyperaceae in Russia (S. pyri) and Japan (S. piricola). 2n=8.

Schizaphis minuta (van der Goot) Apterae are green, BL 1.2-1.8 mm. Alatae have secondary rhinaria distributed III 5-7, IV 2-3, V 0-1. On Cyperus spp. It is usually found on small, short-leaved species growing in sand, for example along dry stream beds. In Africa, Iran (Hodjat 1984b), east and south-east Asia (India to Philippines), Australia and also recorded from Scirpus lacustris in Italy (Patti 1983), and in suction trap samples in Florida, USA (Halbert et al. 2002). In Korea three samples were collected from Mazus miquelii (Lee et al. 2002c). Sexual morphs are apparently unknown.

Schizaphis muhlenbergiae (Phillips & Davis) Apterae are pale green, BL c 1.3 mm. On Muhlenbergia sp., which appears to be the normal host; also on Poa pratensis, and colonies were established on wheat under insectary conditions (original description). In USA (Indiana, Ohio, Colorado, Utah). A record from Lithuania on Leymus arenarius (Rakauskas et al. 2008) needs further confirmation. Monoecious holocyclic with apterous males and oviparae in October.

Schizaphis nigerrima (Hille Ris Lambers) ( = Schizaphis laingi Eastop; see Eastop & Blackman 2005) Apterae are coal black, BL 1.7-2.3 mm. Alatae have secondary rhinaria distributed ANT III 4-7, VI 0-1. On upper sides of leaves of Festuca pratensis, yellowing the tips, and also recorded from Alopecurus spp., Calamagrostis spp. and Elymus spp. In Europe (UK, Netherlands, Finland, Germany, Poland, Austria, Czech Republic, Hungary and north-west Russia), and also recorded from Iran (Mortazavi et al. 2015). Monoecious holocyclic with oviparae and alate males in October (original description). This species is very similar to S. cuprea in Japan.

Schizaphis nigra (Baker) ( = Toxoptera viridirubra Gillette & Palmer; see Eastop & Blackman 2005) Apterae are blackish green (original description; Washington DC) or reddish brown to olive green with orange around bases of siphunculi (original description of viridirubra, Colorado); BL 1.1-2.0 mm. Alatae have secondary rhinaria distributed III 6-14, IV 3-7. V 0-3. On Carex spp. in USA. Monoecious holocyclic, with apterous, brachypterous and alate males (original description) or alate males (viridirubra, original description). Ilharco (2002) redescribed apterous and alate viviparae (as Paraschizaphis nigra).

Schizaphis (Euschizaphis) palustris (Theobald) (Fig.35c,d) Apterae are yellowish brown to olive brown or dark green; BL 1.5-1.8 mm. Described from Triglochin palustre, and also recorded from T. maritima and Poa annua, but the more common hosts are Juncus spp. In marshy situations in Europe (UK, Netherlands, Germany, Slovakia, Spain (BMNH collection, leg. D. Hille Ris Lambers – “Franz Sp.145”), Turkey (Görür et al. 2017), Ukraine (Mamontova 1963) and India (New Delhi; BMNH collection, coll. R. Menon). Monoecious holocyclic with apterous males (Hille Ris Lambers 1947b).

Schizaphis (Paraschizaphis) pashtshenkoae (Ilharco) Apterae are dark green to almost black; BL c.2.0 mm. Alatae have secondary rhinaria distributed (1 specimen) III 15-16, IV 7, V 4. On Carex sp. in east Siberia (Ilharco 2002).

Schizaphis phlei Orlob Apterae are yellow to pale yellowish, to pale green, with siphunculi dark-tipped; BL 1.2-1.9 mm. Alatae have secondary rhinaria distributed ANT III 4-6, IV 0-1. On various grasses (Aegilops, Alopecurus, Apera, Arrenatherum, Phleum) in Sweden and Finland, with Arrhenatherum elatius as a preferred host (Pettersson 1971c, as arrhenatheri), and found causing brown leaf chlorosis on Phleum pratense in New Brunswick, Canada (original description). Monoecious holocyclic with oviparae and alate males in September (original description, and Pettersson 1971b). It is a member of the S. graminum group. Pettersson (1970, 1971a) studied the sex pheromone of this species (as borealis). Pettersson (1971b) provided morphometric data and illustrated four morphs (as arrhenatheri).

Schizaphis pilipes (Ossiannilsson) (Fig.18g) Colour of apterae in life is unknown, probably dark brownish; BL 1.6-2.4 mm. Alatae have secondary rhinaria distributed III 17-20, IV 7-13, V 2 (Heie 1986). On Carex acuta and possibly other Carex spp. in Europe. Monoecious holocyclic; oviparae were found in Sweden in November (Ossiannilsson 1959). Also recorded from Finland, Denmark, Netherlands, Germany, Czechoslovakia and UK.

Schizaphis piricola (Matsumura) Apterae are yellowish-brown or green, with dark antennae and black-tipped siphunculi; BL c. 1.4 mm. Spring colonies live in rolled edges of leaves of Pyrus pyrifolia (Essig & Kuwana 1918), and it has also been recorded from other Pyrus spp. The Schizaphis recorded from Sorbus amurensis (= S. pohuashanensis) in Korea by Paik (1972) was also probably this species. In Japan, Korea and China, and recently introduced to Spain, where it was found colonising P. calleryana, and attended by the ant Tapinoma nigerrimum (Pérez-Hidalgo et al. 2011a). Alatae are produced in May in Japan and migrate to Cyperus rotundus (Miyazaki 1988). They are also found on other Cyperus and Carex spp. Apterae on secondary hosts are yellowish green to deep green, with black siphunculi and antennae black except at bases; BL 0.96-1.85 mm. Alatae have secondary rhinaria distributed III 15-22, IV 7-11, V 0-5 (cf. rotundiventris). S. piricola is very similar to S. pyri and S. mali in Russia. 2n=8 (Chen & Zhang 1985b).

Schizaphis priori Stroyan Colour of apterae in life is unrecorded, probably pale green with darker spinal stripe; BL 2.2-2.6 mm. On Leymus arenarius in Iceland. Monoecious holocyclic, with apterous males collected in late July (original description, as S. geijskesi ssp. priori).

Schizaphis pyri Shaposhnikov Apterae are green, often with darker green longitudinal stripe, and dark siphunculi and cauda; BL 1.4-2.2mm. In spring colonies on Pyrus communis, feeding on upper sides of leaves and folding them longitudinally in half, with the underside turned outward, to form a similar pod-shaped pseudogall to that of S. rotundiventris (Shaposhnikov 1952). Presumed to be heteroecious holocyclic, alatae migrating in May to form colonies on Cyperaceae, although this still needs experimental confirmation. Apterae on secondary hosts are pale to dusky green with black antennae and siphunculi, and a dark cauda; BL 1.5-2.0 mm. They live singly on the undersides of leaves of Carex (Leclant 1967, as fritzmuelleri) and Cyperus. Alatae produced on Carex in France had secondary rhinaria distributed III 20-29, IV 7-15, V 4-5. In continental Europe (France, Spain, Portugal, Czech Republic, Moldova, Russia, Ukraine), Atlantic islands (Azores, Canaries, Madeira), and also recorded from Israel, Turkey, Iran, Kazakhstan, west Siberia, east Siberia (Pashchenko 1988a), and Australia (Victoria). Sexual morphs have not been described; secondary host populations are more widely distributed and are probably mostly anholocyclic.

Schizaphis (Paraschizaphis) rosazevedoi (Ilharco) Apterae are of two colour-forms, orange-brown to rusty brown, or dark green to almost black; BL 1.3-2.4 mm. Alatae have secondary rhinaria distributed III 6-16, IV 0-6, V 0-2. On Typha spp. and Strelitzia reginae. Recorded from Portugal, Madeira, France, Sicily, Israel, Iran, Pakistan, Saudi Arabia, Sudan, Burundi, South Africa and Martinique. (Some of these records were as misidentified S. eastopi). Apparently anholocyclic (Ilharco 2002). 2n=8.

Schizaphis rotundiventris (Signoret) Apterae on Pyrus are greyish or blackish, dusted with white wax, with black siphunculi and cauda; BL c. 1.9 mm. Feeding by fundatrices on developing leaves of Pyrus communis in early spring prevents them from unfolding and causes the edges of the leaves to stick together forming a slightly inflated pod-shaped pseudogall (Das 1918, as S. punjabipyri). Pear-feeding generations are recorded from northern India, Pakistan and Nepal. In Pakistan they are common in the plains and lower hills, especially in humid, monsoon-exposed areas (Naumann-Etienne & Remaudière 1995). Alatae migrate in May to Cyperaceae. Colonies occur most commonly on Cyperus spp., although it can form small colonies in young unfolding leaves of palms such as Elaeis guineensis, and there are also records from Poaceae (Pennisetum, Polytrias)and other monocots (Acorus, Typha). Apterae on secondary hosts are dark green to almost black (see aphids of Karnataka website); BL 1.2-1.7 mm. Alatae have secondary rhinaria distributed III 6-18, IV 0-10, V 0-5. Presumed anholocyclic populations on Cyperus and other monocots are recorded from southern Europe, the Middle East, Africa, Mauritius, east and south-east Asia, Australia, New Zealand, Hawaii, New Caledonia (Mille et al. 2020), south-eastern USA (Halbert et al., 2000) and Costa Rica (Sánchez-Monge et al. 2010). In Pakistan gynoparae and alate males return to Pyrus in early December or earlier, oviparae laying eggs in late December-January which hatch in February-March. [The synonymy with S. punjabipyri (Das) proposed by Shaposhnikov (1979a) is followed here, but still requires experimental confirmation.] 2n=8.

Schizaphis rufula (Walker) Apterae are yellowish green with darker green spinal stripe, slightly wax-powdered, with dark- tipped siphunculi; BL 1.4-3.1 mm. Alatae have secondary rhinaria distributed ANT III 4-8, IV 0-2. Pettersson (1971b) provided morphometric data and illustrated four morphs. On leaves of various grasses, especially in sand-dune habitats (Aegilops, Agropyron, Agrostis, Alopecurus, Apera, Ammophila, Bromus, Elymus, Leymus). Infested leaves become yellowish brown. Pettersson (1971c) studied host preferences, and Van Moorleghem & de la Peña (2016) studied the effects of its feeding on various dune grasses. Records from Salsola, including the original description, are presumably casual occurrences. In north-west and northern Europe, and also recorded from Iran (Rezwani et al. 1994) and Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic with apterous males.

Schizaphis (Paraschizaphis) scirpi (Passerini) Plate 9c Apterae are shiny dark bronze-brown to reddish brown or blackish (see influential points/Gallery); BL 1.5-2.8 mm. Alatae have secondary rhinaria distributed III 7-16, IV 0-8, V 0(-1). In ant-attended colonies at leaf-bases of Typha and Sparganium, also on Cyperaceae (Carex, Eriophorum, Scirpus) and sometimes on Araceae (Colocasia), Juncaceae (Juncus) or Iridaceae. Throughout Europe, and in south-west and central Asia, China, Korea and Japan. Monoecious holocyclic, with alate males (cf. S. caricis). European populations on Eriophorum spp. are regarded as a subspecies, S. scirpi ssp. eriophori Müller. 2n=8.

Schizaphis (Paraschizaphis) scirpicola Hille Ris Lambers Colour of apterae in life is unknown; 1.4-1.7 mm. Alatae have secondary rhinaria distributed III 9-10, IV 3, V 0 (1 specimen, redescribed by Ilharco 2002). On Scirpus atrocinctus in Canada (New Brunswick). It has subsequently been recorded from Scirpus spp. and Carex sp. in Ontario, Nova Scotia, Quebec and Manitoba, and from North Carolina, USA.

Schizaphis thunebergi Heie Apterae are pale, colour in life unknown; BL c.2.1-2.3 mm. The single known alata has 6 secondary rhinaria on ANT III only. On Calamagrostis purpurea in Finland.

Schizaphis variegata Ossiannilsson Apterae are broadly oval, mottled dark green and brown, BL c.1.8-2.0 mm. On Carex juncella in Sweden (Ossiannilsson 1959). Alatae and biology are unknown.

Schizaphis wahlgreni (Ossiannilsson) (Fig.18h) Colour of apterae in life is unknown, probably pale, with black antennae (except at base) and siphunculi, and a pale cauda; BL c.2.1 mm. On Carex vesicaria in Sweden, Poland, Czech Republic and Hungary (Heie 1986), and also found on Eleocharis palustris in Latvia (Rupais 1989a). Alatae have been recorded (Ossiannilsson 1964b), but are undescribed.

Schizaphis weingaertneriae Hille Ris Lambers Apterae are dark green powdered with bluish grey wax; BL 1.2-1.7 mm. Alatae have secondary rhinaria distributed ANT III 5-8, IV 0-2. On undersides of leaves of Corynephorus canescens, usually arranged in single file. In Netherlands, Poland and Czechoslovakia. Oviparae and apterous males were collected in early October (original description). It is not visited by ants.

Schizaphis werderi Börner Colour of apterae in life is unrecorded, BL 1.6-1.9 mm. On Poa alpina in the Alps.

Schizolachnus Mordvilko

Lachninae: Eulachnini

NOTE: Schizolachnus has long been treated as a separate genus, related to Cinara but with some distinctive morphological and biological features. However, molecular phylogenetic studies combining both nuclear and mitochondrial data by R. Chen et al. (2016) clearly placed Schizolachnus as a group within Cinara, and and this was further supported by the work of Havelka et al. (2020). We therefore now follow Aphid Species File in treating Schizolachnus as a subgenus of Cinara.

Schizoneuraphis van der Goot

Hormaphidinae: Nipponaphidini

Eight species are known, at least one of them having host alternation between Distylium galls and Litsea (in Java; Noordam 1991). Closely related to Nipponaphis, but the aleyrodiform exules have long, basally thick dorsal (prosomal) hairs.

Schizoneuraphis gallarum van der Goot Galls on Distylium stellare in Java are oval, balloon-shaped, 25-50 mm long, apparently arising from the stem but formed from the leaf (fig. 130J); pale green to peach-coloured when young, becoming dark grey-brown and woody, but with a thin circular window in the side which becomes the emergence hole for alatae in August-October. Emigrant alatae (BL 1.6-2.2 mm) have secondary rhinaria distributed III 17-22, IV 8-11, V 4-8. Apterous exules feeding along the veins on the undersides of leaves of Litsea glutinosa are aleyrodiform, shiny dark brown to black, with upright pale brown dorsal hairs and a white marginal band of wax; BL 0.7-1.0 mm. Apterae on L. glutinosa in China (Hong Kong; Qiao et al. 2018) and on Litsea sp. in Australia (Queensland; Brumley 2020) have also been identified as this species. The host alternation still needs to be confirmed experimentally. Hille Ris Lambers & Takahashi (1959) claimed to have established it on the basis of the similarity of embryos within emigrant alatae to first instars on Litsea, but Noordam (1991) noted some differences.

Schizoneuraphis himalayensis (Ghosh & Raychaudhuri) Apterae are broadly oval, dark brown to black; BL 0.9-1.1 mm (A.K. Ghosh & Raychaudhuri 1973b, as Nipponaphis). In colonies on leaves and stems of Litsea spp., Lindera sp. and Persea bombycana (= Machilus gamblei) in India (Assam, West Bengal, Manipur). Alatae (sexuparae?) were produced in January. The life cycle is unknown. Nipponaphis machiliphaga of A.K. Ghosh et al. (1971d) seems to be this species. A record from Quercus dilatata (Holman 2009) needs verification.

Schizoneuraphis litseicola Noordam Apterae on the undersides of leaves of Litsea amara (= umbellata) are aleyrodiform, shiny brown-black, similar in life to S. gallarum; BL 0.8-0.9 mm. Only known from Java (Noordam 1991). The life cycle is unknown. The description is very similar to that of S. machiliphaga (Takahashi), which has also now been recorded from Litsea. Schizoneuraphis siamensis (Takahashi) described as shining purplish brown without wax and feeding along mid-rib on the undersides of leaves an unidentified tree of Lauraceae in Thailand (Takahashi 1941a, as Thoracaphis), could be this or a very similar species.

Schizoneuraphis longisetosa Noordam Galls on Distylium stellare in Java are not properly described, but believed to be semiglobular, turning red when mature (possibly gall no. 487 of Docters van Leeuwen-Reijnvaan & Docters van Leeuwen 1926). Emigrant alatae (BL 2.4-2.8 mm, secondary rhinaria distributed III 25-31, IV 12-16, V 8-13) were collected in late May (Noordam 1991). Secondary host populations are unknown.

Schizoneuraphis machiliphaga (Takahashi) Apterae are almost circular, dorsally flattened, reddish-brown, not strongly sclerotised; BL 0.8-1.0mm. On the undersides of leaves of Machilus sp. in Japan (Takahashi 1959b), and recorded from Litsea polyantha and Litsea sp. in India (Assam, West Bengal, Meghalaya; A.K. Ghosh 1988, as Nipponaphis). The life cycle is unknown.

Schizoneuraphis malayna (Takahashi) Apterae are aleyrodiform, almost circular, dorsally flattened, shiny brownish-black, without evident wax; BL c. 1.3mm. On undersides of leaves of Quercus sp. in Cameron Highlands of Malaya (Takahashi 1950, as Thoracaphis malaynus). The life cycle is unknown.

Schizoneuraphis querciphaga (Ghosh and Raychaudhuri) Apterae are yellowish to dark brown; BL 1.3-1.5 mm. On apical stems of Quercus sp., attended by ants, in Sikkim, India (A.K. Ghosh and Raychaudhuri 1973b), and also on Castanopsis in Nepal (Miyazaki 1977). The life cycle is unknown. Very similar to S. malayna.

Schizoneurata Hille Ris Lambers

Eriosomatinae: Eriosomatini

One little-known North American species closely related to Eriosoma but lacking siphunculi in morphs developing on Crataegus, the presumed secondary host. The primary host is unknown, but presumed to be Ulmaceae (Hille Ris Lambers 1973c).

Schizoneurata tissoti Hille Ris Lambers Only immatures and alate sexuparae are described, and its appearance in life is unknown. Apterae and immatures are presumably wax-covered. Sexuparae have BL 1.6-2.2 mm and secondary rhinaria distributed III 15-20, IV 2-5, V 1-3, VI 0-1. On Crataegus uniflora and Crataegus sp. in Florida, USA (Hille Ris Lambers 1973c).

Schizoneurella Hille Ris Lambers

Eriosomatinae: Eriosomatini

One species in the Himalayan region, with empodial hairs reduced (in alatae) or absent (in apterous exules); related to Eriosoma but without siphunculi in the morphs on the secondary host plant, and distinct from the nearctic genus Schizoneurata because it has embryos with spinulose hind tarsi, as in palaearctic Eriosoma (Schizoneura) (Hille Ris Lambers 1973b). A cladistic study by Sano & Akimoto (2011) placed it close to Aphidounguis. [A second species described in this genus, Schizoneurella gei Bozhko, was synonymised with Eriosoma japonicum by Eastop & Blackman (2005).]

Schizoneurella indica Hille Ris Lambers Galls are of the open leaf-roll type, on leaves of Ulmus villosa in India (Kashmir, Jammu) and Pakistan, and possibly China (see below). Bhagat (1982b) reported that it formed clavate, asymmetrical pouch galls on U. wallichiana, but this was probably based on a misidentification (S. Chakrabarti, cited in Sano & Akimoto 2011). Alatae (BL 1.8-2.3 mm) (fig.118B) are produced in May-June. Host alternation occurs to Malus pumila, where colonies live, presumably in woolly wax, on the roots. Qiao et al. (2005d) described apterous and alate exules collected in September-October from Malus in China, and reported that their root feeding may cause considerable economic damage. [However, to judge from alatae and a single aptera representative of this material that have been deposited in the BMNH collection, the identity of these aphids is questionable. The aptera with 4-segmented antennae seems to be Aphidounguis mali; it has wax gland plates of the form typical of that species. Its tarsi all have only one claw fully developed, and embryos within the abdomen have tarsi with the long pointed single claws typical of apterous exules of A. mali. The alatae (collected on sticky bands around the trunks of the apple trees) are sexuparae of a species of Eriosomatinae, not Aphidounguis, and may be those of S. indica, although the differences from the spring migrants from Ulmus galls are not those usually found between these two morphs in one species, so further confirmation seems necessary.]

