The Aphids

Nine species on trees in various families, but especially Myrtaceae, in India, and east and south-east Asia. In several species only alate viviparae are known, but this may be because apterous viviparae are well camouflaged on the stems and easily overlooked, whereas the alatae develop on the leaves. One species (dineni) was described from an unidentified species of Combretaceae. Accounts are available for Java (Noordam & Hille Ris Lambers 1985) and India (A.K. Ghosh & Quednau 1990), and Quednau (2010) keyed the world fauna and illustrated all available morphs.

Taiwanaphis atrovirens Noordam & Hille Ris Lambers Alatae have head, pronotum and sides of abdominal segments 3-8 yellowish or brownish, mesonotum and central part of dorsal abdomen dark green; BL 1.2-1.5 mm. Immature alatae are yellowish-white with a dorsal greenish-black area. Apterous viviparae, other morphs and life cycle unknown. On upper and lower sides of new leaves and developing shoots of Syzygium sp. in Java (Noordam & Hille Ris Lambers 1985).

Taiwanaphis atuberculata Noordam & Hille Ris Lambers Alatae are orange with grey sides to thorax and abdominal segments 1-5, banded antennae and black-bordered forewing veins; BL 1.5-1.7 mm. Other morphs and life cycle unknown. On upper sides and undersides of leaves and sometimes on developing shoots of Syzygium lineatum in Java.

Taiwanaphis decaspermi Takahashi Apterae and alatae are brownish-black to black; BL 1.2-1.6 mm. Alatae have thickly bordered forewing veins. Described from Decaspermum fruticosum in Taiwan, attacking undersides of young leaves and shoots. Subsequently recorded from other Myrtaceae (Syzygium buxifolium, hancei) in China (Guangdong, Hainan, Hong Kong), and Japan. Alate males were collected in Hong Kong in March (BMNH collection, leg. Commonwealth Institute of Entomology).

Taiwanaphis dineni Mandal, Agarwala & Raychaudhuri Alatae have thickly-bordered forewing veins, colour in life is unrecorded; BL 1.8-2.0 mm. On an unidentified species of Combretaceae in Sikkim, India (Mandal et al. 1979). Apterae are unknown, but an alate male and an ovipara were collected in October (A.K. Ghosh & Quednau 1990).

Taiwanaphis kalipadi (Raychaudhuri & Ghosh) Colour in life is not recorded; apterae are rather elongate oval, BL 1.2-1.5 mm. On Annona squamosa in West Bengal, India (Raychaudhuri & Ghosh 1964). Other morphs and biology are unknown, but possibly T. randiae, known only from alatae, is this species.

Taiwanaphis memecyloni Noordam & Hille Ris Lambers Alatae are yellow with black sides to thorax and grey-bordered forewing veins; BL 1.0-1.5 mm. On upper surfaces of young leaves, with immatures positioned close against the main vein, on Memecylon lilacinum (= laevigatum, myrsinoides) in Java (Noordam & Hille Ris Lambers 1985). Other morphs and life cycle are unknown.

Taiwanaphis montanicola Noordam & Hille Ris Lambers Apterae and alatae are brown, the alatae having black-bordered forewing veins; BL 1.7-2.2mm. On the undersides of young leaves of Syzygium racemosum in Java (Noordam & Hille Ris Lambers 1985). The life cycle is unknown.

Taiwanaphis pseudocaudata Noordam & Hille Ris Lambers Alatae have head and prothorax yellow, mesothorax and abdomen orange with grey sides; BL 1.1-1.6 mm. Along veins on upper and lower sides of young leaves and shoots of Syzygium syzygioides in Java (Noordam & Hille Ris Lambers 1985). Other morphs and life cycle are unknown.

Taiwanaphis randiae Ghosh, Banerjee & Raychaudhuri Alatae are yellowish with dark markings and heavily bordered forewing veins; BL 1.7-2.0mm. On upper and lower sides of terminal leaves of Randia sp. in India (Arunachal Pradesh), attended by ants (A.K. Ghosh et al. 1971d). Other morphs and life cycle are unknown; possibly a synonym of T. kalipadi, which is only known from apterae.

Taiwanomyzus Tao

Aphidinae: Macrosiphini

About six palaearctic species mostly on Saxifragaceae or ferns, perhaps related to Utamphorophora but with dorsal spiculation of the head. Apterae usually have secondary rhinaria extending in a row along ANT III.

Taiwanomyzus alpicola (Hille Ris Lambers) (Fig.43m) Apterae are shining black with antennae and femora mainly black, and dark siphunculi and cauda; BL 1.1-1.7 mm. On ferns, (Asplenium, Athyrium, Blechnum, Cystopteris, Dryopteris, Gymnocarpium, Polypodium), particularly those growing in shady situations (Müller 1987). In central and eastern Europe (Germany, Poland, Switzerland, Slovakia, Czech Republic). Monoecious holocyclic with oviparae and apterous males in September (original description, as Utamphorophora).

Taiwanomyzus babai Sorin & Arakawa Colour of apterae in life is unrecorded; BL c.2.3 mm. Alatae have secondary rhinaria distributed III 30-34, IV 5-7, V 0. On Osmunda japonica in Japan (Sorin & Arakawa 2005).

Taiwanomyzus chrysosplenii Miyazaki Apterae are pale brown to reddish brown, with black antennae and cauda, siphunculi also blackish but often with a paler middle section; 1.3-1.6 mm. On Chrysosplenium flagelliferum, feeding on undersides of leaves in a wet environment (Miyazaki 1971), in Japan.

Taiwanomyzus filicis (Miyazaki) (Fig.43l) Apterae are pale yellow with mainly pale appendages; BL 1.2-1.7 mm. Alatae have secondary rhinaria distributed III 30-39, IV 4-6, V 0(-1). On ferns (Athyrium, Dryopteris, Osmunda) in Japan. The life cycle is unknown; alate males were collected on Athyrium in late September (original description).

Taiwanomyzus himalayensis (Chakrabarti & Banerjee) Colour of apterae in life is unrecorded, presumably pale; BL 2.0-2.3 mm. Alatae have secondary rhinaria distributed III 2-6, IV 0, V 0. On ferns (Asplenium, Polypodium) in Uttar Pradesh, north-west India (original description, as Utamphorophora).

Taiwanomyzus montanus (Takahashi) Plate 24f (= T. darjeelingensis M.R. Ghosh, R.C. Basu & Raychaudhuri) Apterae are yellow with extensive black dorsal markings, and black antennae, siphunculi and cauda; BL 1.8-2.4 mm. On Saxifragaceae (Astilbe, Saxifraga) and also (in Japan) on Hydrangeaceae (Hydrangea, Schizophragma). In Taiwan, Japan, Korea, Philippines (Calilung 1967, as Aulacorthum) and West Bengal, India (M.R. Ghosh et al., 1977, as T. darjeelingensis).