Bhagat (1982b) reported a parasitoid and hyperparasitoid of the gall-feeding generations (but note that the identity of the aphid host requires confirmation).

Schlechtendalia Lichtenstein

Eriosomatinae: Fordini

Two east Asian species related to Melaphis, and likewise having host alternation between Rhus and mosses, but alatae emerging from the galls on Rhus have an elongate, curved pterostigma. In the light of recent molecular studies by H.C. Zhang & Qiao (2007) and Yang et al. (2010), as well as numerical taxonomic analysis by Yang et al. (2008), we have reinstated the genus Meitanaphis Tsai & Tang for two of the three species (elongallis and microgallis) treated under Schlechtendalia in Blackman & Eastop (1994).

Schlechtendalia chinensis (Bell) Plate 6f Chinese Gall-nut Aphid Galls (‘Chinese gall-nuts’) are greenish-yellow, densely covered with short, fine hairs, ovoid, becoming woody and irregular in shape with horn-like protrusions, varying from 1 to 6 cm in maximum length (fig. 133). On Rhus chinensis (= javanica, semialata) in China, Japan, Korea, Taiwan and Malaysia. Heteroecious holocyclic, with an unusual one-year cycle; after several generations in the gall, emigrant alatae (BL c. 1.5mm) are produced in October and fly to found overwintering colonies on mosses (Mnium, Plagiomnium). The appearance of the overwintering immature stages on mosses is undescribed. Alatae sexuparae return to Rhus in spring to produce sexuales. Sexual females live in bark crevices for three weeks after mating and then each produces a single fundatrix larva ovoviviparously – a phenomenon not known in any other aphid (Tang 1976; Takada 1991a). The galls have been collected for many years; they have been used in tanning leather, in the preparation of dyes and for medicinal purposes (including a treatment for haemorrhoids; Gardener 1981). Takada (1991a) and P. Liu et al. (2014) studied gall development, and Shao et al. (2013) made a detailed study of the generations developing in the gall. Qiu et al. (1999) examined the influence of environmental factors on development of sexual females. Wang et al.(2008), Ren et al. (2008) and Lv et al. (2010) examined genetic diversity in and between populations. H.-X.Wu et al. (2018) reported large differences in the symbiotic bacteria associated with populations in sumac galls and on mosses. The complete mitochondrial genome has been published (Ren et al. 2016). 2n = c. 36.

Schlechtendalia peitan (Tsai & Tang) was previously treated as a synonym of S. chinensis, but evidently the galls of this species do not develop horn-like protrusions (see Fig. 4d in C. Zhang et al. 2008), and there are differences in the relative lengths of the antennal segments of the emigrant alatae (Yang et al. 2009). On Rhus chinensis and R. punjabensis var. sinica in China.

Schoutedenia Rübsaamen

Greenideinae: Schoutedeniini

Two species on woody Euphorbiaceae/Phyllanthaceae belonging to an ancient group (also including Eonaphis and Paulianaphis) of Gondwanaian distribution. Remaudière (1988, 1990) reviewed the genus, synonymising most of the described species. DNA barcoding also failed to distinguish the two currently recognised species (Q. Liu et al. 2013); further work using nuclear markers is needed. A.K. Ghosh & Agarwala (1993) reviewed the genus in India.

Schoutedenia emblica Patel & Kulkarni Apterae are yellow-green to green (see aphids of Karnataka website); BL 1.4-1.6 mm. In ant-attended colonies, only on Phyllanthus emblica (= Emblica officinalis) in India, Pakistan, Nepal, Thailand and China (D. Zhang et al. 2006). It was recorded from Spiraea sp. in western Himalaya (Bhattacharya & Dey 2001, as ssp. andhraka), but the host is unlikely and there are errors in their morphometric data. Monoecious holocyclic; in southern India, apterous (brachypterous?) males and oviparae are produced in August (David & Hille Ris Lambers 1956; as ssp. andrhaka).

Schoutedenia ralumensis Rubsaamen Plate 10e, f Apterae are lemon-yellow or green; BL 1.4-1.8 mm. Alatae have thickly black-bordered forewing veins. On young shoots, leaves and stems of woody Euphorbiaceae/Phyllanthaceae (Breynia, Bridelia, Fluggea, Glochidion, Phyllanthus), attended by ants. Records from other plants are probably all of vagrants. Widespread in south-east Asia; India; Africa; and down the east coast of Australia. Monoecious and partially holocyclic; oviparae and alate males occur concurrently with parthenogenetic generations throughout summer and autumn. Presumably all eggs laid enter diapause, as fundatrices occur only in spring (Hales & Carver 1976). The oviparae are unusual in having scent glands on the hind femora. The male genitalia were described and illustrated by Wieczorek (2012). Relations with predators and parasitoids in Australia were studied by Hales & Carver (1976) and genetic (allozyme) variation was studied by Tomiuk et al. (1991). 2n = 16 (female), 14 (male) or !5* (sex unknown; Papua New Guinea, leg. J.H. Martin); the highly anomalous spermatogenesis of this aphid was studied by Hales (1989).

Scleromyzus Basu, Ghosh & Raychaudhuri

Aphidinae: Macrosiphini

A genus for one Indian species on Corylopsis with a spinulose head but differing from Myzus in having weakly-developed antennal tubercles and post-siphuncular sclerites (R.C. Basu et al. 1976).

Scleromyzus corylopsis (Basu, Ghosh & Raychaudhuri) Colour in life unrecorded, BL of apterae 1.6-2.5 mm. On unidentified Corylopsis sp(p). in India (Shillong, Assam; R.C. Basu et al. 1973, as Myzus). Other morphs and life cycle are unknown.

Semiaphis van der Goot

Aphidinae: Macrosiphini

About 16 palaearctic species similar to Hyadaphis but with very short siphunculi. They are typically heteroecious between Lonicera and Umbelliferae/Apiaceae, or live without host alternation on one or other of these host‑plant groups. A few species occur on other plants, including two on Impatiens. The genus is not well-known biologically and the life cycles of many species are not completely known. Populations on Lonicera are particularly difficult to assign to species. Accounts are available for north-west Europe (Heie 1992, Blackman 2010), central Europe (Heinze 1960), European Russia (Shaposhnikov 1964), Siberia (Ivanoskaya 1977; Pashchenko 1988a), and Japan (Miyazaki 1971).

Semiaphis aizenbergi (Narzikulov) Apterae are pale green, wax-covered; BL c. 1.7 mm. On Lonicera spp. in Central Asia, and also reported from west Siberia (Ivanoskaya 1977). Oviparae and alate males occur in November (Mukhamediev & Akhmedov 1982).

Semiaphis anthrisci (Kaltenbach) Apterae are dirty green, wax-powdered; BL 1.8-2.2 mm. Recorded from several genera of Umbelliferae/Apiaceae (Aegopodium, Anthriscus, Peucedanum, Torilis), slightly rolling the leaves. In north-west, northern, central and eastern Europe, Iran, Kazakhstan (Kadyrbekov 2012a) and west Siberia (Heie 1992). Differences from several other species (dauci, pastinacae, pimpinellae, sphondylii) are minimal, and therefore host records must be treated warily; possibly the true host range is more restricted. Heikinheimo (1997) gave additional data to distinguish anthrisci from dauci and pimpinellae on the basis of hair lengths. Monoecious holocyclic; oviparae were collected in September on Peucedanum palustre. Stekolshchikov et al. (2008b) provided detailed descriptions of males and oviparae.

Semiaphis cervariae (Börner) (Fig.41c) Apterae are pale yellow-ochre to bright greenish yellow, with fine wax powder; BL 1.9-2.3 mm. On Peucedanum spp., causing yellow and red spotting, and desiccating terminal parts of leaflets. In Europe (Austria, Gemany, Netherlands, Switzerland).

Semiaphis coniumi Bozhko Apterae are greenish with brown legs and dark green siphunculi and cauda; BL c.1.9 mm. On Conium maculatum in Ukraine.

Semiaphis coryspermi Mamontova ex Shaposhnikov Apterae are greenish white, with waxy bloom; BL c.1.5 mm. On Coryspermum sp., in leaves rolled into tubes. In Ukraine, and also reported from Russia (Volga region) and Kazakhstan (Kadyrbekov 2017a, as Hayhurstia coryspermi). Mamontova (1979) provided a more detailed description.

Semiaphis dauci (Fabricius) Plate 13h Apterae are pale blue-green with dark brown head and waxy bloom (see influential points/Gallery); BL 1.3-2.1 mm. On wild and cultivated Daucus carota, feeding on upper sides of rolled young leaves and leaflets in spring, and later in umbels. It is also recorded from Aegopodium and several other genera of Umbelliferae/Apiaceae. In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), the Middle East and Central Asia (south-east Kazakhstan; Kadyrbekov 2005c). Records from USA require confirmation (Foottit et al. 2006), although a carrot-feeding species of Semiaphis does occur there (Skvarla et al. 2017). Monoecious holocyclic (with alate males) on Daucus, or anholocyclic. Similar aphids on Seseli spp. are currently regarded as a subspecies, S. dauci ssp. seselii Börner.

Semiaphis heraclei (Takahashi) (Fig.10k, Fig.38d) Apterae are pale grey-green to yellow-green, wax-dusted, with antennae, legs, siphunculi and cauda rather pale; BL c.2.1 mm. On stems or curled leaves of various Umbelliferae/Apiaceae (Angelica, Apium, Bupleurum, Carum, Changium, Cnidium, Coriandrum, Cryptotaenia, Daucus, Glehnia, Heracleum, Ligusticum, Oenanthe, Osmorhiza, Seseli, Torilis) in east and south-east Asia, India and Pakistan, Algeria (Laamari et al. 2013), and Hawaii. Heteroecious holocyclic with Lonicera spp. as primary hosts in Japan, China and east Siberia. Hyadaphis albus (Monzen), described from Angelica acutiloba in Japan, is possibly this species. 2n=8. [Leycesteria formosana is possibly an alternative primary host in India (Pal & Khuda Bukhsh 1983), but as these authors recorded 2n=10 they possibly had a different species.]

Semiaphis horvathi Szelegiewicz (Fig.41d) Apterae are pale brown, lightly wax-powdered; BL 1.6-2.0 mm. Described from Peucedanum officinale, colonising shoot tips and deforming leaves. In Hungary, also in Czech Republic (BMNH collection, leg. H.L.G. Stroyan) and recorded from P. morisonii and several Ferula species in Kazakhstan (Kadyrbekov 2009b, 2014d), and on P. alsaticum in Russia (Altai; Kadyrbekov 2014g).

Semiaphis iliensis Kadyrbekov Apterae are yellow-green; BL 1.2-1.7 mm. On Lonicera iliensis in Central Asia.

Semiaphis longissima Narzikulov Apterae are green, wax-dusted; BL 1.8-2.2 mm. On Lonicera microphylla in Central Asia. Oviparae and alate males occur in October-November (Mukhamediev & Akhmedov 1982).

Semiaphis moiwaensis Takahashi Apterae are white with pale appendages; BL c. 2mm. On stem and roots of Impatiens noli-tangere in Japan, and Korea (BMNH collection, leg. W.H. Paik). This is likely to be a synonym of S. nolitangere.

Semiaphis nolitangere Aizenberg (Fig.33e) Apterae are pale yellowish green; BL 1.8-2.3 mm. On undersides of young apical leaves of Impatiens noli-tangere, which become slightly rolled, and later on flower stalks (Heie 1995). In Finland, northern Russia (including west Siberia) and Kazakhstan. Heteroecious holocyclic with oviparae and alate males in September in Finland, where transfers were made to the primary host Lonicera tatarica, causing yellowish spots on leaves (Heikinheimo, cited by Heie 1995).

Semiaphis pastinacae Börner Apterae are probably grey-green to yellow-green, with dark antennae, legs, siphunculi and cauda; BL c.1.3 mm. In downwardly rolled leaflets of Pastinaca sativa in central Europe, (Austria, Czech Republic, Switzerland). Host alternation probably occurs with Lonicera spp. as primary hosts, but this needs confirmation (Heinze 1960).

Semiaphis pimpinellae (Kaltenbach) Apterae are dull green mottled with darker green, coated with greyish wax powder; BL 1.4-1.8 mm. On Pimpinella spp., esp. saxifraga, causing leaves to curl upwards, and later in season on umbels. Ossiannilsson (1959) recorded it from Aegopodium podagraria. In Europe (not Iberian peninsula), and Iran (Rezwani & Parvizi 1990). It is very similar to other species in the S. anthrisci group.

Semiaphis rabotkinae Pashchenko Apterae are green, wax-dusted; BL c. 2.2 mm. On Lonicera edulis in east Siberia.

Semiaphis sphondylii (Koch) (Fig.10j) Apterae are greenish white, wax-powdered; BL 2.0-2.5 mm. In rolled leaves on Lonicera nigra in spring, heteroecious holocyclic, migrating to various Umbelliferae/Apiaceae (Angelica, Heracleum, Peucedanum), although some host records may be due to confusion with other similar species such as anthrisci and pimpinellae. In north-west, northern and central Europe (Finland, Latvia, Netherlands, Germany, Austria), west Siberia (Stekolshchikov & Khureva 2020), and also reported to occur in Iran (Rezwani 2010), and in Kazakhstan (on Lonicera tatarica, Bupleurum and Libanotis; Kadyrbekov 2009a,b).

Seneciobium Remaudière

Aphidinae: Aphidini

A genus for one Aphis-like species in south-west Europe completely lacking siphunculi. It has dark legs and antennal sensoriation characteristic of a group of Aphis (incl. jacobeae in Europe and lugentis in North America) that feed on Senecio, which suggests that, despite lacking siphunculi, its separate generic status may not be warranted.

Seneciobium balachowskyi Remaudière Apterae are greenish to black, with a waxy bloom, and appendages all dark, including cauda; BL 1.8-2.2 mm. On stems and flowers of Senecio adonidifolius in the eastern Pyrenees. Monoecious holocyclic with apterous males.

Sensoriaphis Cottier

Taiwanaphidinae

Four species on Nothofagus in Papua New Guinea, Australia and New Zealand, and one on Melaleuca in Western Australia, where Nothofagus occurs only in fossil form. Sensoriaphis is closely related to Taiwanaphis, and regarded by some authors as a subgenus or even a synonym of that genus, but we follow Remaudière & Remaudière (1997) in giving it separate generic status, as it forms a distinct group with biological as well as morphological differences. Carver (1978) provided a key to species, and Quednau (2010) reviewed the genus (as a subgenus of Taiwanaphis) and provided illustrations of all available morphs.

Sensoriaphis furcifera Carver & White (fig. 62A) Alatae have a pale green abdomen, head and pronotum with two longitudinal dark stripes, and mainly black antennae and legs; BL 1.8-2.8 mm. On tender young shoots and suckers of Nothofagus moorei in New South Wales, Australia. Carver & Hales (1974) described all morphs and gave life cycle details; monoecious holocyclic, with oviparae and alate males occurring concurrently with alate viviparae through spring, summer and autumn. Eggs are flat, colourless and wax-covered, and are laid on leaves.

Sensoriaphis melaleucica (Quednau) Colour of apterae in life unknown; BL 1.4-2.2 mm. On Melaleuca spp. (lanceolata, teretifolia) in Western Australia (Quednau 2010). The life cycle is unknown.

Sensoriaphis niuginii Carver (fig. 62B) Both apterae and alatae occur, colour in life not recorded; BL of aptera 1.5-2.2 mm, of alata 2.0-2.4 mm. On undersides of young leaves of Nothofagus carrii in Papua New Guinea (Carver 1978). The life cycle is unknown.

Sensoriaphis nothofagi Cottier (fig. 62D) Apterae are yellowish with brown markings (see influential points/Gallery); BL 1.1-1.5 mm. Well camouflaged, feeding on young stems of Nothofagus spp., generally on trees close to forest edges. Indigenous to New Zealand, where it is rather common and widespread (Teulon et al. 2013). Alatae become mature on the undersides of leaves. Monoecious holocyclic, with oviparae and alate males in September to February (BMNH collection, leg. V.F. Eastop). 2n = 10.

Sensoriaphis tasmaniae Carver & Martyn (fig. 62C) Apterae have a dirty yellow head and prothorax, rest of body olive green and rather shiny (see influential points/Gallery); BL 1.5-2.0 mm. On stems of young shoots of Nothofagus cunninghami in Tasmania (Carver & Martyn 1962). The alatae morph was described by Carver & Martyn (1965). The life cycle is unknown.

Serratocallis Quednau & Chakrabarti

Calaphidinae: Myzocallidini

One Asian oak-feeding species, related to Hoplochaetaphis but with spiculose dorsal hairs and clusters of spinal papillae on the thorax. Quednau (1999) provided illustrations of all available morphs.

Serratocallis takahashii Quednau & Chakrabarti Apterae are brown; BL 1.6-2.3 mm. Alatae have a short broad pterostigma with a black spot at its base. On twigs, leaf petioles and undersides of leaves of Quercus spp. (baloot, dealbata, oblongata, semicarpifolia) in Afghanistan, Pakistan (Naumann-Etienne & Remaudière (1995) and India (Himachal Pradesh, Uttar Pradesh; A.K. Ghosh & Quednau, 1990). Sexual morphs are unknown.

Shenahweum Hottes & Frison

Drepanosiphinae

One North American species on Acer, related to Drepanaphis and considered as a subgenus by some authors; the differences were discussed by Smith & Dillery (1968), and the case for regarding it as a separate genus was convincingly argued by Wieczorek et al. (2013c).

Shenahweum minutum (Davis) (fig. 22D) All viviparae are alate, with brown or black thorax and very pale greenish-yellow to whitish abdomen; BL 1.0-1.25 mm. Immatures are pale greenish-yellow. On the undersides of leaves of Acer saccharum in eastern North America. Oviparae and alate males collected in Maine and North Carolina in November were described by Wieczorek et al. (2013c).

Shinjia Takahashi

Aphidinae: Macrosiphini

One east Asian species with greatly reduced tarsi, host-alternating between Viburnum and ferns. There is a rather strange parallel with a fern-feeding North American species in another monotypic genus, Mastopoda pteridis, with similarly vestigeal tarsi and apparently with similar biology, but not closely related.

Shinjia orientalis (Mordvilko) Plate 22c (Fig.43a) Apterae are shining yellow with black antennae and dark tips to siphunculi; BL 1.3-2.0 mm. Alatae have paired brown patches partially fused across midline. On young growth of Viburnum spp. in spring. Heteroecious holocyclic, migrating to ferns (Athyrium, Eriosorus, Polypodium, Pteridium, Pteris) in Japan (Sorin 1962b, as pteridifoliae). Recorded from east India, Nepal, China, Korea, east Siberia, Japan, Philippines and Australia. It is probably anholocyclic on ferns in Australia (Eastop 1966). 2n=12*.

Shivaphis Das

Calaphidinae: Shivaphidini

Seven Asiatic species associated with Celtis and Pteroceltis. The Indian species were reviewed by A.K. Ghosh & Quednau (1990) and the Chinese species by W. Zhang et al. (1994) and Qiao et al. (2005b). Quednau (2003) reviewed the genus and provided a key to species and illustrations of all available morphs.

Shivaphis catalpinari Quednau & Remaudiere Apterae are green, with long wax filaments (W. Zhang et al. 1994, as S. similicelti); BL of aptera 1.4-2.1 mm, of alata 1.5-1.9 mm. On Celtis australis in Turkey, and on Celtis koraiensis and C. sinensis in China (Quednau 2003, Qiao et al. 2005b). Oviparae and alate males occur in Turkey in October (Quednau & Remaudiere 1985).

Shivaphis celti Das Plate l0c, d Apterae are pale to dusky greenish, sometimes pinkish, covered in bluish-white flocculent wax (see Halbert & Choate 2003); BL 1.9-2.6 mm. Alatae also wax-covered, with forewing veins thickly bordered with fuscous distally and large black patches on pterostigma and distal part of Cu_1b. On undersides of leaves or on shoot tips of Celtis spp. in Turkey, Iran, Central Asia, India, Pakistan, Nepal, Sri Lanka, Korea, Japan, Loochoo, China, Taiwan and Philippines. S. celti was introduced into south-eastern USA in the late 1990s (Halbert & Choate 2003) and had spread to California by 2002. It has more recently been found in Australia (2013, in New South Wales; Carnegie & Nahrung 2019) and South Africa (Giliomee et al. 2018). Das (1918) gave a full account of its biology and natural enemies in Pakistan; oviparae and alate males occurred in December, eggs were laid in January and hatched in March. In Japan, Shibata (1955; as S. celticolens) obtained sexuales in September-November. In Taiwan, S. celti is mainly anholocyclic, but produces some sexuales in December-January. In Australia it seems to be completely anholocyclic (Hales 2020). DNA barcoding studies in southern China (Q. Li et al. 2020) indicate that there may be geographic races or cryptic species. 2n = 10 (Hong Kong; Blackman 1986). [However, Shinji (1931) provided clear images of spermatocytes with n=3, indicating that there may be a separate species in Japan with 2n=6.]

Shivaphis (Sinishivaphis) celticola (Nevsky) All viviparae are alate, pale green to greenish-yellow covered with bluish wax; BL 1.8-2.2 mm. Forewings are marked with fuscous as in S. celti, except that there is a band joining the first fork of the media and Cu_1a (A.K. Ghosh & Quednau 1990). On the undersides of leaves of Celtis caucasica (= australis), recorded from Tadzhikistan, Uzbekistan, Kazakhstan, Pakistan and India (Kashmir). Oviparae and alate males occur in October-November; Narzikulov (1961a) gave a detailed account of the life cycle in Tadzhikistan, with descriptions of all morphs.

Shivaphis (Sinishivaphis) hangzhouensis (Zhang & Zhong) Only alate viviparae are known; yellow, covered with wax, with very thick bands of fuscous along forewing veins, BL 1.9-2.3 mm. Described from Osmanthus asiaticus in China (Zhang & Zhong, 1982c), but subsequently found on the undersides of leaves of more likely hosts, Celtis julianae and C. sinensis (Quednau 2003). Quednau illustrated an ovipara collected on C. sinensis in October. 2n = 10 (Blackman & Eastop 1994).

Shivaphis (Sinishivaphis) pteroceltis Jiang, An, Li & Qiao Apterae are red-brown in life, covered in thick powdery or filamentous white wax; BL 1.5-1.8 mm. In large colonies on leaves of Pteroceltis tatarinowii in China (Shandong and Anhui provinces); colonies with powdery wax are attended by ants, but those with wax filaments are unattended. Fundatrices hatch in March, and oviparae and alate males occur in October. Jiang et al. (2014) provided descriptions of all morphs.

Shivaphis (Sinishivaphis) szelegiewiczi Quednau Only alate viviparae are known; colour in life not recorded, presumably covered in wax, BL c. 2.5 mm. Forewings are marked with fuscous as in S. celti, but with a spot below the pterostigma, next to base of media. On Celtis aurantiaca in Korea (Quednau, 1979), and on C. biondii in Hong Kong. An immature ovipara collected in Hong Kong in December is illustrated by Quednau (2003).

Shivaphis (Sinishivaphis) tilisucta Zhang Only alate viviparae are known; bright green covered with thick wax powder, BL c. 1.9mm. Described from Tilia tuan in China (Zhang & Zhong 1990), but the host was possibly misidentified and it has subsequently been collected from Celtis sinensis in China, and from C. japonica and C. jessoensis in Korea. Quednau (2003) illustrated an ovipara collected on C. jessoensis in Korea in early November.

Siculaphis Quednau & Barbagallo

Calaphidinae: Myzocallidini

One species in the Mediterranean region on Quercus, the apterae of which have remarkable variation in form of dorsal hairs.

Siculaphis vittoriensis Quednau & Barbagallo Apterae are pale yellowish green with longitudinal olive green dorsal stripes leaving a pale spinal line; BL 1.0-1.5 mm. Alatae have smoky wings with Cu_1b strongly bordered with fuscous. On Quercus calliprinos and Q. ilex, forming ant-attended colonies mainly along mid-ribs on basal parts of undersides of leaves. In Italy; described from Sicily, also found in Puglia, Tuscany and Sardinia, and also now recorded from Tunisia (Ben Halima-Kamel 2012). A holocycle occurs in Puglia; the ovipara and apterous male were described from Q. ilex by Patti & Barbagallo (1997). Populations in Sicily were suspected to be anholocyclic (Quednau & Barbagallo 1991).

Sinochaitophorus Takahashi

Calaphidinae: Pseudochromaphidini(?)

One species on Ulmus in east Asia perhaps related to Pseudochromaphis, the apterae having extensive dark dorsal sclerotisation. Quednau (2003) reviewed the genus and decided that it comprised one species with vary variable pigmentation.

Sinochaitophorus maoi Takahashi (= mordvilkoi Quednau & Shaposhnikov) Colour of aptera is not recorded, probably blackish; BL c. 1.3 mm. Alatae are blackish-brown, mottled with red, with maculate wings (Holman & Szelegiewicz 1974a). On young shoots and along mid-ribs on the undersides of the leaves of Ulmus japonica in east Siberia (Quednau & Shaposhnikov 1988, as mordvilkoi), and on U. pumila in Mongolia and China. It is attended by ants. Quednau (2003) illustrated oviparae collected in China and east Siberia in September-October.

Sinolachnus Hille Ris Lambers

Lachninae: Tuberolachnini

Two species described from Taiwan and one from Bhutan, distinguished in particular by the very numerous protuberant secondary rhinaria on the antennae of alatae (Hille Ris Lambers 1956d). One of the Taiwanese species is only known from a trapped alata.

Sinolachnus elaeagnensis Chakrabarti & Das Apterae are undescribed, alatae have BL 3.7-4.0 mm. Collected (with immatures) from young stems of an Elaeagnus sp. in Bhutan (Chakrabarti & Das 2014). Attended by black ants. Sexual morphs and life cycle are unknown.

Sinolachnus niitakayamensis (Takahashi) Apterae are yellowish-brown, blackish on dorsal abdomen; BL c. 2.5mm. On branches of Elaeagnus oldhami in Taiwan (Takahashi 1927a). Sexual morphs and life cycle are unknown.

Sinomegoura Takahashi

Aphidinae: Macrosiphini

Eight species with head ventrally spinulose and with weakly clavate, calf-shaped siphunculi and a long cauda, living on various trees and shrubs in east and south-east Asia. Accounts are available for Japan (Miyazaki 1971), China (Tao 1963), north-east India (Raychaudhuri 1980) and Java (Noordam 1986).

Sinomegoura citricola (van der Goot) Plate 24i Apterae are broadly spindle-shaped, shiny dark brown to reddish-brown with bright red eyes, siphunculi pale at bases and dark distally, and a long dark cauda (see Sasaki 2019b for colour photographs); BL 1.4-2.7 mm. On undersides of leaves or young growth of numerous tropical shrubs and some trees, in about 20 families, the most frequent records being from Lauraceae and Rutaceae. Distributed throughout east and south-east Asia (India, Sri Lanka, China, Thailand, Korea, Taiwan, Japan, Malaysia, Indonesia, the Philippines); also in Australia (Queensland), and recently found in southern California (Halbert 2009). No sexual morphs are known, and it is apparently mainly or entirely anholocyclic. Various karyotypes have been reported, and the variation in chromosome number warrants further investigation; 2n = 12 (Kulkarni 1984, Dutta & Gautam 1993), 2n = 16 (Chen & Zhang 1985b) and 2n = 18 (Kar & Khuda-Bukhsh 1986).

Sinomegoura coffeae Noordam Apterae are broadly spindle-shaped, shining yellow with black antennae and tibiae, siphunculi browner towards apices, and cauda yellow or slightly grey; BL 2.4-3.0 mm. On undersides of leaves of Coffea sp. in Java and Timor (Indonesia). Coffea is not indigenous to south-east Asia and there is presumably a native host (see Noordam 1986).

Sinomegoura nepalensis Das & Raychaudhuri Apterae are yellowish, with siphunculi basally pale and distally dark as in citricola, but with a pale cauda; BL 2.8-2.9 mm. On young shoots of Melia sp. in Nepal (B.C. Das & Raychaudhuri 1983). The life cycle is unknown.

Sinomegoura photiniae (Takahashi) Apterae are green, often with a pinkish tinge (Japan, Takahashi 1960c) or ‘faded orange’ (China, Takahashi 1936a), with black siphunculi and green or yellowish cauda; BL 2.0-2.8 mm. On Photinia spp. and Rhaphiolepis spp. (Miyazaki 1971), and also recorded from Phoebe sp. (Tao 1999). In China, Taiwan, Japan and India. Sexual morphs and life cycle are unknown. 2n = 18 (Khuda-Bukhsh & Kar 1990).

Sinomegoura pyri Ghosh & Raychaudhuri Apterae are pale yellowish, with pale siphunculi and cauda; BL c. 2.4 mm. On undersides of young leaves of Pyrus sp. in Assam, India (A.K. Ghosh & Raychaudhuri 1968b). Ardisia sp. is also recorded as a host plant (Raychaudhuri et al. 1980b). Other morphs and life cycle are unknown.

Sinomegoura rhododendri (Takahashi) Apterae are green, with dark-tipped siphunculi and black cauda; BL c.1.4 mm. On Rhododendron oldhami in Taiwan. Records from other hosts in Philippines and India are likely to be referable to other species.

Sinomegoura symplocois (van der Goot) Apterae are pale reddish brown with eyes, antennae greyish white with apices of segments black, legs pale brown with apices of tibiae and tarsi black, siphunculi pale brownish yellow with dark apices, cauda black; BL c.2.9-3.0 mm. On Eurya japonica and Symplocos spp. (cochinchinensis, sessilifolia). In Java (van der Goot 1917, Noordam 1986) and Malaya (BMNH collection, leg. R. Takahashi). The life cycle is unknown. [The record from Acronychia laurifolia in north-west India (Shillong; A.K. Ghosh & Raychaudhuri 1970) seems to be another, apparently undescribed, species, to judge from two alatae (leg. A.K. Ghosh) with the same date, host and locality data in the BMNH collection.]

Sinonipponaphis Tao

Hormaphidinae: Nipponaphidini

Three or four species in east Asia in which the dorsum of the aleyrodiform aptera is densely covered with spine-like processes.

Sinonipponaphis formosana (Takahashi) Apterae are blackish-brown, with a small yellowish-brown patch on ‘margin near caudal apex’ (Takahashi 1927b); BL c. 1.7mm. Alatae are black with brownish abdomen, with secondary rhinaria distributed ANT III 32-36, IV 18-20, V 10-13. Qiao et al. (2018) provided a redescription. Apterae are fixed firmly to branches of Persea (= Machilus) sp. (Takahashi 1929). It has been found in Taiwan, China (Szechuan, Tao 1966) and Thailand. In Taiwan it is also recorded from Phoebe sp. (Tao 1999). The life cycle is unknown. [The aphid described under the name Sinonipponaphis monzeni from Machilus sp. in Meghalaya, India (A.K. Ghosh 1988) may be this or a related species.]

Sinonipponaphis hispida Noordam Apterae dull black or brownish-black with reddish-brown cauda, without wax; BL 0.8-1.0mm. Living scattered on undersides of leaves of Lithocarpus indutus in Java (Noordam 1991). Alatae and life cycle are unknown.

Sinonipponaphis holboelliae (A.K. Ghosh & Raychaudhuri) Apterae are brownish, covered with mealy wax; BL 1.7-2.1 mm. Heavily infesting stems of Holboellia latifolia in West Bengal. Other morphs are unknown.

Sipha Passerini

Chaitophorinae: Siphini

Eleven species of spiny-haired oval grass-feeding aphids with stump-shaped siphunculi, 5-segmented antennae and a sclerotic tergum. The four species of Sipha s.str. have a knobbed cauda; two are nearctic with a smooth tergum and two palaearctic with a spinulose tergum. The eight species of subgenus Rungsia have a broadly rounded cauda and a smooth tergum. Regional accounts are available for UK (Stroyan 1977), north-west Europe (Heie 1982), the Iberian peninsula (Nieto-Nafría & Mier Durante 1998), European Russia (Shaposhnikov 1964), North America (Halbert et al. 2013b) and USA Rocky Mountain region (Palmer 1952). Wieczorek (2010) fully reviewed the genus with distribution maps and illustrations, host lists and keys to all available morphs. Work by T. Liu et al.(2022) on the molecular phylogeny of Chaitophorinae suggests that there may have been an early origin of Siphini by transition to grasses in the early Eocene.

Sipha agropyronensis (Gillette) Apterae are long-bodied, rusty yellow with brown pleural stripes; BL 1.6-2.2 mm. On upper surfaces of leaves of Elymus glaucus (as Agropyron glaucum), attended by ants (Gillette 1911, as Chaitophorus). Also collected on Elymus smithii (as Agropyron occidentale; BMNH collection, leg. M.A. Palmer) and E. hispidus (Wieczorek 2010). USA; Colorado, with records also from California and Montana (see Halbert et al. 2013b), and Gillette mentions collections “throughout the Northwest”. Monoecious holocyclic, with oviparae in October (Palmer 1952).

Sipha (Rungsia) arenarii Mordvilko Apterae are greenish yellow to brown with pale longitudinal stripes; BL 2.0-2.4 mm. The usual host is Leymus arenarius, but it is also recorded from Elymus hispidus, E. Repens, Calamagrostis epigeios and Avena sterilis ssp. ludoviciana. In Europe, and eastward to west Siberia, Kazakhstan and Inner Mongolia. A record from Turkey (on Triticum aestivum; Akyürek et al. 2011) requires confirmation. Monoecious holocyclic, with oviparae and apterous males in October. The sexual morphs were redescribed by Wieczorek (2010).

Sipha (Rungsia) burakowskii Holman & Szelegiewicz Apterae are greyish green with brownish spots, very slightly pruinose; BL 1.8-2.3 mm. In small colonies on upper sides of leaves of Leymus chinensis (= Aneurolepidium pseudoagropyron) in Mongolia, and also recorded from Elymus repens and Leymus ramosus in Kazakhstan (Kadyrbekov 2002e, 2017a). Wieczorek (2010) provided a redescription of the aptera. Alatae and sexual morphs are unknown.

Sipha (Rungsia) elegans del Guercio (= aegilopis Bozhko; synonymy by Wieczorek 2010) Apterae are yellowish brown to brown or olive green with dark segmental markings leaving a paler spinal stripe (see influentialpoints.com/Gallery); BL 1.4-2.1 mm. Alatae have black dorsal abdominal markings. In small colonies on upper sides of leaves of various grasses and cereals (Aegilops, Agropyron, Agrostis, Ammophila, Arrhenatherum, Bromus, Elymus, Festuca, Hordeum, Phleum, Puccinellia, Setaria, Stipa, Triticum) often causing leaves to roll upwards and develop yellow patches. Havličkova et al. (1996) recorded its differential colonisation of Aegilops and wild Triticum species in Czech Republic. In Europe, across Asia to China and east Siberia, and in North America (see Wieczorek 2010 for full details of distribution). Monoecious holocyclic with slender brown-black apterous males, in both Europe and North America. See also Blackman & Eastop (2000). 2n=6.

Sipha flava (Forbes) Yellow Sugarcane Aphid Plate 5d Apterae are straw to bright yellow (or green at low temperatures) with dusky transverse intersegmental markings; BL 1.3-2.0 mm. Alatae have variably-developed dark dorsal abdominal markings. On leaf blades of grasses in numerous genera including Andropogon, Avena, Cenchrus, Cymbopogon, Cynodon, Digitaria, Holcus, Hordeum, Miscanthus, Panicum, Paspalum, Pennisetum, Saccharum, Setaria, Sorghum and Triticum, often forming large colonies, not attended by ants. Bradshaw et al. (2010) found large populations on young stands of the bioenergy crop Miscanthus ×giganteus in USA. Life table parameters of S. flava on this and another bioenergy crop were measured by Pallipparambil et al. (2014). It is also recorded from Cyperaceae (Carex, Cyperus). In USA (including Hawaii), Central and South America, Madeira, the Azores, and recently found in Morocco (Abdelmajid 2008), Spain (Hernández-Castellano & Pérez-Hidalgo 2014) and South Africa (Way et al. 2014). Monoecious holocyclic with apterous males in areas with cold winters, anholocyclic in warmer climates. Genome sequence data is available at: https://www.ncbi.nlm.nih.gov/genome/annotation_euk/Sipha_flava/100/.

See also Blackman & Eastop (2000).

Sipha glyceriae (Kaltenbach) Apterae are dull pale green to mid-green, sometimes reddish, often with a paler spinal stripe; BL 1.5-2.4 mm. On many species of Graminae, especially in wetlands; genera include Agrostis, Aira, Alopecurus, Glyceria, Elymus, Hydrochloa, Oryza, Paspalum, Phalaris, Poa, Puccinellia. Also collected from Ceratophyllaceae (Ceratophyllum), Cyperaceae (Carex, Cyperus, Eleocharis, Schoenus, Scirpus), Juncaceae (Juncus), Typhaceae (Typha) and Alismataceae (Sagittaria). In Europe, Middle East, eastward to west Siberia and Central Asia, and in USA and Canada (see Wieczorek 2010 for full details of distribution). Monoecious holocyclic, with dark apterous males. 2n=12 (Gut 1976) or 10.

Sipha littoralis (Walker) Apterae are green, dark green or bluish green with brownish markings, head and thorax more brownish; BL 1.4-2.2 mm. In leaf sheaths of grasses in coastal situations (e.g. Spartina spp., Puccinellia maritima, Festuca rubra), where they are able to survive tidal submergence. In north-west and northern Europe. Monoecious holocyclic with brown oviparae and apterous males in October (Heie 1982), or as early as August in northern Russia (Wieczorek 2010).

Sipha (Rungsia) maydis Passerini Plate 5e Apterae are shining dark brown to almost black on dorsal surface; BL 1.0-2.1 mm. Alatae have a solid black dorsal abdominal patch. On numerous species of Poaceae, in more than 30 genera. It feeds on upper sides of leaf blades near bases, sometimes on stems or inflorescences, often attended by ants. In Europe, Mediterranean region, Middle East, Central Asia, India, Pakistan, South Africa, and now in Argentina (Corrales et al. 2007) and USA (Sorensen 2007, Halbert et al. 2013b, Puterka et al. 2019) (for full details of distribution see Wieczorek 2010). Monoecious holocyclic on Poaceae with apterous males recorded from Turkey (Tuatay & Remaudière (1965) and Poland (Wieczorek 2010), but probably anholocyclic in most countries. The male genitalia were described and illustrated by Wieczorek (2011). In drier climates outside north-west Europe it can be a pest of cereal crops (Blackman & Eastop 2000). 2n=12.

Sipha (Rungsia) praecocis (Bozhko) Apterae are yellowish, mottled with green; BL c.1.6 mm. In small colonies on upper sides of leaf blades of Carex preacox in Ukraine, and also recorded from Kazakhstan. Alatae and sexual morphs are unknown.