Takecallis Matsumura

Calaphidinae: Therioaphidini

Seven species of narrow-bodied aphids on bamboos, resembling Myzocallis but with the clypeus bearing an anteriorly directed tubercle, and with the antennal terminal process about equal in length to the base of ANT VI. Usually all viviparae are alate. The genus is oriental in origin, but several species are now widely distributed. Higuchi (1968) revised the species then known, and accounts are available for Japan (Higuchi 1972), India (A.K. Ghosh & Quednau 1990), China (Qiao & Zhang 2004b), Korea (Y. Lee & S. Lee 2018), and for species introduced into Britain (Stroyan 1977). Quednau (2003) reviewed the genus, provided keys to species and illustrations of all available morphs. A specific parasitoid Trioxys remaudierei Starý & Rakhshani has been described that is associated with introduced Takecallis spp. in France and Spain (Rakhshani et al. 2017).

Takecallis affinis L.K. Ghosh Alatae are blackish, wax-covered in life; BL 2.1-2.5 mm. On undersides of leaves of Bambusa sp. and Arundinaria jaunsarensis (= Yushania anceps) in Himachal Pradesh, India. Apterous viviparae are unknown. Apterous oviparae (illustrated by Quednau 2003) and alate males occur in October (L.K. Ghosh 1986). B.C. Das & Chakrabarti (1989b) recorded a new parasitoid species, Trioxys takecallis, from T. himalayensis (= T. affinis) in Garwhal, western Himalaya. Quednau (2003) described a closely related form from Arundinaria sp. (or Yushania niitakayamensis?) in Taiwan with both apterous and alate viviparae as a subspecies, T. affinis niitakayamensis.

Takecallis alba Lee Alatae are pale yellow, covered in white wax; BL 2.1-2.5 mm. The wing veins have conspicuous black spots/triangles at their distal ends. On undersides of leaves of Pseudosasa sp. and Sasa spp. in South Korea (Y. Lee & S. Lee 2018). The life cycle is unknown.

Takecallis arundicolens (Clarke) Alatae are pale yellow or greyish-yellow, wax-dusted, usually without any dark dorsal abdominal markings, but with a black cauda and black-banded antennae (see influentialpoints.com/Gallery); BL 1.8-2.8 mm. [An unpigmented population with a pale cauda occurred at Wisley, UK, in 1967 (BMNH collection, leg. K.M. Harris).] On mature leaves of bamboos (Arundinaria, Bambusa, Phyllostachys, Sasa) in China, Japan, Korea, eastern Russia and introduced into Europe, Africa (Algeria) and USA (California). DNA barcoding has revealed differences between specimens from European and Asian populations which might be indicative of there being more than one species under this name (Y. Lee et al. 2017). The life cycle in east Asia is unknown; probably it is mostly anholocyclic where introduced, although Leclant (1966) reported oviparae in southern France, and Quednau (2003) illustrated an ovipara collected in California. 2n = 18.

Takecallis arundinariae (Essig) (= takahashii Hsu; synonymy by Quednau 2003) Alatae are whitish, pale yellow or greyish-yellow, with paired black elongate spots on abdominal tergite 1-7 and a pale cauda (see influentialpoints.com/Gallery); BL 1.7-2.4 mm. An apterous morph has been described, but only from high altitudes in Taiwan. On undersides of mature leaves of bamboos (especially Arundinaria and Phyllostachys, but sometimes also Bambusa, Dendrocalamus, Pseudosasa and Sasa). In India, China, Taiwan, Korea, Japan and introduced to England, Netherlands (Piron 2009), Hungary (Basky & Neményi 2014), Madeira (Aguiar & Ilharco 1997), Australia (Valenzuela et al. 2010a), New Zealand, North America and Argentina. An ovipara collected on Sasa kurilensis in Korea in early November was illustrated by Quednau (2003); it is anholocyclic where introduced. Coffelt & Schultz (1990) studied its population dynamics in USA. 2n == 18.

Takecallis assumenta Qiao & Zhang Alatae are yellow, with greyish-blue antennae; BL 1.9-2.5 mm. On upper surfaces of leaves of Bambusa spp., causing yellowing. In Qinghai, China (Qiao & Zhang 2004b). Other morphs and life cycle unknown.

Takecallis sasae (Matsumura) Alatae are green; BL c. 1.8 mm. On leaves of Phyllostachys, Pleioblastus, Sasa and possibly also Bambusa, in Japan (Higuchi 1968). Tao (1964) had T. taiwanus in China and Taiwan according to Higuchi (1968). The life cycle is unknown.

Takecallis taiwana (Takahashi) Alatae are pale green with a pale cauda (see influentialpoints.com/Gallery) ; BL 1.4-2.3 mm. Immatures are yellow-green with longitudinal rows of small brown spots. In young, still unrolled leaves and on new shoots of bamboos (Arundinaria and Phyllostachys, sometimes Bambusa, Sasa). In China, Taiwan, Japan and introduced to Europe, South Africa, New Zealand, USA, and more recently to South America (Argentina, Delfino 2001; Chile, Gonzales et al. 2000). Cottier’s (1953) account under T. arundinariae should be referred to this species. Quednau (2003) illustrated an ovipara collected in China in early December, but it is probably anholocyclic where introduced. 2n = 16.

Tamalia Baker

Tamaliinae

A relict nearctic genus with 4-5 species living without host alternation in galls on Arctostaphylos. A detailed diagnosis and review were provided by Remaudière & Stroyan (1984). Oviparae are alate. The male genitalia of an unidentified species of this genus were described and illustrated by Wieczorek (2011). Miller & Crespi (2003) and Miller et al. (2015) studied the molecular phylogeny of the genus and the evolution of inquilinism and host relationships. Their studies included several undescribed species including two nomina nuda collected on other genera of Ericaeae, T. morani from Arbutus arizonica (“Tamalia sp. A” in Miller et al. 2015) and T. cruzensis from Comarostaphylis diversifolia (“Tamalia sp. B”) in Miller et al. 2015).