Sipha (Rungsia) taurica (Mamontova) Apterae are lemon yellow; BL 1.8-2.2 mm. On undersides of leaves of Leymus racemosus, turning them yellow. In Ukraine (Crimea), Russia (Volgograd; Kadyrbekov 2014g) and also recorded from Kazakhstan. Alatae and sexual morphs are unknown.

Sipha (Rungsia) uvarovi Mordvilko Apterae are green to dark green; BL c.1.5-1.6 mm. On wheat (Triticum) and various grasses (Agropyron, Festuca). In Ukraine, (Turkey; Şenol et al. 2017), southern Russia, west Siberia, and Kazakhstan. Alatae and sexual morphs are unknown.

Siphonatrophia Swain

Aphidinae: Aphidini

One nearctic species which is a specialized feeder on Cupressaceae (Remaudière & Munoz-Viveros 1991).

Siphonatrophia cupressi (Swain) Apterae are pale green (concolorous with leaves of host), with body strongly domed dorsally and flattened against leaf ventrally (see Fig.1 in Ortego et al. 2018); BL 1.0-1.7 mm. Alatae have the forewing media once- or twice-branched. On Cupressaceae (Juniperus, Cupressus, Widdringtonia) in western USA, Mexico and Central America (Honduras, Costa Rica), also now recorded from Argentina (Ortego et al. 2018), and introduced to southern Europe (France, Rabasse et al. 2005a; Italy, Bella & Squarcini 2009; Spain, Lumbierres et al. 2015), Tunisia (Halima-Kamel & Mdellel 2017) and Japan (Sano & Yoshitomi 2012). A significant amount of parasitization by Lysiphlebus testaceipes has been observed in southern France and Spain (Rabasse et al. 2005a; Lumbierres et al. 2015). Monoecious holocyclic; oviparae and both alate and apterous males occur in October-November (Palmer 1952).

Sitobion Mordvilko

Aphidinae: Macrosiphini

About 66 species resembling Macrosiphum in the reticulation of their siphunculi, but with shorter hairs, less well‑developed antennal tubercles, and often with a more sclerotic dorsum. The genus is hard to define; the karyotype (2n = 16 or 2n == 18) suggests that the strong similarity to Macrosiphum (which have 2n = 10) may be due to morphological convergence. Probably they have a closer relationship with Metopolophium, which also have 2n=16 or 18 and have a similar biology. Sitobion have a broad range of host plant relationships, with species about equally distributed between monocots (particularly Poaceae) and dicots. Several species host-alternate between dicots (Rubus, Rosa, Akebia) and Poaceae, and some of the other dicot‑feeding species may prove to be the primary host forms of species on grasses. Sitobion are distributed throughout the northern hemisphere, and have also undergone an adaptive radiation in Africa south of the Sahara. The four European species are all associated with Poaceae. The 24 African species include 9 grass‑feeders and 13 from dicots. The 30 Asiatic species include only 6 from Poaceae, but 10 from orchids and other monocots and 13 from dicots. Twelve American and one European species previously in Sitobion were placed in Macrosiphum after a cladistic analysis by Jensen (1997). Five grass-feeding species with characters intermediate between Sitobion and Metopolophium are placed in subgenus Metobion; a key to these species was provided by Stekolshchikov (2011). Accounts are available for Europe (Holman 1961c, added to by Stroyan 1969b), UK and Ireland (Blackman 2010), Fennoscandia and Denmark (Heie 1994), east Africa (Eastop 1958), west Africa (Eastop 1961a), Africa south of the Sahara (Remaudière & Autrique 1985), Japan (Miyazaki 1971, in Macrosiphum), China (Tao 1963), India (David 1976), north-east India (Raychaudhuri, 1980, in Macrosiphum) and east Siberia (Pashchenko 1988a). Many of the African species are parasitised by Aphidius camerunensis (see Remaudière & Autrique 1985). Sunnucks et al. (1996) and Hales et al. (2010) discussed the introduced species in Australia.

Sitobion africanum (Hille Ris Lambers) Plate 26d Apterae are variably coloured yellowish green to green, reddish, yellowish, or greenish brown, reddish or brownish forms being usually densely wax‑covered; BL 1.6‑3.1 mm. There is also a very variably developed dorsal abdominal pattern of dark pigmentation; there may be an extensive, broken pattern of sclerotization of the whole dorsal surface or, at the other extreme, the cuticle of the paler green forms may be almost completely pale. Alatae have a pattern of transverse, narrow, often interrupted dark bars on the dorsal abdomen. On Poaceae (Aristida, Bothriochloa, Brachyaria, Bromus, Calamagrostis, Cenchrus, Cynodon, Dichanthium, Digitaria, Eragrostis, Exotheca, Hyparrhenia, Panicum, Pennisetum, Setaria, Sorghum, Sporobolus, Stenotaphrum, Tricholaena, Triticum, Zea) in Africa, Indian Ocean islands, and also recorded from Yemen. A record from Chile was based on a misidentification (E. Fuentes-Contreras, pers. comm.), and a record from Ficus sp. in Turkey (Akyürek et al. 2019 is doubtful and requires additional confirmation. In Africa S. africanum is recorded frequently on various other monocots (Asparagus, Cyperus, Gladiolus, Watsonia) and also on dicots (Aeschynomene, Aster, Clematis, Desmodium, Droogmansia, Euphorbia, Hakea, Hypericum, Lactuca, Lefeburea, Pavetta, Tephrosia, Thesium, Ximenia), including some trees (e.g. Dombeya, Ficus, Phyllanthus). Euphorbiaceae-feeders identified as S. africanum have lateral abdominal tubercles; possibly more than one species is being identified as africanum, and it may eventually transpire that this name is being applied to a complex of species. Müller & Schöll (1958) gave an account of the variation and host plants of S. africanum in South Africa. The life cycle is unclear; it is probably mainly anholocyclic, but Müller & Schöll (1958) described oviparae from Ficus.

Sitobion akebiae (Shinji) Apterae are variably coloured green, yellowish green or orange red, with dorsal abdomen pale or variably pigmented. Heteroecious holocyclic between Akebia and secondary hosts mainly in Poaceae according to Moritsu (1944, as granarium) and Takahashi (1964), but Miyazaki (1971) found fundatrices on Stellaria and Platanus, so overwintering eggs are apparently laid on various plants. Summer host plants include Elymus, Festuca, Oryza, Poa, and Triticum; also Gladiolus, Plantago, Ranunculus and Trifolium. In Japan and Korea. Kanehira et al. (1988) studied population dynamics of aphids identified as S. akebiae in wheat fields in Japan. The more widely distributed S. miscanthi is very difficult to separate from S. akebiae and may perhaps be the same species. For further information see Yano et al. (1983).

Sitobion alopecuri (Takahashi) (= Sitobion salicornii Richards; synonymy by Jensen, 1997) Apterae are yellowish brown to mid apple-green with pale to dusky siphunculi, darker at apices; BL 1.8-3.2 mm. On grasses, particularly Alopecurus, but also in sandy habitats on Ammophila, Elymus, Hordeum and Poa. Asia (Taiwan, north India), Europe (Netherlands, UK, Poland) and North America (California, Hille Ris Lambers 1966a, as ssp. sylvesteri; Oregon, A. Jensen, aphidtrek.org; British Columbia, Richards 1963, as salicornii – vagrants?; Manitoba, BMNH collection, leg. A.G. Robinson). Monoecious holocyclic on Alopecurus, with oviparae and alate males produced in October (Stroyan 1991, and BMNH collection). It is characteristic of the European and North American populations to have marginal abdominal tubercles on all or most of abdominal tergites 2-5, whereas these are not developed in the single oriental specimen seen (from India). S. pseudoalupecuri Chakrabarti, described from an unidentified grass in Himachal Pradesh, India, is very similar but has a shorter, blunter cauda. 2n=18 (sample from British Columbia).

Sitobion anselliae (Hall) Apterae are grass green to yellow green with black siphunculi; BL 1.4-2.2 mm. Alatae have no dark dorsal abdominal markings. On wild and cultivated orchids (Ansellia, Cattleya, Epidendrum, Eulophia). Common on Eulophia streptopetala in Burundi (Remaudière & Autrique 1985). In Africa south of the Sahara (Angola, Burundi, Kenya, Rwanda, South Africa and Zimbabwe). Colonies are ant-attended (Pheidole sp.; van Harten & Ilharco 1976).

Sitobion asirum Aldryhim & Ilharco Apterae are pale green; BL 1.7-2.0 mm. In small colonies on upper sides of leaves of Phragmanthera regularis (a parasite on Acacia) in Saudi Arabia. Closely related to an African species on Loranthaceae, S. loranthi.

Sitobion aulacorthoides David, Narayanan & Rajasingh Apterae are green, with dark brown apices to femora and tibiae, and siphunculi also dark brown on distal half; BL 1.8-2.4 mm. Described from undersides of leaves of an unidentified plant (a “weed”) in NW India (Himachal Pradesh). There are subsequent records of this species from various plants in north-west and north-east India, mainly Lamiaceae (Ajuga, Elsholtzia, Perilla, Plectranthus), but also from species of Commelina, Rubus and Valeriana, which are presumably not normal host plants. Biology is unknown. 2n=18.

Sitobion autriquei Remaudière Apterae are shining brown-black, with black SIPH, and cauda pale except for dusky tip; BL 2.0-2.7 mm. On various Compositae (Conyza, Crassocephalum, Helichrysum), and also collected from species of Plectranthus (Lamiaceae) and Laportea (Urticaceae). In Africa; as yet known only from Burundi (Remaudière & Autrique 1985). The life cycle is unknown.

Sitobion avenae (Fabricius) Grain Aphid Plate 26e Apterae are yellowish green or dirty reddish brown, sometimes rather shiny, with black antennae and siphunculi (see influentialpoints.com/Gallery); BL 1.3‑3.3 mm. On numerous species of Poaceae, including all the cereals and pasture grasses of temperate climates, and many other monocots. Certain dicots may also occasionally be colonised, e.g. Capsella bursa-pastoris and Spergula arvensis. On cereals the aphids prefer to feed on the upper leaves, and on the ears once these have emerged. Throughout Europe, the Mediterranean, the Middle East, Central Asia, India, Nepal, Pakistan, Africa; North, Central and South America (oriental records refer to S. miscanthi). Monoecious holocyclic, with alate males, on many species of Poaceae (Müller 1977a). Wu et al. (2022) described differences between numbers and morphology of secondary rhinaria in different morphs. Anholocyclic overwintering is common in regions with mild winters. Rautapää (1970) studied preferences among a range of host plants. DNA studies in southern England (Sunnucks et al. 1997) and France (Haack et al. 2000) have revealed host specialisation including races preferentially colonising wheat and Dactylis glomerata, the latter form apparently hybridising naturally with S. fragariae. However in England there is molecular evidence from suction trap material that an insecticide-resistant clone of S. avenae may now predominate (Morales-Hojas et al. 2020). Economic implications of limited genetic exchange between populations on different hosts were discussed by Vialatte et al. (2005). The genetics of introduced populations in Chile was studied by Figueroa et al. (2005).

S. avenae, S. miscanthi and S. akebiae form a complex of closely related species. Choe et al. (2006) compared variation in morphology and in an mtDNA sequence of S. avenae from England and North America with the oriental taxa S. miscanthi and S. akebiae, and concluded that the latter two should be treated as synonyms of S. avenae. As a result, in some recent papers cereal-feeding populations of this complex in east Asia have been identified as S. avenae (e.g. Xu et al. 2011; Xin et al. 2014; B. Zhang et al. 2016; D.. Wang et al. 2019). However we agree with Hales et al. (2010) that there is insufficient evidence for such a step, and a more thorough study of oriental Sitobion is needed. It is possible that there is a complex situation in China involving both introduced S. avenae and indigenous S. miscanthi, which could explain the high levels of genetic diversity revealed in recent studies using microsatellite markers in populations identified as S. avenae (Xin et al. 2014, D.Wang et al. 2020), as well as those identified as S. miscanthi (Y. Wang et al. 2016b; Morales-Hojas et al. 2020). This could also explain the genetic differentiation between populations on wheat and barley in China (D. Wang et al. 2019, 2020a,b) The ongoing confusion may also need to be taken into account in studies of secondary symbionts (Alkhedir et al. 2015; Hu et al. 2019). The complete mitochondrial DNA of a sample from China identified as S. avenae has been published (B. Zhang et al. 2016). See also Blackman & Eastop (2000, 2007). 2n=18.

Sitobion bambusicola (L.K. Ghosh) Apterae are yellowish; BL c. 3.7mm. In a colony under young, tender leaves of an unidentified Bambusa sp. in Himachal Pradesh, India (L.K. Ghosh 1986). Other morphs and life cycle are unknown.

Sitobion bamendae (Eastop) Apterae are green with dark dorsal abdominal cross-bands, pinkish green or olive-green to almost black; BL 1.7-2.5 mm. On grasses (Pennisetum, Setaria) in Africa (Burundi, Cameroon).

Sitobion (Metobion) beiquei (Hille Ris Lambers) Apterae are pale green with bright green spinal and marginal longitudinal stripes; BL 3.9-4.2 mm. Originally beaten from mixed vegetation, subsequently collected from Poaceae (Agrostis, Calamagrostis) (BMNH collection, leg. G.W. Simpson and M.E. MacGillivray). In north-eastern USA (Maine) and Canada (Quebec). There are also recent records from western USA (Idaho, Oregon, Utah) including oviparae collected from a lakeside sedge in October (A. Jensen, aphidtrek.org).

Sitobion berchemiae (Takahashi) Apterae are blue-green, BL c. 1.8 mm. On Berchemia lineata and B. floribunda in China. S. berkemiae (Shinji) is possibly a synonym.

Sitobion berkemiae (Shinji) Apterae are deep green with mainly black antennae and legs, black siphunculi and a pale yellow cauda; BL 1.3-1.8 mm (Miyazaki 1980a). On Berchemia racemosa in Japan. The life cycle is unknown.

Sitobion (Metobion) brevirostre (Heikinheimo) Apterae are blackish green with dark antennae and pale siphunculi; BL c. 4.1 mm. The host plant is unknown, but assumed here to be a species of Poaceae. Only known from Finland.

Sitobion breyniae Noordam Apterae are dull green with pale brown head, black siphunculi and brownish yellow cauda; BL 1.8-2.3 mm. On Breynia microphylla in Java (Noordam 2004).

Sitobion burundiense Remaudière Apterae are shining dark green with black siphunculi and rather dark antennae, legs and cauda; BL 2.1-2.9 mm. It differs from other African Sitobion in having 4 hairs (not 3) on first tarsal segments. On Kotschya africana and Cassia sp. in Burundi (Remaudière & Autrique 1985). It apparently only occurs on Leguminosae/Fabaceae. The life cycle is unknown.

Sitobion (Metobion) calvulum (Ossiannilsson) Apterae are dull bluish green, immatures yellowish green; 2.1-2.5 mm. Originally described from Poa arctica and there have been subsequent collections from this plant, but colonies have also been found apparently feeding on Salix polaris and above-ground stems of its root parasite Pedicularis hirsuta. Only known from Spitzbergen. Monoecious holocyclic; the life cycle is abbreviated with oviparae and apterous males in late July-early August (Hodkinson et al. 2004, as Acyrthosiphon). It is parasitised by an endemic aphidiine species (Chaubet et al. 2013).

Sitobion caricis (Glendenning) Only alatae are described (possibly of more than one species), on Carex sp. in northern British Columbia, Canada.

Sitobion cissi (Theobald) Apterae are variable in colour, from pinkish brown to almost black, with some whitish wax intersegmentally; BL 2.0-2.5 mm. The extent of dark dorsal pigmentation varies from total absence in samples from South Africa, to broad dark bars or paired patches on all tergites in a collection at over 2000m altitude in Kenya (BMNH collection, leg. J.H. Martin). Alatae are brown to black with similarly variable dorsal pigmentation. On Cissus and Rhoicissus in Kenya, Zimbabwe and South Africa. Monoecious holocyclic, with oviparae and alate males collected in June in Pretoria (Eastop 1958).

Sitobion colei (Eastop) Apterae are probably blackish in life; BL 1.7-2.3 mm. On several genera of Lamiaceae (Aeollanthus, Ocimum, Plectranthus, Salvia, Solenostemon, Tetradenia) in southern Africa (Remaudière & Autrique 1985).

Sitobion congolense (Doncaster & Hille Ris Lambers) (Fig.57b) Apterae are mid- to dark green; BL 2.2-2.5 mm. On young growth of Compositae (Vernonia spp., Elephantopus scaber), with records also from the guttiferous tree Harungana madagascarensis, which seems able to act as a reserve host for both this aphid and S. nigeriense (Remaudière & Autrique 1985). In Africa (Angola, Burundi, Cameroon, Congo, Ghana, Ivory Coast, Kenya, Nigeria, Zaire). Attended by ants (Duviard 1969), and parasitized by Aphidius camerunensis (Remaudière & Autrique 1985). The life cycle is unknown.

Sitobion cuscutae (Holman) Apterae are broadly spindle-shaped, orange with black appendages; BL 2.5-3.0 mm. On Cuscuta umbellata in Cuba; this appears to be the main host, and its colour is identical to that of the aphid, but smaller colonies have been also found on plants in genera in other families (Cissus, Pavonia and Ruellia; original description).

Sitobion dismilaceti (Zhang) Apterae are yellowish; BL c.2.7 mm. On Smilax china in Yunnan, China. This is possibly a synonym of S. sikkimense (q.v.).

Sitobion eulophiae Remaudière Apterae are black in life (but macerated specimens are pale with black siphunculi and femoral apices and a dark cauda; BL 2.1-2.6 mm. Alatae are without dark dorsal abdominal markings. On the orchid species Eulophia horsfallii (= rosea) in Burundi. (Not attacking E. streptopetala, the host of S. anselliae at the same locations). It was also found on Liparis neglecta (Remaudière & Autrique 1985).

Sitobion fragariae (Walker) Plate 26f Apterae on Rubus and other Rosaceae in spring are yellowish green with small brown intersegmental markings, with siphunculi either entirely dark or pale with darker apices (see influentialpoints.com/Gallery); BL 2.0-3.0 mm. Heteroecious holocyclic, migrating to Poaceae, where apterae are dirty greenish yellow with black siphunculi, and BL 1.6-2.9 mm. Although the principal primary hosts are Rubus, especially R. fruticosus, the sexual phase may also sometimes occur on Fragaria, Rosa and Geum. In Europe, Asia, and introduced to South Africa, North and South America, and Hawaii. Type specimens of Uroleucon qinghaiense described from vagrants on Artemisia sp, in China (Zhang et al. 1999) are S. fragariae or a close relative. An aphid introduced into Australia and New Zealand and studied as Sitobion sp. “near fragariae” (e.g. Sunnucks et al. 1996) has a relatively long cauda and very few microsatellite alleles in common with S. fragariae from southern England, but morphometric studies indicate that these are probably founder effects (Hales et al. 2010). 2n=18.

Sitobion (Metobion) graminearum (Mordvilko) Apterae are light green or pink, with dark antennae ; BL 3.1-4.0 mm. On Poaceae (Arrhenatherum, Calamagrostis, Deschampsia, Festuca, Phalaris, Poa) in Europe (Sweden, Finland, north-west Russia, west Siberia, Czech Republic, and possibly Ukraine). Stekolshchikov (2011) redescribed apterae, alatae and sexual morphs and designated a lectotype. Monoecious holocyclic, with oviparae and alate males in September.

Sitobion graminis Takahashi Apterae are dark reddish brown covered with grey wax dust, often of a bluish appearance (red in alcohol), with black antennae, legs, siphunculi and cauda; BL 1.8-2.9 mm. On many genera and species of Poaceae, usually feeding on the flower stems. Also recorded from Cyperus and Juncus. In Africa, Réunion, India, Malaya, Java, New Guinea and the Philippines. Apparently anholocyclic. 2n=18.

Sitobion gravelii (van der Goot) Apterae are pale yellow or pale brownish, with distal segments of antennae dark brown, siphunculi dark brown and cauda pale; BL 3.6-4 mm. Described from an unidentified host, but there are later records from Artemisia and unidentified Compositae (Raychaudhuri, 1980, as Macrosiphum spinotibium). However these seem unlikely hosts, and it is more probably a grass feeder. In northern India. Oviparae were collected from an unknown host in late November (A.K. Ghosh et al. 1971d, as M. spinotibium). 2n=12 (Khuda-Bukhsh & Basu 1987, as M. spinotibium on Artemisia vulgaris).

Sitobion halli (Eastop) Apterae are uniformly pale green, with black-tipped siphunculi and pale cauda; BL 1.8-2.9 mm. Immatures are somewhat wax-powdered (Eastop 1959). On young growth of trees in various families (Apocynaceae, Rubiaceae, Moraceae, Euphorbiaceae, Melianthaceae, Myrtaceae) in central and southern Africa (Burundi, Tanzania, Zimbabwe, South Africa; Remaudière & Autrique 1985). The life cycle is unknown.

Sitobion hillerislambersi van Harten Apterae are bright grass-green with a black oval dorsal abdominal patch and black siphunculi; BL 1.7-2.2 mm. On undersides of leaves of Cyperus esculentus in Angola.

Sitobion himalayensis L.K. Ghosh Apterae are greenish with dark siphunculi; BL 3.7-4.3 mm. On inflorescences of Oryzopsis lateralis in Himachal Pradesh, India.

Sitobion hirsutirostris (Eastop) Apterae are pale green with a slightly darker spinal stripe, siphunculi black and cauda pale; BL 2.5-3.1 mm. On Anisopappus africanus, and less commonly on Helichrysum setosum (Remaudière & Autrique 1985). In Africa (Burundi, Cameroun).

Sitobion ibarae (Matsumura) Apterae yellowish green or yellowish brown with pale brownish head, mainly dark antennae and legs, long blackish siphunculi and a long pale cauda; BL 2.1-3.1 mm. On wild and cultivated Rosa spp. in east and south-east Asia, and introduced to Hawaii (BMNH collection, leg. E. Hardy). The life cycle is unknown, perhaps anholocyclic at least in warmer countries. Dwarf, brownish apterae may occur in Japan in midsummer (Takahashi 1964).

Sitobion indicum A.N. Basu (Fig.25c) Apterae are probably yellowish green, with an irregular dark dorsal abdominal patch, black siphunculi and pale brown cauda; BL c. 2.0-2.1 mm. On orchids, mostly Cymbidium spp., in hilly regions of north India (West Bengal, Himachal Pradesh, Uttar Pradesh. Possibly it is entirely anholocyclic. A record from Rosa indica (Kurl 1980) is likely to be based on a misidentification. 2n=18.

Sitobion javanicum Noordam Apterae are shiny brownish red, with brown to blackish siphunculi and brown cauda; BL 2.0-2.3 mm. On Themeda arguens, living between spikelets, in Java (Noordam 2004).

Sitobion kamtshaticum (Mordvilko) Apterae are green with black antennae and siphunculi; BL 1.5-2.9 mm. Alatae have rhinaria on ANT IV as well as III. On young growth of Rosa multiflora and Rubus sp. in spring. Heteroecious holocyclic, migrating to grasses (Digitaria adscendens and Microstegium vimineum; Miyazaki 1971, as rubiphila). In Japan, Korea and east Siberia.

Sitobion krahi (Eastop) Apterae are green, with siphunculi pale only on basal third and a pale cauda; BL 2.5-3.0mm. Alatae have siphunculi wholly black and dark marginal abdominal sclerites, small antesiphuncular sclerites and large postsiphuncular sclerites. Collected on growing points of saplings of a forest tree, probably Rauvolfia vomitoria but possibly Tabernaemontana chippii, in Ghana (Eastop 1959). Specimens trapped or on unidentified hosts have been collected in Nigeria, Sierra Leone and Tanzania (BMNH collection), and Remaudière & Autrique (1985) recorded krahi from Mangifera indica in Burundi. The life cycle is unknown.

Sitobion lambersi David Apterae are pale green with brown siphunculi and a pale cauda; BL c.1.8 mm. On stems and in flower heads of tropical grasses (Bothriochloa, Chloris, Cynodon, Dicanthium, Digitaria, Ischaemum, Panicum, Paspalum, Themeda), and occasionally on Cyperaceae. Widely distributed in the Old World tropics; Africa, southern India, Sri Lanka, Java, and also recorded from Fiji. A record from Turkey (Görür et al. 2017) requires further confirmation. Two alatae have also been trapped in Brazil (C.L. Costa, pers. comm.).The name is possibly being used for a complex of species in the Old World tropics, which share the distinctive character of a cauda with only 2 long hairs.

Sitobion leelamaniae (David) Banded Cereal Aphid (= chanikiwiti Eastop 1959, = howlandae Eastop 1959; synonymy by Eastop & Blackman 2005) Apterae are pale yellowish green to green or bright yellow, antennae ringed with brown, legs pale, siphunculi pale brown, darker towards apices (see aphids of Karnataka website); BL 1.6-2.3 mm. On undersides of leaves of many species of grasses and cereals (Brachiaria, Cynodon, Eleusine, Pennisetum, Sorghum, Zea). Described from south India, and also in Sri Lanka, Réunion (as S. chanikiwiti), and widely distributed in southern Africa, where most records are as S. chanikiwiti and S. howlandae. Possibly heteroecious holocyclic with Hagenia abyssinica as primary host in Burundi (Remaudière & Autrique 1985, as S. chanikiwiti).

Sitobion leonidasi Remaudière Apterae are variably pigmented but usually with a shiny black abdomen and both siphunculi and cauda black; BL 1.8-2.5 mm. Alatae have dark wing veins and variably developed dark abdominal markings. On Bersama ugandensis, living particularly in leaf galls formed by a psyllid. Only known from Burundi (Remaudière & Autrique 1985).

Sitobion loranthi van Harten Apterae are green with siphunculi and apices of antennal segments black; BL 1.7-2.7 mm. On upper and undersides of leaves of Loranthus sp. in Angola. There are similar aphids in the BMNH collection from Loranthus dregei in South Africa, but with a longer and more pointed R IV+V.

Sitobion luteum (Buckton) Yellow Orchid Aphid (Fig.25b) Apterae are bright greenish yellow to pale yellowish green, with an oval black dorsal abdominal patch, antennae black except for base of ANT III, black siphunculi and a pale cauda; BL 1.3-2.1 mm. Alatae have only dark intersegmental markings. On many species of orchids (Dendrobium, Epidendrum, etc.) with an almost world-wide distribution, in temperate regions usually under glass. In Europe, Madagascar, Mauritius, India, Singapore, Java, New Guinea, Australia, Fiji, Tahiti, USA (New Mexico, New York), Central and South America. Apparently entirely anholocyclic; no sexual morphs have been recorded. 2n = 12.

Sitobion martorelli (Smith) Apterae are bright yellow-green with siphunculi dark except at bases; BL 2.7-2.9 mm. On flowers, young leaves and pods of Theobroma cacao in the Caribbean (Cuba, Dominican Republic, Jamaica). Attended by the ant Solenopsis geminata (original description).

Sitobion matatum (Eastop) Apterae are shining black dorsally; BL 2.0-2.5 mm. On Tricholaena rosea (=Melinis repens) in Kenya.

Sitobion mesosphaeri (Tissot) Apterae are yellowish green to bottle green, with dark antennae, legs and siphunculi; BL 1.9-3.2 mm. On Hyptis and Ocimum spp. in southern USA (Florida), Mexico, Panama (Quirós et al. 2009), Puerto Rico, Cuba, Venezuela and Argentina (the latter as M. hyptidis Blanchard 1944), on Plectranthus sp. in Jamaica (Tannice Hall, pers. comm), and on Hyptis spp. and Pogostemon cablin in Costa Rica (Villalobos Muller et al. 2010). Large specimens that are apparently this species have also been collected on Quercus in Colombia, Mexico (BMNH collection, leg. R. Peña) and Costa Rica (BMNH collection, leg. J.H. Martin). The life cycle is unknown. It has been treated as a synonym of Macrosiphum salviae, which possibly belongs in Sitobion; differences between apterae of the two species are given by Remaudière & Remaudière (1997), and in the key to aphids on Ocimum.

Sitobion microspinulosum David, Rajasingh & Narayanan Apterae are olive green, BL c.2.8-3.0 mm. Described from Arthraxon lancifolius in West Bengal, feeding on stalk and inflorescence. Only known from types (holotype aptera and paratype immature alata), which are in BMNH collection.

Sitobion milii Remaudière Apterae are grey with black segmental cross-bands; BL 1.7-2.5 mm. On Euphorbia milii in Burundi. Closely related to S. phyllanthi; see Remaudière & Autrique (1985) for a comparison of the two species.

Sitobion miscanthi (Takahashi) Apterae are variably coloured green, reddish brown to dark brown with shiny black siphunculi, pale cauda, and dorsal cuticle variably tanned from pale with only very small intersegmental sclerites to very dark (in the latter case the pigmented area may be entire or segmentally divided); BL 1.7‑3.0 mm. On many genera and species of Poaceae. Also on Cyperus, and sometimes found in large numbers on dicots, particularly semi‑aquatic species, e.g. Polygonum hydropiper (Eastop, 1966). In Indian subcontinent, east and south-east Asia, Australia, New Zealand, and Pacific islands, including Hawaii. Records from Turkey (Görür et al. 2011b, 2012) require additional confirmation. Anholocyclic almost everywhere, but an ovipara (on Polygonum chinense) and males collected in India may be this species (see David 1976), and oviparae and males have been reared from the introduced populations in Australia (Hales et al. 2010). S. akebiae in Japan and Korea is closely related and on secondary hosts not practically distinguishable from S. miscanthi; it may be a synonym, in which case one would expect the holocycle to involve overwintering as eggs on various dicots, as in Japan. The taxonomy of the miscanthi/akebiae group requires clarification by a combination of molecular and biological studies (see also under S. avenae). Sunnucks et al. (1996) studied DNA and karyotype variation in the introduced populations in Australia, and concluded that probably two successful colonizations had occurred. Genetic evidence suggests that hybridisation may have occurred between introduced populations of S. miscanthi and Sitobion ‘sp. near fragariae’ (see above) in New Zealand (Hales et al., 1998), although the sexual morphs of neither of these species have so far been found in the field there. Recent studies on S. miscanthi populations in China have revealed two strains of Wolbachia (Z.Wang et al. 2009) and a new Rickettsia-like secondary symbiont (T. Li et al. 2011, 2016).

Choe et al. (2006) compared variation in morphology and in an mtDNA sequence of S. avenae from England and North America with S. miscanthi and S. akebiae, and concluded that the latter two should both be treated as synonyms of S. avenae. This has led to confusion in the literature, with some molecular workers identifying cereal aphids as S. miscanthi (Y.Wang et al. 2016b, X.Jiang et al. 2019, Morales-Hojas et al. 2020), and others as S. avenae (q.v.). All three groups reported high levels of genetic diversity in and especially between Chinese populations, which suggests that both species may be present in China. X.Jiang et al. (2019) have published a chromosome-level draft genome of a Chinese cereal aphid that they identified as S. miscanthi. A more thorough study of oriental Sitobion is clearly needed to resolve the confusion that exists at present in the literature. See also Blackman & Eastop (2000, 2017). 2n = 18 in India (Kurl & Chauhan 1988), but varies from 17 to 21 in Australian populations.

Sitobion mucatha (Eastop) Apterae are of two colour forms, both appearing translucent and delicate, one very pale yellow with dark siphunculi and the other pale yellow-green with dusky siphunculi; BL 2.0-2.5 mm. Living singly or in small colonies on undersides of leaves of Vernonia lasiopus in Kenya.

Sitobion neusi (Eastop) Apterae are shining black; BL 1.7-2.3 mm. On various tropical grasses (Digitaria, Eragrostis, Panicum, Setaria, Tricholaena), and also found on Canna and Ensete (BMNH collection). In Kenya, Ethiopia, Burundi (at 2000m) and Madagascar.

Sitobion nigeriense (Eastop) (Fig.57c) Apterae are green with dark siphunculi; BL 1.5-2.2 mm. On young growth of Vernonia spp. (BMNH collection), and also recorded from Guttiferae (Harungana madagascarensis and Psorospermum febrifugum), which possibly act as reserve hosts (cf. S. congolense), although Remaudière & Autrique (1985) recorded it twice colonizing H. madagascariensis in Burundi. Parasitism by a ?Trioxys sp. was observed. In Africa (Burundi, Congo, Nigeria, Kenya, Ethiopia). The life cycle is unknown.

Sitobion nigrinectarium (Theobald) Apterae are green with black siphunculi and a long pale cauda; BL 1.8-2.9 mm. On various Leguminosae/Fabaceae (Cajanus cajan, Glycine javanica (= Neonotonia wightii), Crotalaria, Dolichos, Pisum, Rhynchosia) in Africa, and also on Indian Ocean islands (Madagascar, Mauritius, Réunion, Anjouan). See also Blackman & Eastop (2000). 2n=18.

Sitobion niwanistum (Hottes) Apterae are pale green, dusted with white wax powder (see aphidtrek.org); BL c. 2.8-3 mm. On undersides of leaves of Mertensia paniculata in western USA, and there is also a record from Ohio (Smith & Parron 1978). Monoecious holocyclic, with oviparae and apterous males in late August-September (Palmer 1952, and BMNH collection, leg. D. Hille Ris Lambers).

Sitobion ochnearum (Eastop) Apterae are salmon-pink (but this colour was possibly due to fungal infection); BL 1.7-2.3 mm (Eastop 1959). On Ochna spp. in eastern and southern Africa (Angola, Kenya, South Africa, Zambia, Zimbabwe). The life cycle is unknown. 2n=18.

Sitobion orchidacearum (Franssen & Tiggelovend) Apterae are shiny black with dark red around the siphunculi; BL 1.4-2.7 mm. On Orchidaceae (Coelogyne, Dendrobium) in Java (Noordam 2004).

Sitobion paludum Müller Apterae are shining green or brownish, with black head and antennae, legs with femora black except at bases, black siphunculi and pale cauda; BL 1.7-3.0 mm. On Vaccinium uliginosum in northern Europe. Heteroecious holocyclic, migrating to Rhynchospora alba (Cyperaceae), where apterae are darker than on the primary host. Müller (original description) gave a full account of its biology.

Sitobion papillatum Remaudière Apterae are matt dark brown, with black siphunculi sometimes paler at base; BL 1.7-2.5 mm. In inflorescences of Pennisetum purpureum in Burundi (Remaudière & Autrique 1985). A shorter-haired form on a bamboo, Arundinaria (= Yushania) alpina, was distinguished as a subspecies, S. papillatum ssp. subnudum Remaudière (although if the differences remain consistent after further collections this form should probably be given separate species status). The life cycle is unknown.

Sitobion pauliani Remaudière Apterae are pale green or pink, usually with a shiny dark olive green or brown dorsal patch; BL 1.5-2.1 mm. In inflorescences of grasses, especially Pennisetum, Setaria and Eleusine. In Africa south of the Sahara, Madagascar, Réunion (BMNH collection, leg. G. Remaudière), Andaman & Niccobar Isles, India (Orissa), Sri Lanka, Indonesia, West Malaysia (BMNH collection, leg. J.H. Martin), Philippines (BMNH collection, leg. V.J. Calilung), Central America (Venezuela, Honduras, Panama, Cuba, Jamaica, Costa Rica; Villalobos Muller et al. 2010 and BMNH collection, leg. J.H. Martin) and South America (Brazil, S.P.; S. Matsuoka, pers. comm.).

Sitobion phyllanthi (Takahashi) Apterae are dirty olive green with dark brown head and dorsal markings (Müller 1959; as S. adgnatum); BL 1.5-2.1mm. Prepared specimens of apterae have variably developed narrow dorsal abdominal cross-bands between dark marginal sclerites. Described from Phyllanthus sp. in Mauritius (Takahashi 1937c). An alata probably of this species was trapped in Sri Lanka (BMNH collection). Remaudière (1988) proposed that S. adgnatum Müller, widely distributed on various Euphorbiaceae (Euphorbia, Breynia, Phyllanthus) in Africa (Millar 1994), is this species, and the BMNH collection contains specimens collected by A. van Harten on Phyllanthus maderaspatensis in Angola. It is also reported to occur in Hawaii. Müller (1959) gave a account of this species (as S. adgnatum). The life cycle is unknown.

Sitobion plectranthi M.R. Ghosh, A.K. Ghosh & Raychaudhuri Apterae are pale-coloured; BL 1.7-2.1 mm. On stems and flowerstalks of Plectranthus coetsa, and also recorded from Perilla sp. In north-east India.

Sitobion pseudoluteum A.K. Ghosh Colour of apterae in life is unknown; BL 1.7-2.3 mm. On Masdevallia sp. (? – recorded as Madenvallia) and Cymbidium spp. in West Bengal and Sikkim, India. The consistency of the unusual first tarsal chaetotaxy (4,4,4) needs to be confirmed with additional material. It appears closely related to S. orchidacearum in Indonesia, and could be a synonym. The specimens with a dark cauda and long hairs referred to by David (1976) under the name S. indicum may be this species. 2n=18 (Kar et al., 1990).

Sitobion raoi (Kulkarni) Apterae are pale brown (in life?); BL 2.1-2.5 mm. On Iseilema anthephoroides in Maharashtra, India.

Sitobion rosaeiformis (Das) Apterae are bright yellowish green, green or pinkish brown with antennae and legs mainly pale, and siphunculi entirely or mainly dark with pale bases (see aphids of Karnataka website); BL 2.1-3.3 mm. On both cultivated and wild Rosa in India, Pakistan, Bhutan (Chakrabarti & Das 2014) and Vietnam (Szelegiewicz 1968). Also recorded from Rubus lasiocarpus and Spiraea corymbosa. Probably heteroecious holocyclic with migration to grasses, as it has been collected on Digitaria adscendens (= ciliaris), although host alternation has not been confirmed experimentally by host transfers, and populations can persist on roses into the cold season. Oviparae and alate males were found on Rosa in November-March (David 1976). 2n=18.

Sitobion rosivorum Zhang Colour of apterae in life is unrecorded; BL c. 4.2 mm. On Rosa chinensis and Rosa sp. in China. 2n=18 (Chen & Zhang 1985a).

Sitobion schoelli (Müller) Apterae are green, with head pale brown, antennae and legs with dark apices to segments, siphunculi black and cauda dusky; BL 1.4-2.2 mm. On Kniphofia sp. in South Africa, and Angola (BMNH collection, leg. A. van Harten).

Sitobion (Metobion) scoticum (Stroyan) Apterae are dull apple-green with dark antennae and pale brownish siphunculi; BL 2.3-3.0 mm. On Poa trivialis, living on uppersides of leaves and hidden in inflorescences, in marshy habitat. For many years only known from Scotland, but now recorded from P. pratensis in north-west Russia (Nenets region; Stekolshchikov 2017). Monoecious holocyclic, with alate males.

Sitobion sikkimense (A.K. Ghosh & Raychaudhuri) Apterae are undescribed, probably pale green with black siphunculi, and BL 2.5-3 mm. On undersides of leaves of Smilax sp. in northern India, and subsequently recorded from S. ferox (David 1976, as M. smilacifoliae ssp. ferocis). Monoecious holocyclic with oviparae and alate males in late October (Raychaudhuri et al. 1980). S. dismilaceti, described from Smilax in China, may be a synonym.

Sitobion smilacicola (Takahashi) Apterae are pale yellow with black siphunculi and yellow cauda; BL c. 3 mm. Alatae have 6-8 secondary rhinaria on ANT III. On young leaves of Smilax stenopetala in Taiwan, and recorded also from S. china in Korea (Paik 1965). Both apterae and alatae are described as having femora and tibiae with numerous small circular sensoria or pores. Records from China and India may apply to other species.