Tamalia coweni (Cockerell) Plate 3h Apterae are dirty yellow to blackish, with variably-developed dark green cross-bands both dorsally and ventrally (see influentialpoints.com); BL 1.25-1.5 mm. In reddish leaf-galls on Arctostaphylos spp. in western North America, across boreal Canada to Ontario, and south to Mexico. Monoecious holocyclic, with alate oviparae and males produced from June to October. The peculiarities of the life cycle were discussed by Remaudière & Stroyan (1984), and studied further by Miller & Avilés (2000). T. keltoni Richards, described from Arctostaphylos sp. in Mexico, is doubtfully distinct. 2n=6.

Tamalia dicksoni Remaudière & Stroyan Colour of apterae in life is unknown; BL 1.4-2.1 mm. In leaf-galls on Arctostaphylos pringlei in California, USA (Miller & Crespi 2003). Presumably monoecious holocyclic; males were collected in late June (original description).

Tamalia inquilinus Miller Apterae are dark grey to brown or black; BL 1.4-1.7 mm. Living as an inquiline in leaf-galls of T. coweni on Arctostaphylos spp. (Miller & Sharkey 2000). On west coast of North America from north California to Baja California Norte, Mexico. Alate oviparae and males were collected in late July. Miller & Crespi (2003) compared the relative rates of mtDNA evolution of T. coweni and T. inquilinus, Miller (2004) studied the ecology of the relationship between the two species, and Miller et al. (2015) analysed their phylogenetic relationship.

Tamalia milleri Kanturski & Wieczorek Colour of apterae in life is unknown (abdomen has broad dark spinopleural cross-bands); BL 1.8-2.1 mm. On Arctostaphylos columbiana in western USA (California, Oregon). Apterous viviparae and alate oviparae were collected in August (Kanturski & Wieczorek 2015a).

Taoia Quednau

Calaphidinae: Calaphidini

Two oriental species on Betulaceae, possibly related to Euceraphis and Cepegillettea, characterized by the small rounded secondary rhinaria, ANT PT a little shorter than BASE VI and siphunculi with swollen bases and a single appended hair. Both apterous and alate viviparae occur. Quednau (1973) gave an account of the genus.

Taoia chuansiensis (Tao) Apterae and alatae are yellowish-green, the alatae having darker legs and antennae, and dark-tipped siphunculi; BL 2.9-3.1 mm. Living solitarily on undersides of leaves of Alnus spp. (cremastogyne, japonica) and Betula spp. in China (Tao 1964, as Euceraphis chuansiensis; Qiao et al. 2003). Sexual morphs and life cycle are unknown.

Taoia indica (Ghosh and Raychaudhuri) Apterae are pale green; BL 2.7-4.1 mm. Alatae have darker antennae and legs, and dark-tipped siphunculi. On Alnus nepalensis and Betula alnoides in India, Nepal (A.K. Ghosh & Quednau 1990), on Alnus nitida in Pakistan (Naumann-Etienne & Remaudière 1995), and on Alnus sp. in China (Qiao et al. 2003). There are many Indian records from other genera, presumably due to “casual visits”. Oviparae were collected in December and apterous (or alatoid) males in April, July and December (R.C. Basu & Raychaudhuri 1980).

Tauricaphis Mamontova

Aphidinae: Macrosiphini

One species from Ukraine related to Lipamyzodes but with dorsal abdominal hairs arising from tubercles and apterae with secondary rhinaria on ANT III.

Tauricaphis arabisi Mamontova Apterae are greenish with mainly yellowish brown appendages; BL c. 1.2 mm. On Arabis caucasica in Ukraine. Monoecious holocyclic; oviparae were collected in September (original description).

Tenuilongiaphis Zhang

Aphidinae: Macrosiphini

One species in China with elongate body, short 4- or 5-segmented antennae, long thick dorsal hairs and numerous short dorsal cylindrical processes.

Tenuilongiaphis stata Zhang & Zhong Apterae are pale green, with two longitudinal rows of darker green spots; BL 1.7-2.0 mm. Sedentary in habit, and only occurring in very small numbers, on Artemisia sp. in Gansu province, China. Believed to be heteroecious holocyclic, as a fundatrix(?) described from an unidentified shrub (G. Zhang 1999, as Sportaphis sporta; see G. Zhang & Qiao 2002), appears to be this species, and alatae and oviparae described as Ceruraphis hippophaetropis collected on Hippophae rhamonoides (G. Zhang 1999) seem likely to be be the autumn forms. Specimens with 4-segmented antennae found on Artemisia lavandulaefolia in Shanxi province, were described as a subspecies, Tenuilongiaphis stata ssp. shanxiensis (L. Zhang et al. 2008).

Tetraneura Quednau

Eriosomatinae: Eriosomatini

About 30 species in which the apterae are usually very globose and have one-segmented tarsi and the alatae have a simple, unbranched media in the forewing and usually ANT IV much shorter than ANT V. At least seven species have host alternation between leaf galls on Ulmus and roots of Poaceae, and the lesser-known species are described either from Ulmus or grass roots. The elm galls are invariably stalked, pouch-like outgrowths on upper sides of leaves. Tetraneura seems to be predominantly oriental and eastern palaearctic but T. ulmi appears to be native to northwest Europe, T. caerulescens and T. africana are mainly around the Mediterranean and in south-west Asia. T. fusiformis and T. nigriabdominalis are probably east Asian although populations of these species are now widespread. Hille Ris Lambers (1970b) revised the world fauna and there are more recent accounts from India (A.K. Ghosh 1984b), China (G. Zhang et al. 1991a, G. Zhang et al. 1999c), the UK (Blackman et al. 2019b), and the Iberian peninsula (Nieto Nafría et al. 2002a). Akimoto (1995) studied the interactions within the complex of six species found on Ulmus japonica, and Suzuki et al. (2009) compared amino acid concentrations in their galls. The genus is in need of further revision. The karyotype variation may be useful in this respect. (See also Blackman & Eastop 2000, pp. 348-351.

Tetraneura aequiunguis Zhang & Zhang Only the fundatrix is described, from an unidentified Ulmus sp. in China (Zhang et al. 1991).

Tetraneura africana van der Goot Apterae are buff‑coloured to pale brownish with a dark brown head and prothorax, often bluish due to waxy bloom, and with whitish wax secreted posteriorly; BL 2.0-3.3 mm. In root‑feeding colonies on Cynodon dactylon, often well below ground level, and also recorded from Sorghum halepense, Calamagrostis spp., Phragmites australis, and rarely on cereals. Recorded from Italy, Morocco, Egypt, Turkey, Ukraine, Iran, and east to Kazakhstan and Pakistan. A smaller aphid collected on the roots of Calamagrostis epigeios in Poland may also be this species (Hille Ris Lambers 1970b). Galls of T. africana on Ulmus have not been identified, although Mordvilko (1935) described sexuparae collected from Ulmus campestris in Italy. Populations on Cynodon roots are probably mainly or entirely anholocyclic.