Sitobion smilacifoliae (Takahashi) Apterae are bright green to orange-red with very variably developed shining brownish black dorsal sclerotisation, femora with distal half black, siphunculi black and cauda yellow; BL 2-3 mm. Alatae have 5-8 secondary rhinaria on ANT III. On undersides of leaves of Smilax spp. in east Asia (Taiwan, China, Korea, Japan). Specimens from Smilax spp. in Australia, and possibly those previously recorded from Hong Kong and Thailand, seem to be a different species, the identity of which needs further study (Hales et al. 2010).

Sitobion takahashii (Eastop) The colour of apterae in life is not recorded in the original description (Takahashi 1938a, as Macrosiphum phyllanthi). A photomicrograph and line illustrations of an aphid identified as this species are available on the aphids of Karnataka website; BL 1.7-2.3 mm. On Phyllanthus spp. in China, Thailand and India, and perhaps also on Mimosa pudica in India (M.R. Ghosh et al. 1977a). [Specimens from Phyllanthus (maderaspatensis, urinaria) from southern India, Java and Singapore in the BMNH collection have a longer antennal PT and shorter R IV+V than takahashii s. str., and seem to be the form referred to as a subspecies by David (1958); possibly the name mimosae Ghosh, Basu and Raychaudhuri is applicable to this form, but further work is necessary to confirm that there are indeed two very similar species, both living on Mimosa and Phyllanthus.] 2n = 18 (Khuda-Bukhsh & Kar 1989).

Sitobion thalictri Remaudière Apterae are dark green with black antennae and siphunculi, and dark head, legs and cauda; BL 2.0-3.1 mm. On Thalictrum rhynchocarpum in Burundi.

Sitobion triumfettae Remaudière Apterae are very pale yellowish green with a darker green spinal stripe and sometimes dusky dorsal cross-bands; BL 1.7-2.3 mm. On Triumfetta rhomboidea in Burundi.

Sitobion wikstroemiae (Mamet) Colour of apterae in life is unrecorded; BL 1.8-2.4 mm. On Wikstroemia viridiflora in Mauritius (original description) and Sri Lanka (BMNH collection, leg J. Edirisinghe). Specimens collected in Kenya on Gnidia sp. (BMNH collection, leg. J.H. Martin) and trapped in Zimbabwe (BMNH collection, leg. C.E. Taylor) also seem to be this species. 2n=16.

Sitobion yakini (Eastop) Apterae are green, with distal parts of antennae dusky to black, and siphunculi dark distally, sometimes paler at base; BL 1.5-2.2 mm,. On various tropical grasses (Andropogon, Eleusine, Eragrostis, Pennisetum, Poa, Tricholaena, Triticum) in Africa (Burundi, Kenya, South Africa).

Sitobion yasumatsui (Moritsu) Apterae are shiny yellowish brown with a variably-developed dark brown-black dorsal patch (see Moritsu 1983); BL c.2.1 mm. Alatae have more than 40 rhinaria on ANT III. On Spodiopogon (= Eccoilopus) cotulifer in Japan, and Andropogon ascinodis in Thailand (BMNH collection, leg. H. Banziger).

Slavum Mordvilko

Eriosomatinae: Fordini

Four species closely related to Aploneura, producing sac-like or coral-like galls on Pistacia spp. in south-west and central Asia. Three species produce emigrant alatae and are apparently host-alternating, with a two-year life cycle as in most other host-alternating Fordini, but their secondary host generations are still unknown. The fourth lives exclusively on Pistacia. Davatchi (1958) revised the genus.

Slavum esfandiarii Davatchi and Remaudiere Galls on twigs of Pistacia mutica in Iran are coral-like, red when mature, with branches slightly larger than those of S. wertheimae and S. mordvilkoi (Davatchi & Remaudiere 1957). Galls were also recently found on Pistacia sp. in Turkey (Şenol et al. 2014). Heteroecious holocyclic; alatae (BL 2.0-2.2 mm) emerge in November and fly to an unknown secondary host.

Slavum lentiscoides Mordvilko Galls (fig. 126N) are bag-like outgrowths of the leaves of Pistacia spp., similar in position to those of Geoica, but with pointed apices. In Crimea, Iran, Kazakhstan, Tadzhikistan, Turkmenistan and Pakistan . Heteroecious; alatae (BL 1.6-2.2 mm, fig. 81B) emerge in October and are reported to migrate to roots of various grasses (Kadyrbekov 2002d, 2014d), but the secondary host generations are undescribed. It is possible that S. lentiscoides is the primary host form of Aploneura ampelina, which lives on roots of vines in southern Europe, Africa and south-west and central Asia, and is also reported from grasses, but experimental work is needed to confirm this.

Slavum mordvilkoi Kreutzberg Galls on twigs of Pistacia vera are coral-like, similar to those of S. wertheimae. Recorded only from wild P. vera, in Turkmenistan, Afghanistan and eastern Iran (Kreutzberg, 1953). Apparently heteroecious, but only the primary host part of the life cycle is known.

Slavum wertheimae Hille Ris Lambers Galls are branching, coral- or cauliflower-like (fig. 126O), reddish when mature, 5-15 cm in diameter, on twigs and branches of Pistacia atlantica and P. mutica in Iran, Israel, Jordan and Syria. Monoecious holocyclic, presumably with a one-year cycle; alatae (BL 1.7-2.2 mm) emerging from galls in September are sexuparae and presumably give birth to sexuales on the trunk, although this has not been recorded. Wool & Manheim (1983) studied morphological variation, Aloni et al. (1989) studied the differentiation of plant tissue in the galls, Álvarez et al. (2016) studied their ultrastructure in comparison with other Fordini, and Martinez (2008) studied the impact of the galls on subsequent growth of the host trees. Avrani et al. (2012) found genetic differentiation between northern xeric and southern mesic Israeli populations of S. wertheimae indicative of cryptic allopatric speciation..

Smiela Mordvilko

Aphidinae: Macrosiphini

Four palaearctic crucifer-feeding species related to Brevicoryne, with broad dark volcano-shaped siphunculi, a short cauda, and apterae usually with rhinaria on ANT III.

Smiela fusca Mordvilko Plate 12g Apterae are grey-brown, wax-powdered; BL 1.4-1.8 mm. On Berteroa spp., deforming leaves and flower buds (Müller 1975a), and now also recorded from Armoracia rusticana (Basky 2016). In eastern Europe (Germany, Poland, Hungary, Bulgaria, Serbia, Czech Republic, Ukraine). Oviparae occur in October (BMNH collection, leg J. Holman).

Smiela mongolica Holman & Szelegiewicz Apterae are green, densely covered in wax powder, with mainly dark appendages; BL 0.9-1.2 mm. On Alyssum (= Ptilotrichum) canescens, colonising terminal parts of shoots, in Mongolia. Other morphs are undescribed.

Smiela schneideri (Hille Ris Lambers) Apterae are blackish, probably wax-covered in life; BL 1.2-1.5 mm. Alatae have dark transverse bars on the dorsal abdomen. On Isatis glauca, colonising the stem apices and inflorescences, in Syria. A population found on Alyssum tortuosum in Crimea, with fewer secondary rhinaria on the antennae (of apterae) and a shorter antennal terminal process, was described as a subspecies, S. schneideri ssp. alyssii Mamontova-Solukha. Sexual morphs and life cycles are unknown.

Smiela syreniae Bozhko Apterae are greenish brown-black, covered with dense white wax powder; BL c.1.6-1.7 mm. On Erysimum spp. and Syrenia spp. In south-west Russia, Ukraine, Slovakia and Central Asia.

Sminthuraphis Quednau

Calaphidinae: Saltusaphidini

One species related to Iziphya but with a very different pattern of dorsal sclerotisation, all tergites bearing numerous wax glands.

Sminthuraphis ulrichi Quednau (Fig.18j) Apterae are plump-bodied, covered in bluish white wax: BL c.1.5-1.6 mm. Living on ground-level stems of Carex ligerica in Germany, and also recorded from Carex spp. in France, Poland, Hungary and Czech Republic (BMNH collection), and east to Kazakhstan (Kadyrbekov 2011b). Alatae have similar dorsal sclerotisation and wax gland distribution to apterae, bordered wing veins with dark spots at the ends, and secondary rhinaria distributed III 9-11, IV 1-3 (BMNH collection). Living all year on Carex, with sexuales collected in October in France (Leclant 1966).

Smynthurodes Westwood

Eriosomatinae: Fordini

One species characterized by the elongate second antennal segment and the thick sclerotic rims of the primary sensoria on the last two segments.

Smynthurodes betae Westwood Galls on Pistacia spp. (atlantica, mutica and, rarely, vera) are yellow-green or red, spindle-shaped, about 20mm long, formed by rolling of the edge of the leaflet near its base (fig. 126P). These are secondary galls, produced by the progeny of the fundatrix, which lives in a small red mid-rib gall (Burstein & Wool 1991). The ultrastructure of the galls was studied by Álvarez et al. (2016). Host-alternating, with a two-year cycle; alatae (BL 1.3-1.6 mm, fig. 81O) emerge in September-November and migrate to roots of numerous, mostly dicotyledonous, plants. Secondary hosts are particularly Compositae/Asteraceae (Artemisia, Arctium), Leguminosae/Fabaceae (Phaseolus, Vicia, Trifolium), and Solanaceae (Solanum tuberosum, S. nigrum, Lycopersicon esculentum); also sometimes on Beta, Brassica, Capsella, Gossypium, Heliotropum, Rumex, etc. Only rarely is it found on monocots (Poaceae, Cyperaceae). Apterae on roots of secondary hosts are dirty yellowish white, wax‑dusted, with light brown head, prothorax, ANT and legs; BL 1.6‑2.7 mm (Plate 2f). The holocycle is recorded throughout the range of the primary hosts; Algeria, Morocco, Israel, Syria, Iran, southern Crimea, Transcaucasus (Mordvilko, 1935, as Trifidaphis phaseoli; Davatchi, 1958) and Pakistan (Naumann-Etienne & Remaudière 1995). Anholocyclic populations occur commonly on secondary hosts in other parts of the world (see also Blackman & Eastop 2000, p.347). Wool & Burstein (1991a, b) studied phenology and ecology of gall generations and interactions with a parasitoid in Israel. Kurzfeld-Zexer & Inbar (2021) showed that the galls were protected by defoliating caterpillars.2n=8.

Sorbaphis Shaposhnikov

Aphidinae: Macrosiphini

One species perhaps related to Sappaphis, but with 4 instead of 3 hairs on 1st segments of fore and mid-tarsi.

Sorbaphis chaetosiphon Shaposhnikov Plate 10g Spring colonies curl leaves of Sorbus spp. downward and turn them reddish. Host-alternating; emigrant alatae (BL 2.1-3.0 mm, with shining black abdomen due to an extensive sclerotic patch), migrate in late May-June to Ligularia (Stekolshchikov & Shaposhnikov 1998), where they live in ant shelters on stems or at bases of leaves. Apterae on secondary hosts are grey-brown to blackish, with immature stages dirty yellow; BL 1.7-2.6 mm. Stekolshchikov & Shaposhnikov (1998) provided a redescription including fundatrix, emigant alata and apterous exule. In Japan, Korea, China, west and east Siberia and the southern Urals. 2n = 38.

Spatulophorus Müller

Aphidinae: Macrosiphini

Two species in Europe related to North American Landisaphis but with 2 hairs (no sense peg) on HT I, and without the development of the spinal sclerites on abdominal tergites 6-8 into rugose conical processes.

Spatulophorus alyssi Holman Apterae are greyish green with slight pruinosity, especially ventrally; BL 1.1-1.3 mm. Found in September feeding (with sexual morphs) on the upper sides of the youngest terminal leaves of young shoots of Alyssum saxatile in central Bohemia, Czech Republic. The inflorescences are possibly colonised earlier in the season. Monoecious holocyclic, with apterousmales. Alatae are unknown.

Spatulophorus incanae Müller Apterae are pale green to yellowish green or grey-green, BL 1.5-1.9 mm. Living inconspicuously on flower stalks and in inflorescences of Berteroa incana, and after flowering on undersides of leaves. Also recorded from Capsella bursa-pastoris. In eastern Europe (Germany, Hungary, Ukraine). Monoecious holocyclic on Berteroa, with apterous males. Müller (original description) gave a very full account of this species.

Staegeriella Hille Ris Lambers

Aphidinae: Macrosiphini

One or more palaearctic species on Galium and related Rubiaceae, differing from Hydaphias in the shape of the siphunculi. A species in Tasmania, two oviparae of which were collected on Coprosma (BMNH collection, leg. M Williams) may also belong to this genus. The unnamed species from Sorbus aucuparia described as a Staegeriella by Bozhko (1976, p.30) probably does not belong in this genus, and could perhaps be the primary host form of Pseudacaudella rubida.

Staegeriella necopinata (Börner) Apterae are plump-bodied, dark greyish green to lead-coloured, powdered ventrally with grey wax (see influential points/Gallery); BL 1.3-2.2 mm. On Galium spp., and probably also Asperula spp., colonising the stems and flowerheads, causing stunting and twisting of new growth. Throughout Europe, and also recorded from western Siberia, Iran (Momeni Shahraki et al. 2019), Kazakhstan (Kadyrbekov 2017a) and Tunisia (Boukhris-Bouhachem et al. 2007). Oviparae and alate males were collected in October (Hille Ris Lambers 1947c). A second species, S. asperulae Bozhko, was described from Asperula cynanchica in Ukraine, and this name was also applied to populations on A. aristata in Switzerland (Jörg & Lampel 1988). However, the characters used for separation of apterae of S. asperulae from those of S. necopinata (ratio of body width to length and shape of siphunculi) are inconsistent and unreliable, and another collection of Staegeriella from A. aristata in Switzerland (BMNH collection, leg. D. Hille Ris Lambers) differs from both S. necopinata on Galium and from S. asperulae according to Jörg & Lampel (1988), so it appears that a single, variable taxon may be involved.

Staticobium Mordvilko

Aphidinae: Macrosiphini

About 12 species similar to Macrosiphoniella, associated with Plumbaginaceae in salt-marsh and coastal habitats. The taxonomy is very problematic, as several of the nominal species are poorly known and not clearly differentiated in the literature. The better-known species exhibit much geographic variation, accentuated by the fact that populations are distributed linearly along coasts or isolated in salt marshes separated by deserts. Kadyrbekov (2003b) reviewed and keyed the western palaearctic species.

Staticobium caspicum Bozhko Apterae are shining dark brown; BL c. 2.0 mm. On Limonium spp. in Russia (north Caucasus). Decribed as a subspecies of S. latifoliae, but probably just a geographical variant of that species.

Staticobium caucasicum Bozhko Apterae are shining dark brown; BL c.2.2 mm. On upper and undersides of leaves of a Limonium sp. in Ukraine. Stekolshchikov (2005) recorded it (as limonii ssp. caucasicum) from L. gmelinii in southern Russia (Astrakhan), and Kadyrbekov (2011c) from the same host in west Kazakhstan.

Staticobium gmelini Bozhko Apterae are brown, with pale cauda; BL c. 1.9-2.0 mm. On flowerstalks and root collars of Limonium and Goniolimon spp. in Ukraine, west Siberia and Kazakhstan. S. zolotarenkoi (Ivanoskaya) is a synonym according to Kadyrbekov (2003b), and its host was probably Limonium gmelinii rather than Helichrysum arenarium. S. strongilosiphon Bozhko, and perhaps S. insularum, are probably also synonyms.

Staticobium insularum Bozhko Apterae are dark brown with greenish tinge; BL c. 2.0-2.1 mm. On Goniolimon tataricum in Ukraine (Bozhko 1961 – but on Limonium sp. according to Bozhko 1976), and Kadyrbekov (2011c, 2014g) recorded it on Limonium gmelinii in Kazakhstan and Russia. Closely related to S. gmelini, and possibly a synonym.

Staticobium latifoliae Bozhko Apterae are dark brown to blackish; BL 1.6-2.6 mm. On undersides of leaves and flowerstalks of Limonium spp. in Pakistan, Kazakhstan, Tajikistan (Kadyrbekov 2013c), Iran (Rezwani 1990), Lebanon, Russia (north Caucasus, west Siberia), Ukraine, Bulgaria, Romania, Hungary, Greece and Italy (Barbagallo & Stroyan 1982). S. tauricum Bozhko is a synonym according to Kadyrbekov (2003b). S. smailovae Kadyrbekov has morphology intermediate between latifoliae and staticis on the basis of characters given in Kadyrbekov’s (2003b) description and key.

Staticobium limonii (Contarini) Apterae are dark olive-green to dark brown; BL 1.5-2.2 mm. On Limonium spp. in southern Europe and the Mediterranean region. Accounts from north-west Europe are referable to S. staticis (q.v.). The distinction from S. latifoliae seems unclear, and the separate species status of these two taxa requires further confirmation. 2n=12*.

Staticobium longisetosum Kadyrbekov Colour of apterae in life is unknown; BL c.2.0 mm (one specimen). On Limonium gmelinii in south-east Kazakhstan (Kadyrbekov 2003b).

Staticobium loochooense (Takahashi) Colour of apterae in life is unrecorded, probably dark; BL c. 1.4-1.5 mm. On Limonium wrightii in Loochoo (Japan), and on L. tetragonum in Korea (Paik 1972). A sample collected from Aster tripolium were presumably vagrants. Redescribed by Sorin (1967).

Staticobium otolepidis Nevsky Apterae are brown, with antennae dusky except yellow III-IV, siphunculi black with pale bases, and cauda pale yellow; BL 1.6-2.3 mm. In large colonies on flowerstalks of Limonium otolepis and L. perfoliatum in Central Asia and north-west China. S. nevskyi Hille Ris Lambers is probably the fundatrix of this species, and hence a synonym (Kadyrbekov 2003b).

Staticobium smailovae Kadyrbekov Apterae are brownish; BL 1.8-2.2 mm. On stems of Limonium gmelinii in Kazakhstan (Kadyrbekov 2003b) and Russia (Kadyrbekov 2014g).

Staticobium staticis (Theobald) Plate 27f Apterae are shiny, deep green, brown, or dirty greenish or reddish, with antennae paler with dark apices, siphunculi black with pale bases, and cauda green, pale red or yellow (see influentialpoints.com/Gallery); BL 2.3-2.7 mm. On leaves and on upper parts of stems under inflorescences of Limonium spp. especially L. vulgare (= Statice limonium), often flooded by high tides and covered with mud (Foster 1984). In coastal areas of north-west and north Europe. A similar undescribed species but with a shorter antennal terminal process occurs in the Mediterranean region. Oviparae and apterous males occur in September in Germany (Müller 1975b, as S. limonii). The sexuales described by Jacob (1948a, as S. limonii) from L. humile in North Wales were possibly of another closely-related species.

Staticobium suffruticosum Kadyrbekov Apterae are dark greenish or light brownish; BL 1.9-2.3 mm. On stems of Limonium suffruticosum in Kazakhstan (Kadyrbekov 2003b, 2011c). Very close to S. otolepidis.

Stegophylla Oestlund

Phyllaphidinae

Seven species on nearctic Quercus. Alate females apparently do not occur and males may be apterous or alate within a species. Hille Ris Lambers (1966a) keyed the species then known and discussed the confused nomenclature. Quednau (2010) fully reviewed the genus and provided illustrations of all available morphs.

Stegophylla brevirostris Quednau Apterae pale greenish white, producing abundant wax (Halbert 2013); BL 1.2-1.7 mm. On Quercus spp., seeming to prefer Q. virginiana, in eastern USA. An ovipara and an alate male were collected in late September on Q. alba in Maryland (Quednau 2010). Populations on Q. virginiana in Florida seem to be anholocyclic. Ammar et al. (2013) studied the ultrastructure of the wax glands and the wax secretions of this species.