Tetraneura asymmachia Zhang and Zhang Described from alatae (BL c.2mm) obtained from galls on Ulmus pumila and Ulmus sp. in June-July in China (Zhang et. al. 1991). Appears closely related to T. chinensis.

Tetraneura basui Hille Ris Lambers Colour of apterae in life is unrecorded; BL 1.5-1.8 mm. On roots of Poaceae (Capillipedium, Echinochloa, Eleusine, Eragrostis, Oryza, Pagonantherum, Paspalum, Pennisetum, Polypogon, Setaria) in north-east India (Raychaudhuri 1980). It is not known whether there is a gall generation on Ulmus.

Tetraneura brachytricha Zhang & Zhang Colour of apterae in life is not recorded; BL c.2.7 mm. On roots of Phragmites communis in Liaoning Province, China. Presumed to be heteroecious holocyclic; alate sexuparae collected on Ulmus in Xinjiang Province were assigned to this species, but the life cycle needs conformation.

Tetraneura caerulescens Passerini Galls are stalked, hazelnut- or walnut- sized (fig. 134G), red when mature, on Ulmus spp. (glabra, laevis, procera, minor, montana) in southern and south-eastern Europe, north Africa and south-west and central Asia. Alatae (BL 1.5-2.2 mm) emerge in late May-July and migrate to found colonies on roots of Poaceae (Alopecurus, Cynodon, Echinochloa, Eragrostis, Festuca, Milium, Poa, Setaria). Apterae on grass roots are orange-brown to brown, secreting bluish flocculent wax; BL 1.5-2.2 mm. Records of secondary host populations in China (Zhang et al. 1985, 1991a) need further confirmation.

Tetraneura capitata Zhang & Zhang Apterae are pale reddish brown; BL c.2.2 mm. On roots of Sorghum vulgare (= bicolor) in Hebei Province, China. A population on Agropyron cristatum in Gansu Province was described as a subspecies, T. capitata ssp. agropyricena Zhang (in Zhang & Qiao 1997b).

Tetraneura chinensis Mordvilko Galls are apparently rather large, sac-like, broader than high, green and slightly hairy (fig. 134F). Recorded from Ulmus sp. in China (Mordvilko 1935, as T. hirsuta) and U. pumila in Mongolia (Holman & Szelegiewicz 1974a). Alatae (BL 2.2-2.6 mm) were obtained from galls in May-June. Secondary host populations on roots of Poaceae have been identified by comparison of COI sequence (H. Zhang et al. 2008).

Tetraneura fusiformis Matsumura (= T. nigriabdominalis of many authors, incl. Hille Ris Lambers 1970b; = T. hirsuta Baker) Apterae are greenish‑ or brownish‑white; BL 1.5-2.5 mm. In colonies on roots of many genera and species of Poaceae, (Agropyron, Axonopus, Cenchrus, Chloris, Cynodon, Dactyloctenium, Echinochloa, Eleusine, Eragrostis, Oryza, Panicum, Paspalum, Pennisetum, Saccharum, Setaria, Sorghum), their presence often indicated by a reddish‑purple discoloration of the leaves. In Middle East, Africa, India, Nepal, Bangladesh, Pakistan, Sri Lanka, Andaman Isles, Thailand, Japan, Korea, Indonesia, Malaysia, the Philippines, New Britain, Australia, New Caledonia, Fiji, Tonga, Brazil, Argentina, Columbia and Central America. Heteroecious holocyclic in Japan, with a sexual phase and galls on Ulmus japonica; the galls are upright spindle-shaped, up to 12mm in height, petiolate at base, with numerous granular protuberances and short white silky hairs, usually crimson when mature. Alatae (BL 1.7-2.3 mm) emerge in June-July through a lateral opening.

T. fusiformis is probably entirely anholocyclic outside the range of Ulmus japonica, which seems to be its only primary host; its distribution suggests that parthenogenetic overwintering is only possible in tropical and subtropical conditions. The taxonomic confusion involving the application of the name T. nigriabdominalis was discussed by Eastop & Blackman (2005). All African records of T. nigriabdominalis as well as many of those from Asia (Blackman & Eastop 2000) should be referred to T. fusiformis, although past pest outbreaks on upland rice in Japan, and perhaps elsewhere in east Asia, were of the true nigriabdominalis (S. Akimoto, pers. comm.). Hille Ris Lambers (1970b, as T. nigriabdominalis) distinguished several root-feeding populations on the basis of abdominal chaetotaxy, and erected a subspecies, bispina, for his material from Africa and North America. However, most of the African material in the BMNH collection does not conform to bispina, and probably there are several distinct clonal populations distributed around the world. Raychaudhuri (1980) distinguished two populations in north-east India. 2n = 18 (gall generation); 2n = 17, 18, 19 and 20 (permanently parthenogenetic populations).

Tetraneura indica L.K. Ghosh Apterae are yellow; BL 1.9-2.3 mm. Alatae are unknown. In ant-attended colonies on roots of Calamagrostis pilosula in Himachal Pradesh, north-west India. According to Chakrabarti (2001) this species produces leaf-fold galls on an Ulmus sp., but the primary host generations have apparently not been described.

Tetraneura javensis van der Goot Apterae are whitish to yellowish‑white (see aphids of Karnataka website); BL 1.5-2.5 mm. On roots of Poaceae, especially sugar cane (Saccharum officinale), and also recorded from Capillipedium, Echinochloa, Eleusine, Neyraudia, Oryza, Panicum, Setaria and Sorghum. Anholocyclic populations of T. javensis occur on grass roots in Pakistan, India, Japan, Java and New Guinea, and it is a pest of sugar-cane in India and through east and south-east Asia (Blackman & Eastop 2006, p.1310). Probably T. javensis is heteroecious holocyclic in Pakistan, where there are galls believed to be of this species on Ulmus wallichiana (Hille Ris Lambers 1970b, Naumann-Etienne & Remaudière 1995). Alatae from these galls had BL 1.8-2.0 mm and secondary rhinaria distributed ANT III 11-14, IV 3-5, V 4-7. The morphology of embryos inside these alatae corresponded well with that of T. javensis on grass roots (Hille Ris Lambers 1970b), but the life cycle needs to be confirmed by host transfers. Setokuchi (1993) studied populations on sugar cane in Japan. We have not attempted to include in the key four other species described from India that were differentiated from T. javensis mainly on the basis of dorsal hair chaetotaxy, as this can show much intraspecific variation; Tetraneura kalimpongensis Raychaudhuri, Pal & M.R. Ghosh from roots of Saccharum and Pennisetum in West Bengal, Tetraneura lambersi Chakrabarti & Maity on an unidentified grass in Uttar Pradesh, Tetraneura multisetosa Raychaudhuri, Pal & M.R. Ghosh from roots of Capillipedium and Imperata in West Bengal, and Tetraneura utpali Chakrabarti, Maity & Bhattacharya from roots of Apluda mutica in Uttar Pradesh. The group is clearly in need of further study.

Tetraneura nigriabdominalis (Sasaki) (= T. akinire) Plate 5f Galls are stalked, hairy, elongate, spindle- or pouch-shaped, 15-40 mm high, usually with a pointed apex (fig. 134E), usually bicoloured green and rose-red when mature (see influentialpoints.com). They occur on various Ulmus spp. (canescens, japonica, parvifolia, procera, pumila) in southern and south-east Europe (including Malta – Mifsud et al. 2009), south-west Asia (Iran, Georgia), Kazakhstan (Kadyrbekov 2014e), China, Korea and Japan. It is also in eastern USA on U. minor (Stoetzel 1987, as akinire), and in western USA (California) on U. pumila and hybrid elms (S. Dreistadt, pers.comm.). Galls on Ulmus sp. on high ground in Pakistan (Naumann-Etienne & Remaudière 1995, as akinire) are probably also this species. Alatae (BL 1.4-2.3 mm) (fig. 117E) leave the gall through lateral slits in May-July to found ant-attended colonies on the roots of Poaceae (including Cynodon, Digitaria, Echinochloa, Oryza, Saccharum, Setaria). Apterae on grass roots are greenish‑ or brownish‑white; BL 1.5-2.5 mm. Their presence on the roots of some hosts is indicated by a reddish-purple discoloration of the leaves.

This species is known particularly as a pest of rice (Blackman & Eastop 2000), but many of the references to nigriabdominalis, e.g. in Africa, the Middle East, India and south-east Asia, are referable to T. fusiformis. W. Lee et al. (2012b) distinguished Japanese populations of T. nigriabdominalis (referred to as “Type A” in their paper) from those of T. fusiformis (“Type B”) by their differing mitochondrial COI sequences. Tanaka (1961) studied its host preferences, although he may have had a mixture of T. nigriabdominalis and T. fusiformis. Galli & Bonvicini-Pagliai (1998) studied its relations with ants. Enoki (1986) studied resistance to this aphid in U. japonica. The male genitalia were described and illustrated by Wieczorek (2012). Hille Ris Lambers’ (1970b) descriptions of gall generations under this name (his ‘Group I material’) should be referred to T. fusiformis (Eastop & Blackman 2005). Populations in China were described as a subspecies, T. nigriabdominalis ssp. shanxiensis Zhang & Zhang (as T. akinire ssp.). 2n = 18 (in gall generations); however Galli & Manicardi (1998) reported karyotype variation from 2n =13 to 19.

Tetraneura paiki Hille Ris Lambers Described from a single alate sexupara trapped in Korea (Hille Ris Lambers 1970b). Moritsu’s (1983) account and illustrations of an aphid under this name in Japan should be referred to T. nigriabdominalis (S. Akimoto, pers. comm.).

Tetraneura polychaeta Hille Ris Lambers Galls are conical, bag-like (Chakrabarti et al. 1985), on Ulmus laevigata (= villosa) in Pakistan. Alatae (BL 2.0-3.2 mm) were collected from galls in April, but probably emerge later in spring. A sexuparae trapped in Korea was also tentatively assigned to this species (Hille Ris Lambers 1970b). Secondary host generations are unknown.

Tetraneura polychorema Zhang Colour of apterae in life is not recorded; BL c. 2.1 mm. On roots of an undetermined grass species in Gansu Province, China (Zhang & Qiao 1997).

Tetraneura pumilae Lee, Seo & Hwang Galls (undescribed) are formed on Ulmus pumila. Emigrant alatae are grey, BL 1.6-1.8 mm, emerging in early June (Lee et al. 1993). The secondary host is unknown. T. iriensis Lee, Seo & Hwang was described from an unidentified Ulmus sp. in the same paper, with emigrant alatae of BL 1.2 mm, but the differences from T. pumilae are perhaps attributable to smaller body size.

Tetraneura radicicola Strand Stalked pouch galls on Ulmus japonica in Japan are similar to those of T. nigriabdominalis, but hairless or with only sparse hairs (see http://www.agr.hokudai.ac.jp/systent/akimoto/gall/T.%20radic%20gall.jpeg). Alatae (BL 1.7-2.5 mm) migrate in June-July to found colonies on roots of Poaceae. Presumed anholocyclic populations of this species on grass roots are recorded from Australia, Malaya, Philippines, Korea, Taiwan, India, Nepal and Sri Lanka, although on its secondary hosts T. radicicola cannot be reliably distinguished from T. yezoensis (q.v.). Tong et al. (2021) studied a mermithid nematode in Japan that parasitises alate sexuparae. 2n = 14. [Tetraneura persicina Zhang & Zhang, described from alatae (BL c.2.6mm) from galls on Ulmus pumila and Ulmus sp. in China (Zhang et al. 1991, Chen & Qiao 2012), appears very similar to radicicola, but the alatae have fewer dorsal cephalic hairs and their embryos have fewer spinopleural hairs. See also T. triangula.]

Tetraneura sorini Hille Ris Lambers Galls are stalked, hairy, pouch-shaped, often somewhat globular, yellow and red (Moritsu 1983 and http://www.agr.hokudai.ac.jp/systent/akimoto/gall/T.%20sorini%20gall.jpeg). On Ulmus spp. (laciniata, japonica, parvifolia) in Japan, Korea and China. Usurpation of galls of their own and other Tetraneura spp. by first instars was studied by Akimoto & Yamaguchi (1996). Alatae (BL 1.6-2.0 mm) emerge in June. Secondary host generations are unknown. 2n = 16.

Tetraneura triangula Zhang & Zhang Colour of apterae in life is not recorded; BL c. 2.5 mm. On roots of various Poaceae (Echinochloa, Imperata, Phragmites, Sorghum) in Liaoning, Shanxi and Shandong Provinces, China. Described from exules and sexuparae, which are similar to T. radicicola but much more hairy (W. Zhang & G. Zhang 1991). Galls and spring generations are unknown, but possibly this species occurs in Japan, on U. japonica (S. Akimoto, pers. comm.).