Stegophylla davisi Quednau (new name for Phyllaphis? querci Fitch of Davis 1911a) Apterae pale greenish, BL not recorded. On Quercus spp. of the white oak group (alba, macrocarpa, stellata) in eastern North America, forming small colonies on upper and undersides of leaves (Davis 1911a). Pale brownish oviparae and both apterous and alate males occur in September-October, the apterous male having an alatiform, sclerotised thorax (Quednau 2010).

Stegophylla essigi Hille Ris Lambers Apterae are broadly oval, pale grey-green to olive, covered in white wax wool; BL 1.2-1.8 mm. Feeding by fundatrices in spring causes edges of young leaves to curl upward and become thickened and red. On native Quercus spp. in California, USA, with oviparae and both apterous and alate males produced in November, but populations may continue parthenogenetically on evergreen oaks such as Q. agrifolia, especially in leaves spun together by caterpillars (Hille Ris Lambers, 1966a). It is also recorded from Oregon (A. Jensen, aphidtrek.org). 2n = 12.

Stegophylla mugnozae Remaudiere & Quednau Apterae are broadly pear-shaped, creamy white, covered with white or bluish-white wax wool; BL 1.4-1.8 mm. On leaves of Quercus crassipes and Q. castanea in Mexico (Remaudiere & Quednau 1985), and subsequently recorded from Panama (Quirós et al. 2009). Other morphs and life cycle are unknown.

Stegophylla quercicola (Monell) (= querci of Blackman & Eastop 1994) Apterae are oval, pale greenish- or brownish-yellow, covered in dense white wax wool; BL 1.0-1.5 mm. In small, scattered colonies on upper and undersides of leaves of Quercus spp., usually of the white oak group (e.g. alba, stellata) in eastern North America. Oviparae and alate males occur in September-October (Hottes & Frison 1931; as quercicola Baker).

Stegophylla quercifoliae (Gillette) Apterae are yellowish to yellowish-green with brown head, wholly covered in white wax wool; BL 1.1-1.5 mm. On leaves of Quercus spp., especially Q. gambellii, large colonies causing leaf curl. Oviparae and alate males occur in October. In western USA (California, Colorado, Utah, Idaho).

Stegophylla quercina Quednau (= quercicola of Blackman & Eastop 1994) Apterae are pale yellowish-green with brown head, covered with bluish-white wax wool; BL 1.6-1.9 mm. On leaves of Quercus spp., usually of the red oak group, in dense flocculent masses which may cause curling and yellowing (Quednau, 1966). Oviparae and apterous males occur in October (the males are true apterae without a sclerotised thorax). In eastern North America. Quednau (2010) distinguished two subspecies, with S. quercina s.str. known from Quebec and New York (BMNH collection, leg. R.L. Blackman), and S. quercina acutirostris Quednau, known from Wisconsin and North Carolina. The latter subspecies makes pseudogalls on the infested Quercus spp. (Quednau 2010). 2n > 30 (for quercina s. str.).

Stellariopsis Szelegiewicz

Aphidinae: Macrosiphini

One species in Mongolia possibly related to Spatulophorus, but with more numerous body hairs arising from tubercles, short conical siphunculi and rather hairy R IV+V and cauda.

Stellariopsis songini Szelegiewicz Apterae are pale green, lightly dusted with wax; BL 1.5-2.1 mm. Alatae have secondary rhinaria distributed III 29-34, IV 5-6, V 0-1 (1 specimen). On stems and undersides of leaves of Stellaria dichotoma in Mongolia.

Stomaphis Walker

Lachninae: Stomaphidini

About 30 palaearctic species of very large, stem- or root-feeding aphids, remarkable for their very long rostrum which enables them to probe down through crevices in thick bark of oak and other trees. The morphology of these remarkable mouthparts was studied by Brożek et al. (2015) and their mode of use by Rakitov et al. (2018). Males are small, apterous, lack mouthparts, and often exhibit morphological aberrations. The evolutionary significance of male dwarfism in Stomaphis was discussed by Depa et al. (2015). This could be a stage in the evolutionary loss of males leading to permanent parthenogenesis, but it seems more likely to be due to the sheltered, confined habitat in which males reach maturity in close proximity to females, so that their only function is to mate before they die. The male genitalia of four species were compared by Wieczorek (2012). In some species (subgenus Parastomaphis) the males adopt an unusual, ventro-ventral mating position, with associated morphological modifications in both sexes (Depa et al. 2014). Colonies are invariably attended by ants (usually Lasius spp.). Protaphidius wissmannii is a specialized parasitoid. Accounts and keys are available for Europe (Heinze 1962, Szelegiewicz 1962, Czylok & Blackman 1991), Iberian peninsula (Nieto Nafría et al. 2002), Ukraine (Mamontova 1972), Korea (Lee et al. 1994, Seo 1994), China (Qiao & Zhang 1999d) and Japan (Sorin 2012a). The Acer-feeding species were reviewed and keyed by Binazzi & Blackman (2004). Depa & Kanturski (2014) provide a key to known fundatrices of European species, and Depa et al. (2015) provided illustrations and keys for oviparae and males of European species. Hodgson et al. (2019) provided a key to alatae. Depa & Mróz (2013) studied mitochondrial DNA and morphometric variation in Stomaphis populations on Acer and Salix in Europe and discussed their taxonomic status. These studies were extended to other Stomaphis spp. on other hosts by Depa et al. (2017a). Similar research on Stomaphis spp. in Japan was undertaken by Yamamoto et al. (2020a,b), who investigated how interactions with host plants and ants influenced diversification within the genus; their studies included three undescribed species, and provided first evidence of host alternation in a member of the Lachninae,with migration from a coniferous primary host (see under S. japonica).

Stomaphis abieticola Sorin Colour in life was not recorded; BL of aptera c.6.2 mm. On Abies firma in Japan (Shikoku). Oviparae and apterous males were collected in late October (Sorin 2012a).

Stomaphis aceris Takahashi Colour in life was not recorded; BL of aptera 4.2-4.7 mm. On Acer sp. in Japan (Takahashi, 1960b), attended by Lasius umbratus (Yamimoto et al. 2020). Oviparae and apterous males were collected in November (Sorin 1995).

Stomaphis acquerinoi Binazzi Apterae are elongate oval, greyish white powdered with wax, immatures shiny yellowish white; BL 4.3-6.4 mm. On Acer pseudoplatanus in Italy (Tuscany; Binazzi 2001), in colonies on smooth exposed trunk of young trees, or in bark crevices of older trees. No sexuales were found, and populations are believed to overwinter as alate viviparae (Binazzi & Pennacchio 2002). Ant attendant behaviour and parasitism by Protaphidius wissmannii were studied by Binazzi & Pennacchio (2002, 2005). Depa & Mróz (2013) suggested that this might be a geographical variant of S. graffii.

Stomaphis alni Sorin Apterae are dark brown; BL c. 6mm. On Alnus hirsuta in Japan. Sexual morphs occur in September (Sorin 1965).

Stomaphis aphananthae Sorin Apterae are blackish-brown; BL c. 4.7 mm. On Aphananthe aspera in Japan, attended by Lasius hayashi (Yamimoto et al. 2020). Sexual morphs occur in November (Sorin 1979a).

Stomaphis asiphon Szelegiewicz Colour in life is unknown; BL of aptera 4.8-5.9 mm. Described from roots of an undetermined tree species in Korea (Szelegiewicz 1975). Pashchenko (1988b) recorded it living in cracks in bark at bases of trunks of Juglans mandshurica in eastern Siberia. Sexual morphs of S. asiphon s.str. are undescribed. However, Sorin (2012a) described fundatrices, apterae (fundatrigeniae), oviparae and apterous males of Japanese populations on Juglans mandshurica ssp. sieboldiana (= J. ailanthifolia) as a subspecies, S. asiphon ssp. sakuratanii. The apterae had fully developed siphuncular cones, although these were absent in fundatrices; they will run to S. matsumotoi in our key to aphids on Juglans, and the distinction from that species seems to require further confirmation. The aphids were living on the surface of the trunk at or just below ground level, attended by ants, and sexual morphs were produced in November.

Stomaphis betulidahuricae Zhang & Qiao Apterae are powdered with greyish-white wax; BL c. 5.6 mm. In bark crevices of Betula dahurica in Hebei, China (Qiao & Zhang 1999d). Alatae and sexual morphs are undescribed.

Stomaphis bratislavensis Czylok & Blackman Apterae are brownish, coated with powdery wax; BL 4.3-4.8 mm. On bark of Quercus petraea in Slovakia, and subsequently recorded from Bulgaria and Turkey (Şenol et al. 2014). Sexual morphs occur in October (Czylok and Blackman, 1991). The ovipara was redescribed by Depa et al. (2015). 2n = 8 (Blackman, 1990).

Stomaphis carpini Sorin Colour in life is unrecorded; BL of aptera c. 5.5 mm. On Carpinus tschonoskii in Japan (Sorin, 1965). Sexual morphs are undescribed.

Stomaphis cupressi Pintera Apterae are yellowish-brown, powdered with grey wax; BL 4.9-6.2 mm. Described from specimens collected on trunk of Cupressus benthami in Kenya (Pintera, 1965), and since found on C. macrocarpa in France (BMNH collection, leg. G. Remaudiere), on Cupressus spp. in Italy (BMNH collection, leg. D. Roberti), on Juniperus spp. in Spain (Nieto Nafría et al. 2002), and on C. sempervirens in Turkey (Akyürek et al. 2010). Mamontova (1982) described a population on C. sempervirens in Abkhazia (Caucasus) as a subspecies, S. cupressi sp. caucasica. Sexual morphs are unknown. 2n = 14.

Stomaphis fagi Takahashi Colour in life was not recorded; BL of aptera c. 5.3mm. On Fagus sp. in Japan (Takahashi, 1960b). An oviparae and an apterous male were described by Sorin (1995), collected in September .

Stomaphis (Parastomaphis) graffii Cholodkovsky Apterae are yellowish, covered in white wax powder; BL 4.2-6.5 mm. In bark crevices on trunks of Acer spp., usually A. campestre, A. platanoides or A. pseudoplatanus, often at or below soil level, attended by Lasius spp. (brunneus, balcanicus). In Europe (UK, France, Italy, Poland, former Czechoslovakia, Rumania, Ukraine), and east to Georgia. Sexual morphs have been collected from mid-August to November; they were redescribed by Depa et al. (2015). Depa & Kanturski (2014) described the fundatrix. Depa (2012, 2013) studied populations and life cycle in Poland. A population in Ukraine was described by Mamontova-Solukha (1963, 1972) as a subspecies, S. graffii ssp. acerina, but this was placed as synonym of S. graffii graffii by Depa & Mróz (2013), who compared mitochondrial DNA and morphometric data (see also under S. longirostris).

Stomaphis hirukawai Sorin Appearance in life unrecorded: BL c. 5.4 mm. On Chamaecyparis obtusa in Japan (Sorin 1995), attended by Lasius spp. (Yamimoto et al. 2020). Oviparae collected in September were included in the original description. Matsuura & Yashiro (2006) studied the behaviour of ants tending the eggs of this species.

Stomaphis japonica Takahashi Plate 15f Apterae are shiny brown, sometimes wax-dusted; BL c. 5 mm. On trunks of Quercus spp. in Japan (Takahashi 1960b; as S. quercus japonica), attended by Lasius spp. (Yamimoto et al. 2020), and also recorded from China (Qiao & Zhang 1999d) and Korea (Seo 1994). Sexual morphs occur in Japan in October-November. Takada (2008) found that sexual reproduction in the region of Kyoto only took place on Q. serrata. He observed a partial migration to other Quercus spp. (acutissima, variabilis) for the summer months, with alatae leaving these hosts in October-November. Yamamoto et al. (2020b) have now used a combination of mtDNA and multivariate morphometric analysis to demonstrate that Japanese populations identified as S. japonica comprise lineages with different life-cycles. One of these lineages migrates from primary host Pinus densiflora to secondary host Quercus acutissima (and is probably a junior synonym of S. pini), and another lives without hosrt alternation on Q. serrata (and possibly Q. dentata). The taxonomy and nomenclature of this group still needs to be resolved. Kamiju & Takada (1983) studied a parasitoid and hyperparasitoid of S. japonica. 2n = 10 (Blackman 1990).

Stomaphis (Parastomaphis) juglandis Petrović Apterae are elongate oval, shiny dark brown, sometimes wax-dusted; BL 5.7-6.7 mm. On trunk of Juglans regia in Serbia (Petrović 1998). The original description included alate viviparae collected in July and oviparae and apterous males collected in October; the sexual morphs were redescribed by Depa et al. (2015).

Stomaphis knechteli Binazzi & Blackman Colour in life unknown; BL of aptera 4.5-5.6 mm. On trunk of Acer campestre in Rumania (Binazzi & Blackman 2004). An alata was collected in August. Depa & Mróz (2013) suggested that this might be a geographical variant of S. graffii.

Stomaphis liquidambaris Takahashi Apterae are blackish; BL c. 5.2mm. On stems of Liquidambar formosana in Taiwan (Takahashi, 1925) and China (Ningxia; Qiao & Zhang 1999d). Sexual morphs are unknown.

Stomaphis (Parastomaphis) longirostris (Fabricius) Apterae are whitish powdered with greyish-white wax (see influential points/Gallery); BL 6.0-6.5 mm. On trunks of Populus spp. (nigra, tremula) and Salix spp. (alba, viminalis). There is also a record of a population on Acer campestre in Tuscany (Binazzi & Blackman 2004; but see Depa & Mróz 2013). Tashev’s (1961) record from Quercus sp. in Bulgaria may be an error; however, the species resembling S. graffii that he found on Populus nigra is possibly longirostris. In continental Europe (France, Netherlands, Germany, Austria, Czech Republic, Slovakia, Italy, Spain, and probably UK), and east to Turkey and western Siberia. Depa & Kanturski (2014) described the fundatrix. Depa et al. (2015) described the sexual morphs, collected in September- October. Depa & Mróz (2013) compared mitochondrial DNA and morphological parameters of S. longirostris with those of the closely-related European Acer-feeding species (graffii, acquerinoi, knechteli), and concluded that the key characters used for distinguishing species in this group were unreliable.

Stomaphis malloti Sorin Apterae are dark brown; BL c. 5.2mm. On Mallotus japonicus in Japan, attended by Lasius hayashi (Yamimoto et al. 2020). Sexual morphs occur in October-November (Sorin 1979a, Takada 2009).

Stomaphis matsumotoi Sorin Apterae are dark brown; BL c. 6.2 mm. On (trunk of?) Juglans mandshurica var. sachalinensis (= S. ailanthifolia) in Japan, attended by Lasius spp. (Yamimoto et al. 2020). Sexual morphs occur in October (Sorin 1995).

Stomaphis mordvilkoi Hille Ris Lambers Apterae are dark brown; BL 5.3- 7.3mm. On trunk of Juglans regia; described from West Bengal, India (Hille Ris Lambers 1933b) and subsequently rediscovered in northern Italy (Colombo 1982). It is also now recorded from Hungary (Szathmáry et al. 2005). Sexual morphs are unknown.

Stomaphis pini Takahashi Colour in life unknown, probably blackish; BL c. 4.5 mm. On basal part of trunk or roots of Pinus densiflora in Japan (Takahashi, 1960b; Inouye, 1970). There is also a record from P. halepensis in Algeria (Laamari et al. 2013; it needs to be confirmed that this is the same species). Oviparae and apterous males collected in early December were described by Sorin (1995). Apterae and an ovipara collected in late August on Abies firma in Japan were described by Sorin (2012a) as a subspecies, S. pini ssp. takaoensis. [Following the work of Yamamoto et al. (2020b), it now seems probable that the host-alternating form currently under the name S. japonicus (q.v.) is a synonym of S. pini.]

Stomaphis pistacicola Zhang & Qiao Apterae are greyish brown; BL c. 4.3 mm. On Pistacia sinensis in China (Qiao & Zhang 1999d). Other morphs and life cycle are unknown.

Stomaphis pterocaryae Sorin Colour in life was not recorded; BL of aptera c.5.3 mm. On Pterocarya rhoifolia in Japan (Honshu), attended by Lasius sp(p) (Yamimoto et al. 2020). Oviparae and apterous males were collected in October (Sorin 2012a).

Stomaphis quercisucta Qiao & Zhang Apterae are elliptical, greyish white, dusted with white powder; BL c. 5.2 mm. On Quercus sp. in Hebei, China (Qiao & Zhang 1999d). Apart from the appearance in life the description does not indicate any clear distinction from the western palaearctic species S. quercus.

Stomaphis quercus (Linnaeus) Plate 15e Apterae are elliptical to elongate oval, shining dark green or brown (see influential points/Gallery); BL 5.3-7.0mm. In bark crevices on trunk of Quercus spp., also on Betula, and possibly Alnus glutinosa (although the latter record should perhaps be applied to S. wojciechowskii). Throughout Europe, and in west Siberia and Kazakhstan (where all the records are from Betula; Kadyrbekov 2017a). Sexual morphs occur in September-November; they were redescribed by Depa et al. (2015). Goidanich (1958) made a detailed study of its life cycle and relation with the attendant ant Lasius fuliginosus, and Lorenz & Scheurer (1998) made a comparable study of populations on Betula pendula. In Sardinia it is associated with L. brunneus on Q. suber, causing damage to cork bark (Loi et al. 2012). Depa et al. (2012) confirmed using mitochondrial DNA that the populations on Quercus and Betula are conspecific, and that S. betulae Mamontova, described from Betula in the Ukraine, is a synonym. 2n = 10.

Stomaphis radicicola Hille Ris Lambers Apterae are broadly oval, powdered with grey dust; BL 4.5-6.0mm. On roots of Betula sp., deep underground, attended by Lasius umbratus,in the Netherlands (Hille Ris Lambers 1947b). Depa et al. (2013) have now described the ovipara, collected in October on Alnus glutinosa in Slovakia, and the male was described by Depa et al. (2015). This specimen was in a deep bark crevice above ground level, attended by Myrmica rubra.

Stomaphis rhusivernicifluae Zhang Appearance in life is unknown; BL of aptera c. 5.4mm. On Toxicodendron vernicifluum (= Rhus verniciflua) in Yunnan, China (Zhang & Zhong 1985e). Other morphs and biology are unknown.

Stomaphis sinisalicis Zhang Apterae are without wax powder; BL c. 6.0 mm. On Salix matsudana, collected at Beijing, China (Zhang & Zhong 1982b); subsequently recorded from Populus sp., and from other parts of China (Qiao & Zhang 1999d). Other morphs and biology are unknown.

Stomaphis takahashii Sorin Appearance in life is not recorded; BL of aptera 4.7- 5.0 mm. On Acer spp. and Betula latifolia in Japan (Hokkaido). Sexual morphs occur in September (Sorin 1965).

Stomaphis ulmicola Inouye Apterae are shiny brown-black; BL c. 6.3mm. On trunk of Ulmus japonica in Hokkaido, Japan (Inouye, 1938; Takahashi, 1960b), attended by Lasius spp. (Yamimoto et al. 2020), and also recorded from Hebei, China (Qiao & Zhang 1999d). Sorin (1995) described oviparae and an apterous male collected in October.