Tetraneura ulmi (L.) Elm‑Grass Root Aphid Galls are stalked, approximately bean-shaped (fig. 134D), smooth and shiny, reddish-green and/or yellow (see influentialpoints.com/Gallery), on leaves of various Ulmus spp. (campestris, glabra, minor). Their histology was investigated by Álvarez et al. (2013). Urban (2003a) made a detailed study of its bionomics on elms. T. ulmi occurs commonly in Europe, across Asia to eastern Siberia (Pashchenko 1988a) and northern Japan, and was introduced to North America. Alatae (BL 1.8-2.6 mm) (fig. 117E) emerge in June-July to colonize roots of Poaceae. Apterae on grass roots (Plate 1d) are pale orange yellow, yellowish white or reddish, with head, prothorax, and appendages brown; the body lightly dusted with wax; BL 1.7-2.8 mm. They are found on numerous species of Poaceae (especially Agropyron, Bromus, Dactylis, Deschampsia, Festuca, Holcus, Hordeum, Lolium, Poa, Zea), are invariably ant‑attended, and often in ants’ nests. Overwintering occurs commonly on grass roots or in ants’ nests, root‑feeding colonies producing rather few sexuparae in autumn. Ivens et al. (2012a,b) used microsatellite DNA analysis to investigate clonal diversity of root-feeding populations in the Netherlands. Zwölfer (1957) made a detailed study of root-feeding populations, including comparison with the closely related and perhaps synonymous Tetraneura longisetosa (Dahl), collected from roots of Brachypodium, Deschampsia and Festuca in Germany. Tetraneura chui Zhang, described from Sorghum vulgare (= bicolor) in China, may also be a synonym. 2n (T. ulmi) =14 (gall generation); Galli & Manicardi reported 2n =14 to 16.

Tetraneura ulmicema Zhang Known only from alatae emerging from galls on Ulmus glabra in Ningxia, China; BL c. 2.3 mm (G. Zhang & Qiao 1997a). The galls are smooth and hairless (Chen & Qiao 2012).

Tetraneura yezoensis Matsumura Stalked pouch galls on Ulmus japonica and U. laciniata in Japan resemble those of T. radicicola (see http://www.agr.hokudai.ac.jp/systent/akimoto/gall/T.%20yezoensis.jpg). Alatae (BL 1.8- 2.1 mm) migrate in late June-July to colonize roots of various Poaceae (Echinochloa, Eleusine, Eragrostis, Imperata, Miscanthus, Oryza, Saccharum, Setaria, Triticum, etc.). Apterae on grass roots are pale yellowish, brownish or pinkish white, with brown head and prothorax. Anholocyclic populations on grass roots assigned to this species also occur in Australia (Carver & Basu 1961, as T. heterohirsuta; Hille Ris Lambers 1970b). However there are no really reliable morphological criteria for distinguishing adults of T. yezoensis on grass roots from those of T. radicicola, because the adult chaetotaxy of this group is extraordinarily variable. Hille Ris Lambers (1970b) noted that embryonic (and first instar) chaetotaxy is more stable within species, but embryonic hair counts are almost impossible unless the embryos are dissected out prior to mounting.

Akimoto (1990) studied local adaptation to host plant and possible host-race formation in T. yezoensis. Suzuki et al. (2009) found exceptionally high asparagine concentrations in its galls. 2n = 12 in Japan (cf. T. radicicola); Chen & Zhang (1985b) reported 2n == 18 from China, but probably had another species. [T. sikkimensis Raychaudhuri, Pal & M.R. Ghosh, described from a small sample on Eragrostis nigra in Sikkim, India, is distinguished only by characters of adult chaetotaxy and may be a synonym of T. yezoensis. Tetraneura changaica, described from sexuparae only in Mongolia (Szelegiewicz & Holman 1980), resembles T. yezoensis but has shorter R IV+V and different abdominal chaetotaxy.]

Thalictrophorus Zhang & Qiao

Aphidinae: Macrosiphini

One species in China on Thalictrum, perhaps related to Longicaudus but with a short cauda bearing only 5-6 hairs.

Thalictrophorus thalictrophilus Zhang, Qiao & Zhao Apterae are white in life, BL c.1.9 mm. On Thalictrum aquilegifolium in Shanxi Province, China (Qiao et al. 2000). Other morphs and life cycle are unknown.

Thecabius Koch

Eriosomatinae: Pemphigini

About 12 species closely related and very similar morphologically to Pemphigus, but with some biological differences, and retained here as a separate genus despite the reservations of Aoki (1975), with Parathecabius Börner as a subgenus. Spring generations form galls by folding the leaves of Populus. The fundatrix inhabits a small gall set apart from that housing its progeny. Known secondary hosts are Ranunculus, Lysimachia and Salix. Secondary hosts of several species are unknown. Accounts are available for North America (Smith 1974a), north-west Europe (Heie, 1980), the UK (Blackman et al. 2019b), Japan (Aoki 1975, as Pemphigus) and China (G. Zhang et al. 1995a, G. Zhang et al. 1999c; their subgenus Oothecabius Zhang is here synonymised with Epipemphigus).

[Note: The type species of Thelazocallis Zhang in Qiao & Zhang 2000c, T. ranunculicola, is evidently based on exules of a Thecabius sp.; Thelazocallis Zhang 2000 is therefore a new synonym of Thecabius Koch 1857.]

Thecabius affinis (Kaltenbach) Plate 6d, (fig. 3), (Fig.45a) Galls occur on Populus spp., in May-June, mainly on P. nigra and its varieties, and are formed by folding of young leaves in half along mid-ribs; the outer, upper side of the leaf becomes blistered and yellowish or reddish (see influentialpoints.com/Gallery). These are secondary galls containing only the alatiform progeny of the fundatrix, which inhabits a small gall of its own formed by folding the edge of a leaf (fig. 131I). Alatae (BL 2.2-3.1 mm, with green abdomen, fig. 86B) leave the gall in late June-July to found colonies at stem bases and on runners of Ranunculus spp. (fig. 3). Apterae on Ranunculus (Plate 2i) are dirty yellowish white or yellowish green with shiny wax-wool (see influentialpoints.com/Gallery); BL 2.1-2.9 mm. There may be rare occurrences on other Ranunculaceae, e.g. Caltha palustris (Barbagallo et al. 2011). Mordvilko (1935) gave an account of the life cycle and described all morphs. Sexuparae are produced from late August to fly back to poplars, but alate virginoparae are also produced that colonise other Ranunculus plants, and overwintering on buttercup roots is common. T. affinis occurs on a wide range of Populus spp. throughout Europe and across Asia to east Siberia, Japan and Korea, and also in North America (see below). According to Chakrabarti (1987, 2001), in northern India T. affinis induces leaf galls on Ranunculus hirtellus in spring, and migrates to form subterranean colonies on Geranium spp. in late summer and winter; the life cycle is still unclear, and warrants further investigation, as this form would appear to be functioning as a distinct biological species. Populations on east Asian Populus spp. are often treated as a separate species, T. orientalis Mordvilko, here regarded as a synonym. T. populisuctus Zhang & Zhong, described from sexuparae collected on P. cathayana in China (G. Zhang et al. 1995a) may also be a synonym. However, T. orientalis ssp. minensis Zhang, described from Ranunculus roots in China (G. Zhang et al. 1995a), differs from T. affinis by having 4 wax plates instead of 2 on abdominal tergite 7, and could be a distinct species. The karyotype of T. affinis is 2n=38 in both England and Japan.

[This chromosome number has been determined additionally for a sample from British Columbia, Canada, leg. C.-k. Chan, and COI barcode sequences have confirmed that T. affinis is present in North America (Skvarla et al. 2017). DNA data have also confirmed the existence of a separate North American species, Thecabius populiconduplifolius (Cowen), which has 2n=28 (Harper & Macdonald 1966). T. populiconduplifolius is virtually indistinguishable from T. affinis and has an identical biology, but with native poplars as primary hosts (Palmer 1952).]

Thecabius anemoni (Shinji) Apterae are dark brick-brown, with white wax secreted from posterior abdomen; BL unrecorded (“rather large”). On Anemone hupehensis (roots?) in Japan. The generic position is uncertain.

Thecabius (Parathecabius) auriculae (Murray) Apterae are white to dusky white or dirty brownish, secreting wax-wool from posterior abdomen; BL 1.3-1.5 mm. Living all-year-round on the roots of Primula spp. in England, where it can be a pest of cultivated auriculas. It has also been collected on Androsace sempervivoides and Lysimachia (= Glaux) maritima. This species is very similar to T. lysimachiae, (see Stroyan 1964a), but unpublished data from more recently collected specimens of apterae and alate sexuparae (the latter collected in March and presumably redundant) has revealed some consistent differences (see key to aphids on Lysimachia). 2n=16.

Thecabius beijingensis Zhang Alatae from galls on Populus cathayana in China are yellow in life, BL c. 2.8 mm. They appear very similar to emigrant alatae of T. affinis except for the slightly higher rhinarial count. However, the sausage-shaped mid-rib gall illustrated (G. Zhang et al. 1995a, fig. 2) looks more like that of an Epipemphigus, and is probably one previously vacated by a species of that genus.

Thecabius (Parathecabius) cerastii (Börner) Apterae with wax-wool; BL 1.4-1.6 mm. On underground stems and roots of Cerastium uniflorum in Austria. The life cycle is unknown.

Thecabius (Parathecabius) gravicornis (Patch) Galls are formed by folding leaves downward along the mid-rib to form a yellowish-green, swollen pocket which becomes filled with the developing emigrant alatae. The fundatrix inhabits a separate small pocket on the upper side of a leaf lamina (Palmer 1952). On Populus balsamifera in North America, widely distributed but apparently rather rare; also collected on P. angustifolia in Utah (BMNH collection, leg. R. Danielsson). Alatae (BL 1.7-2.8 mm, with olive-yellow abdomen) migrate in July-August, presumably to an unknown secondary host.

Thecabius (Parathecabius) latisensorius Hori Galls like those of T. gravicornis are found on Populus maximowiczii in Japan and Sakhalin, and on P. laurifolia and P. talassica in Russia (Altai region) and Kazakhstan (Kadyrbekov 2017a). Alatae (BL 3.0-3.5 mm, with dirty yellow abdomen, fig. 86A) migrate in late July-September, presumably to an unknown secondary host (Hori 1938a; Aoki 1975, as Pemphigus latisensorius). 2n=18 (Blackman 1986).

Thecabius (Parathecabius) lysimachiae Borner Galls containing developing emigrant alatae are formed by folding and convoluting the leaf of Populus nigra (fig. 131J). The fundatrix inhabits a separate oval pocket-gall on the upper side of the leaf lamina, as in other Parathecabius. Alatae (BL 2.3-2.6 mm, with dark, dirty greenish abdomen) migrate in late June-July to roots and runners of Lysimachia spp., especially L. nummularia. Other species, e.g. Lysimachia (= Glaux) maritima, may also be colonised. Apterae on Lysimachia are brownish or greyish green, with wax; BL 1.1-1.8 mm. Alate sexuparae return to poplars in September, but anholocyclic overwintering on Lysimachia is also common. In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007) and central Asia, and recently (2000) recorded from New York State, USA (Foottit et al. 2006). T. luppovae (Narzikulov), described from Populus densa (= P. talassica) in Tajikistan, may be a synonym. 2n=18.

Thecabius populimonilis (Riley) The galls on Populus spp. (angustifolia, balsamifera, fremontii, trichocarpa) are rows of ovoid, bead-like cells, pale green or with a reddish tinge, developed from upper side of leaf lamina on both sides of mid-rib (see influentialpoints.com). Each gall in spring contains one developing aptera or alata. The fundatrices inhabit similar, solitary galls. This species is apparently facultatively monoecious; Gillette (1913) suggested that the large alatae (BL 2.7-3.0mm) produced in individual galls in June-July might recolonize cottonwoods, but it seems more likely that they are emigrants and found colonies of apterous exules in wax on Salix roots such as those observed by Lange (1943) in California. Presumably alatae produced in these colonies in October are sexuparae and return to Populus. However, alate sexuparae (BL 1.7-2.0mm) are also produced in galls on Populus, up to 12 per gall, in August-October (Maxson & Knowlton 1929, Palmer 1952, Harper 1959a). A more recent study by Aoki et al. (1996b) has confirmed most of these observations; they found heteroecious and monoecious life cycles at different locations, but their data on sex ratios indicate that some gene flow is probably occurring, rather than that two separate species are involved. Pike et al. (2012) have also now confirmed by DNA barcoding that colonies on willow roots (probably Salix dodgeana) in Washington are T. populimonilis. This species is widely distributed in USA, western Canada and Mexico. [Pemphigus saliciradicis may be a synonym; see note under that species.]

Thelaxes Westwood

Thelaxinae

Four oak-feeding species, three in Europe and the Mediterranean region, and one in North America. Remaudiere (1983b) provided a key including characters of alatae as well as apterae, Ilharco (1966a) keyed first instar nymphs, and Wieczorek et al. (2013) provided key couplets to distinguish between the oviparae and males of T. dryophila and T. suberi.