Stomaphis wojciechowskii Depa Apterae are elliptical, pale fuscous to pale brownish, dull, slightly powdered with wax, and with a series of paired brown pleurospinal sclerites on either side of a pale spinal band; BL 6.1-6.8 mm. On basal part of trunk of Quercus robur, at ground level or up to c.1.8m above ground, in ant shelters constructed within bark crevices and covered by soil (Depa et al. 2012). Recent work has shown that it has a wide host range including other Quercus spp., Alnus glutinosa, Juglans regia, Salix sp. and Tilia cordata (Depa et al. 2017a). The attendant ants are usually Lasius brunneus, never L. fuliginosus (cf. S. quercus). Recorded from Central and Eastern Europe (Poland, Hungary, Romania, Slovakia). Alatae collected in the UK and Poland were described by Hodgson et al. (2019). Sexual morphs occur in September to early November; they were redescribed by Depa et al. (2015). Depa & Mróz (2012b) described the fundatrix, collected from Q. robur in early June. Depa et al. (2017a) proposed a possible role of specific ant-aphid relationships in driving the divergence between this species and S. quercus.

Stomaphis yanonis Takahashi Apterae are grey-brown with two rows of dark brown dorsal patches; BL c. 5.0mm. In crevices on bark of trunk of Celtis sinensis and Zelkova serrata, in Japan, Korea, China and Taiwan, attended by Lasius spp. (Yamimoto et al. 2020). Takahashi (1960b) described a population on Aesculus turbinata as a subspecies, S. yanonis ssp. aesculi. Sexual morphs occur in October-November (Sorin, 1965, Takada 2009). 2n = 20 (Honda 1921) or 16? (Blackman 1990).

Strenaphis Quednau

Calaphidinae: Saltusaphidini

A genus for one distinctive aphid related to Saltusaphis but with a dome-like protrusion of the front of the head. The name is a replacement for Stenaphis Quednau 1954 which is preoccupied by Stenaphis del Guercio 1913 (Quednau in Favret et al. 2007, p.397).

Strenaphis elongata (Baker) Apterae are very long-bodied, yellow to orange-yellow with paired dark orange flecks on dorsal abdomen; BL 2.5-3.1 mm. An alata trapped in Canada (New Brunswick) was illustrated by Quednau (2010). On Carex spp. (stricta, strictior) in eastern North America. Oviparae (original description) and apterous males (Hottes & Frison 1931) occur in October. 2n=10.

Subacyrthosiphon Hille Ris Lambers

Aphidinae: Macrosiphini

One species living on Trifolium in north-west Europe, with characters between Acyrthosiphon and Aulacorthum. The peculiar habitat, the presence of antesiphuncular sclerites in the aptera, and the unusual antennal sensoriation of the alatae which have 2-14 rhinaria usually restricted to the basal half of ANT III, are distinctive features.

Subacyrthosiphon cryptobium Hille Ris Lambers Plate 24a (Fig.53c) Apterae are pale olive-green with head sometimes faintly reddish; BL 1.6-2.3 mm. Living hidden on older parts of prostrate stems of Trifolium repens, and dropping when disturbed so that they are rarely observed or collected. The plant shows no reaction. Monoecious holocyclic, with apterous males. In Europe, and introduced to USA.

Subiziphya Quednau

Calaphidinae: Saltusaphidini

One nearctic species related to Iziphya and Saltusaphis but with exclusively mushroom-shaped dorsal hairs and a distinctive, rounded and very hairy abdominal tergite 8.

Subiziphya clauseni Quednau Apterae are long-bodied, colour in life unknown but probably dark, with antennae black except at bases; BL c.1.8 mm. Described from Poa sp., but the true host must surely be Cyperaceae, probably a Carex species (hence inclusion in the key to sedge aphids). Alatae have broad dark dorsal abdominal cross-bars, bordered wing veins with spots at their apices, and 15-17 secondary rhinaria on ANT III (see also Quednau 2010).

Subovatomyzus A.N. Basu

Aphidinae: Macrosiphini

One species on Lamiaceae in India resembling Capitophorus, with short capitate dorsal hairs and smooth clavate siphunculi.

Subovatomyzus leucosceptri A.N. Basu Apterae are spindle-shaped, pale translucent green with a darker green spinal stripe, and whitish appendages; BL 1.5-1.9 mm. Alatae have blackish head, thorax and antennae, a dark central abdominal patch, and numerous secondary rhinaria on ANT III-V. On undersides of leaves of various labiate species in several genera (Callicarpa, Colebrookia, Elsholtzia, Leucosceptrum, Pogostemon), and also recorded from Vernonia (Compositae; as Capitophorus vernoniae A.K.Ghosh & Raychaudhuri), but these were possibly vagrants. Described from West Bengal, India, where it is most common in April to June and reproduces parthenogenetically throughout the year (original description). Also recorded from Sikkim and Meghalaya (A.K. Ghosh 1973), and from Nepal (BMNH collection).

Subsaltusaphis Quednau

Calaphidinae: Saltusaphidini

About 12 Carex-feeding long-bodied aphids with spatulate empodial hairs, and with dorsal hairs mostly very short and mushroom-shaped, with an incised “cap” so that they are stellate in dorsal view. Apterae lack secondary rhinaria. Alatae have a dark central abdominal patch on tergites 3-5, incised intersegmentally. Richards (1971) reviewed the world fauna, and there are reviews of the European species by Stroyan (UK, 1977), Heie (north-west Europe, 1982), and Nieto Nafría & Mier Durante (Iberian peninsula, 1998) and of the Japanese species by Sorin (2005). Quednau (2010) reviewed the world fauna, provided keys and illustrated all available morphs.

Subsaltusaphis aquatilis (Ossiannilsson) Apterae are long-bodied, pale yellowish, with dark intersegmental muscle sclerites; BL 2.1-2.6 mm. On Carex spp. (aquatilis, acutiformis, riparia) in northern and central Europe. Alatae have 10-17 secondary rhinaria on ANT III (Heikinheimo 1984). It is almost indistinguishable from the early seasonal form of S. picta. 2n=8 (cf. picta).

Subsaltusaphis canadensis Richards Apterae are long bodied, pale yellow with black antennae; BL c. 2 mm. On Carex sp. in Ontario, Canada. Alatae have 8-12 secondary rhinaria on ANT III. Apart from its small size and lack of dorsal pigmentation this species closely resembles S. lambersi. It could also be a variant form of S. virginica (Quednau 2010).

Subsaltusaphis flava (Hille Ris Lambers) (Fig.20c) Apterae are dull yellow, with dark intersegmental muscle sclerites (no longitudinal dark stripes), and antennae black except at bases; BL 1.8-2.1 mm. Alatae have 8-9 secondary rhinaria on ANT III. On Carex nigra in Europe. Oviparae occur in October in Denmark (Heie 1982), and the male genitalia were described and illustrated by Wieczorek (2011). 2n=8.

Subsaltusaphis intermedia (Hille Ris Lambers) Apterae are yellowish white, with rather indistinct dark intersegmental muscle sclerites, and antennae black except at bases; BL 1.9-2.2 mm. Alatae have 9-12 secondary rhinaria on ANT III. On Carex hirta, living between the leaf bases on “dry waysides”. In Europe (Netherlands, Austria, Germany, Czech Republic).

Subsaltusaphis lambersi (Quednau) Apterae are pale yellowish, with narrow brownish intersegmental muscle plates, and a pair of brownish, sometimes rather vague and incomplete, pleural longitudinal stripes; BL 2.4-2.8 mm. On Carex acuta in Sweden, apparently preferring shady habitats and waterside situations (Ossiannilsson 1959). It is also recorded from Finland (Albrecht 2010), Poland (Achremowicz 1972) and Spain (Nieto Nafría & Mier Durante 1998). Oviparae were found in Sweden in early September. Alatae are undescribed. Similar aphids described from Carex ?reinii in Japan are treated by Quednau (2010) as a subspecies, S. lambersi kamijiensis Sorin. This Japanese form has alatae with 12-18 secondary rhinaria on ANT III, and its oviparae were found in late January (Sorin 2005). 2n=6 for ssp. kamijiensis (Blackman 1980, erroneously listed as S. saracola).

Subsaltusaphis ornata (Theobald) (Fig.20e) Apterae are yellow to ochreous yellow with two very distinct pleural longitudinal dark lines, clearly defining a pale spinal band; BL c. 2.7-2.8 mm. Alatae have c.13 secondary rhinaria on ANT III. On various Carex spp., with a record also from Scirpus sylvaticus. In Europe, Iran (BMNH collection), and now also recorded from Kazakhstan (Kadyrbekov 2014e). 2n=8.

Subsaltusaphis pallida (Hille Ris Lambers) (Fig.20b) Apterae are pale yellow, with dark intersegmental muscle sclerites; BL c.2.2-2.7 mm. Alatae are unknown. On Carex spp. (gracilis, stellulata) on river banks, and there is also a record from Juncus (Shaposhnikov 1964). In Europe, Central Asia ((Kazakhstan) and east Siberia (Quednau & Shaposhnikov 1988).

Subsaltusaphis paniceae (Quednau) Apterae are pale yellow, with dark intersegmental muscle sclerites and variably developed longitudinal dark markings similar to those of S. picta (q.v.), and with antennae banded on segments III-V, VI being wholly dark; BL 1.3-2.3 mm. On Carex spp. in wet meadows in Europe (UK, Sweden, Poland, Germany, Italy, Spain). Very similar to the North American S. virginica.

Subsaltusaphis picta (Hille Ris Lambers) Plate 4h Apterae are whitish yellow, with dark transverse intersegmental muscle sclerites, and somewhat less dark and variably developed longitudinal dark markings, the most heavily marked specimens having dark marginal and pleural stripes and a shorter spinal stripe; antennae black beyond basal half of segment III (see influentialpoints.com/Gallery); BL 2.4-3.0 mm. Alatae have 11-16 secondary rhinaria on ANT III. On Carex spp. growing at the edge of water, and sometimes also on Scirpus in the same situation. Widely distributed in Europe; also recorded from Kazakhstan, and east Siberia (Quednau & Shaposhnikov, 1988). Oviparae and apterous males occur in September-October in Sweden and Scotland (Heie 1982 and BMNH collection, leg. H.L.G. Stroyan). 2n=10.

Subsaltusaphis (Primoriaphis) primoriensis Quednau & Shaposhnikov Apterae are undescribed, alatae are pale yellow with blackish grey markings, and 4 rhinaria on ANT III; BL c 1.6 mm. On Carex laevirostris in east Siberia.

Subsaltusaphis rossneri (Börner) (Fig.20d) Apterae are ochreous yellow, usually with dusky pleural longitudinal stripes on head and thorax, and paired segmental patches plus a faint spinal stripe on abdomen; BL c.2.3-2.5 mm. Alatae have 10-13 secondary rhinaria on ANT III. On Carex spp. (elata, nigra, rostrata, ?vesicaria) in Europe, and one alata is recorded from Pakistan (Naumann-Etienne & Remaudière 1995).

Subsaltusaphis (Primoriaphis) taoi Hsu ex Tao (= pulchra Quednau & Shaposhnikov; synonymy by Quednau 2010) Apterae are lemon yellow, slightly shiny, with four longitudinal black stripes; BL c. 2.3mm. Alatae have 6-9 secondary rhinaria on ANT III. On Carex sp. In Taiwan, and on C. rubra in east Siberia (Quednau & Shaposhnikov 1988). Sorin (2005) redescribed apterae, alatae and oviparae from Japan, collected on C. dispalata.

Subsaltusaphis virginica (Baker) Apterae are pale yellow, with or without faint dark longitudinal pleural and marginal stripes, and sometimes a very faint spinal stripe. The antennae are pale basally and dark distally, with the base of segment IV often paler than the distal part of III but clearly darker than I and II. BL 1.9-2.9 mm. Alatae have 11-15 secondary rhinaria on ANT III. On Carex spp. in North America (records mainly from eastern US and Canada, but also from Utah) and Japan (as S. saracola Higuchi). Oviparae were found in Illinois in early November, on both Carex and Scirpus eriophorum (= cyperinus); BMNH collection, leg. T.H. Frison and H.H. Ross. [Normally this species has a pair of long posterior marginal hairs on abdominal tergite 7, and the posterior edge of this tergite projects laterally, as is typical for Subsaltusaphis. However, a collection of specimens from North Carolina (BMNH collection., leg. D. Hille Ris Lambers, no. 234) has no long marginal hairs on abdominal tergite 7, only mushroom-shaped ones, and the posterior margin does not project laterally. It is possible that two species are involved, but no other distinguishing features between the two forms were apparent.]

Subtakecallis Raychaudhuri & Pal

Calaphidinae: Therioaphidini

Two bamboo-feeding species in India, resembling Takecallis in the anterior projection of the clypeus, but differing in that the ANT PT/BASE ratio is only 0.24-0.55 and the tibiae are evidently glandular. A.K. Ghosh & Quednau (1990) reviewed the genus and Quednau (2003) illustrated all available morphs.

Subtakecallis brevisetosa Raychaudhuri & Pal All viviparae are alate, appearance in life not recorded, probably wax-covered; BL 1.9-2.3 mm. On unidentified bamboo in West Bengal, India (Raychaudhuri & Pal 1974). Sexual morphs are unknown.

Subtakecallis pilosa (David, Rajasingh & Narayanan) All viviparae are alate, covered in white powdery wax, with brown antennae, legs and dorsal abdominal sclerites; BL 2.3-2.6 mm. On Bambusa sp(p). in north-east India (Meghalaya, West Bengal). Oviparae and males were described by Agarwala & Mahapatra (1990).

Sumatraphis Takahashi

Greenideinae: Cervaphidini

One oriental species on Ulmaceae with flattened apterae having corrugated dorsal cuticle and protruding spiracles.

Sumatraphis celti Takahashi Apterae are dull green (in Sulawesi; J.H. Martin, pers. comm.) or yellow to deep brown or brownish green (in India; A.K. Ghosh et al. 1971e, A.K. Ghosh & Agarwala 1993); BL 1.5-1.7 mm. On new growth of Celtis tetrandra or ?Trema orientalis; the apterae are flattened into angles of new shoots (J.H. Martin, pers. comm.). In India it is also recorded from Mallotus sp., Quercus sp. and Sida sp., but these may be “casual” visits. In Indonesia (Sulawesi, Sumatra), India (Manipur, Meghalaya, Sikkim, Tamil Nadu, West Bengal), Nepal and China (Yunnan). The alate vivipara was described by A.K. Ghosh & Agarwala (1993), who also noted that S. celti is active in India during the winter months. Sexual morphs are unknown. Sumatraphis tubercaudatus Qiao & Zhang, described from Celtis sp. in China (Qiao & Zhang 2000a), is probably a synonym.

Swirskiaphis Hille Ris Lambers

Aphidinae: Aphidini

One species in the Middle East apparently related to Aphis, but with long thick dorsal hairs. A second species described in this genus from China, S. bambuciepula Zhang & Zhang is probably a Melanaphis, perhaps M. pahanensis.

Swirskiaphis polychaeta Hille Ris Lambers Plate 8e Colour of apterae in life is unknown; BL 1.9-2.6 mm. Alatae, described by Remaudière et al. (2006), have c.15-16 secondary rhinaria on ANT III. On Umbelliferae/Apiaceae (Dorema, Ferula, Prangos) in the Middle East (Iran, Lebanon, Turkey). The life cycle is unknown.

Symydobius Mordvilko

Calaphidinae: Calaphidini

About eight species of medium to large, rather shiny dark brown aphids associated with Betula and Alnus, forming colonies on the branches and twigs, invariably attended by ants. All the apterae (even the oviparae) are ‘alatiform’, with a similar pattern of sclerotization and pigmentation to that of the fully-winged adults. Monoecious holocyclic; males are apterous, and oviparae have the posterior abdominal segments extended into an ovipositor-like structure. Qiao & Zhang (2002b) reviewed the genus for China; they erected a subgenus, Antisymydobius, which becomes a junior synonym of Yezocallis Matsumura 1917, this having the same type species (S. kabae). Sorin (2014) reviewed and keyed the Japanese species. Trioxys betulae is a specialized parasitoid.

Symydobius alniarius (Matsumura) Reddish or yellowish-brown, with dark brown cross-bands and often with white powder on posterior segments; BL 3.1-3.4 mm. On stems and twigs of Alnus spp. in Japan (Higuchi 1972), Korea and eastern Siberia (Primorskiya Kray). The ovipara, collected in Japan in September, was described by Sorin (2014), who also described a subspecies (nipponica), collected not only on Alnus spp. but also on Betula ermanii. 2n = 20.

Symydobius americanus Baker Apterae are shiny dark brown to black (see influential points/Gallery); BL 2.4-2.7 mm. On stems and twigs of Betula papyrifera in northern USA and Canada. Sexual morphs occur in September-October (Baker 1918).

Symydobius intermedius Gillette and Palmer (fig. 32B) Olive green to dark grey-brown; BL 2.2-2.6 mm. On stems and twigs of Betula occidentalis in western USA (Colorado, Utah, Idaho) and also recorded from British Columbia. Sexual morphs occur in September-October (Gillette & Palmer 1931). 2n = 16.

Symydobius kabae (Matsumura) (fig. 29C, fig. 32C) Dark brown; BL 2.5-3.0mm. On bark of last year’s branches and twigs of Betula maximowicziana and B. platyphylla var. japonica in Japan (Higuchi 1972); also recorded from B. gmelini in Mongolia (as alniarius – see Holman and Szelegiewicz 1974), from B. platyphylla var. latifolia and Alnus (hirsuta var. sibirica, japonica) in Korea and from B. dahurica in eastern Siberia (Primorskiya Kray). Sorin (2014) described ovipara and apterous/alatiform males, collected in Japan in late September-October. Paik (1972) recorded oviparae in Korea in November. S. paucisensorius Zhang & Zhang, described from Betula sp. in China (G. Zhang et al. 1993a), is probably a synonym, and S. careofasciatus, described from apterae found on grasses (Qiao & Zhang 2002b), are probably vagrant individuals of S kabae. 2n = 26? (Shinji 1931; but see Blackman 1986).

Symydobius minutus Quednau and Shaposhnikov Yellowish-brown, with dark transverse bars and white wax powder posteriorly; BL 2.2-2.4 mm. On shoots of Alnus japonica in eastern Siberia (Primorskiya Kray) and Korea (Quednau & Shaposhnikov 1988). Sexual morphs are unknown.

Symydobius nanae Holman Brown to dark brown, with dark appendages except for two pale rings on distal halves of antennae; BL not recorded, probably 2-3 mm. In small aggregations on young branches of Betula nana; only known from apterous viviparae, oviparae and males collected in September in Czech Republic (Holman 1996).

Symydobius oblongus (von Heyden) (fig. 32D) Shiny dark brown (see influentialpoints.com/Gallery); BL 2.0-3.5 mm. On twigs, young stems and branches of Betula spp. throughout Europe and across Asia. Sexual morphs occur in October-November (Heie 1982). Holman (1990) studied developmental morphology of fundatrices and their progeny, and Fowler & Macgarvin (1985) studied the effects of ant attendance on population growth. See Heie (1982) and Stroyan (1977) for general accounts. Two species described by Qiao & Zhang (2002b) from Betula platyphylla in China, S. fumus and S. brevicapillus, are close to S. oblongus and may be variant populations of that species. 2n(female) = 15, 2n (male) = 14 (Blackman 1988).

Symydobius quednaui Sorin Dark brown with blackish-brown dorsal abdominal cross-bands, and dark brown antennae and legs; BL 2.5-3.4 mm. On Alnus spp. in montane regions of Japan (Sorin 2014).

Szelegiewicziella Holman

Aphidinae: Aphidini

One species in east Asia apparently related to Aphis, but apterae and alatae both have 4-segmented antennae, and apterae have a smooth dorsal sclerotic shield.

Szelegiewicziella chamaerhodi Holman Apterae are green with greyish wax exudation; BL 1.2-1.45. Alatae have just 1-3 secondary rhinaria on ANT III. Described from specimens collected on flower stems of Chamaerhodis erecta in August in Mongolia, and also recorded from eastern Kazakhstan (Kadyrbekov 2009a). Similar aphids were collected on Artemisia vulgaris in June in China (Liaoning province; Zhang & Zhong 1990). The dates of collection do not support a host alternation from Chamaerhodes to Artemisia, so this possibility needs to be tested experimentally.