Thelaxes californica (Davidson) Appearance in life is not recorded, but presumably much like T. dryophila; BL of aptera 1.2-2.0mm. On Quercus spp. in western North America, from British Columbia to Mexico. Records from Turkey (e.g. Görür et al. 2011a) should probably be applied to T. suberi. Monoecious holocyclic; sexual morphs aestivate in first instar (Davidson 1917, as ?dryophila), adult oviparae occur in September-October (Palmer 1952). 2n = 12*. [Note: there is an important error in the printed version of Key F to the aphids on Quercus in Blackman & Eastop (1994, pp. 430-431); the first proposition in couplet 5 should refer to T. californica, the second to T. suberi.]

Thelaxes dryophila (Schrank) Apterae are broadly oval, rather flattened, dark brownish-red to purplish-grey with a paler spinal stripe (see influentialpoints.com/Gallery); BL 1.1-2.3 mm. Alatae have dark marginal sclerites and dorsal cross-bands on posterior abdominal segments. Ant-attended colonies form at tips of shoots, extending onto stems, leaf petioles and along mid-ribs on undersides of leaves. On Quercus spp. in Europe, the Mediterranean region and south-west Asia. Monoecious holocyclic; wax-coated sexual morphs aestivate from June to September on undersides of leaves in forks of veins (Polaszek 1986), and under cynipoid marble galls. Lombard et al. (1988) studied composition of the honeydew. 2n = 8.

Thelaxes suberi (del Guercio) Plate 4a, b Apterae vary in colour from pale green to dark brown (see influential points/Gallery); BL 0.9-1.5 mm. Living abundantly on young shoots, leaves and developing acorns of numerous Quercus spp. (but not robur) in England, Wales (Baker 2009), southern Europe, Mediterranean region and southwest Asia. Also recorded from Castanea sativa (in Italy). Monoecious holocyclic, but the life cycle is unstudied; Polaszek (1986) collected presumed sexuparae on Q. cerris in England in November-December, and Wieczorek et al. (2013b) described single specimens of the ovipara and small apterous male collected in Turkey in early November. 2n = 8. [Note: there is an important error in Key F to the aphids on Quercus in Blackman & Eastop (1994, pp.430-431); the first proposition in couplet 5 should refer to T. californica, and the second to T. suberi.]

Thelaxes valtadorosi Remaudière Apterae are light brown, somewhat waxy; BL 1.0-1.9 mm. On Quercus spp. in Greece (Remaudiere 1983b), and also recorded from Turkey (Şenol et al. 2014a, in abstract only). The life cycle is unknown. 2n = 8.

Therioaphis Walker

Calaphidinae: Therioaphidini

About.30 maculate aphid species with a knobbed cauda living on Leguminosae/Fabaceae of the tribes Trifoliae, Loteae, Galaeagneae and Coronillae. Half the species are known only from south-east Europe and the Middle East, four extend into north-west Europe, and four or five extend across Russia, one as far as Japan. The world fauna was revised by Quednau (2003), who provided keys and illustrations of all available morphs. He raised Bicaudella to the status of a full genus, and introduced a new subgeneric classification that is followed here. Earlier accounts are available for Europe (Pintera 1957a), Hungary (Szelegiewicz 1969b), north-west Europe (Heie 1982), the Iberian peninsula (Nieto Nafrìa & Mier Durante 1998) and the Middle East (Remaudière 1989c).

Therioaphis (Pterocallidium) aizenbergi Ivanoskaya & Tomilova Apterae (and alatae) are yellowish, covered dorsally with numerous large dark spots; BL c.1.4-1.7 mm. On Caragana arborescens in Siberia and Mongolia. Monoecious holocyclic; all the morphs are included in the original description.

Therioaphis alatina Hille Ris Lambers & van den Bosch Apterae are pale dirty greenish yellow with dark brown markings; BL 1.3-2.2 mm. On Ononis spp. in France, Italy and Switzerland. Although all adult viviparae have wings, these are often small and non-functional (original description). Sexual morphs occur in October.

Therioaphis (Rhizoberlesia) arnaultae Remaudière Apterae are probably pale yellowish with variably developed segmental markings; BL c.1.7 mm. On Astragalus sp(p). in Iran, Turkey and Lebanon. Oviparae and alate males were present in November (Remaudière 1989c). Remaudière & Talhouk (2000) provided further information on morphology of apterae.

Therioaphis astragali (Dzhibladze) Apterae seem not to occur, alatae often being brachypterous; alatae are yellowish with black dorsal abdominal spots, and dark spots at ends of wing veins, BL 1.5-1.9 mm. On Astragalus caucasicus in Georgia, and also recorded from Astragalus sp(p). in Azerbaijan, Iran and Turkey. Oviparae and alate males occur in October (original description).

Therioaphis (Pterocallidium) azerbaidjanica Remaudière Apterae are probably yellowish, BL 1.3-1.7 mm. On an Astragalus sp. with long, pointed leaves in Iran (Tabriz). Alatae and sexual morphs are unknown.

Therioaphis beijingensis Zhang Apterae are unknown, alatae are probably yellow with four rows of dark dorsal abdominal spots (two spinal, two marginal), and dark siphunculi; BL c. 2.2 mm. On Caragana frutex in China. Apart from the pigmentation this species seems close to T. tenerum.

Therioaphis (Pterocallidium) bonjeaniae Hille Ris Lambers & van den Bosch (Fig.40g) Apt. whitish to yellowish; BL 1.4-1.8 mm. On undersides of leaflets of Lotus (Dorycnium) spp. (hirsutus, rectus) in France (Alpes Maritimes), Spain and Portugal. The life cycle is unknown; parthenogenetic reproduction continued into November (original description). Sousa Silva & Ilharco (2016) provided a redescription and morphometric data for aptera and alatae collected in Portugal.

Therioaphis (Rhizoberlesia) brachytricha Hille Ris Lambers & van den Bosch (Fig.40c) Apterae are very pale buff to pale yellowish with dusky spots; BL 1.3-2.0 mm. On upper sides of leaves of young growth of Lotus spp., especially L. corniculatus growing in warm, dry positions. It can also feed on Trifolium pratense (BMNH collection, leg. H. Szelegiewicz, and see original description). Widely distributed in Europe, east to Iran. Monoecious holocyclic with alate males.

Therioaphis cana Zhang, Chen, Qiao & Zhong Only alatae are described; pale grey and leaf green in life, BL unrecorded. On Medicago sativa in Xinjiang province, China (G. Zhang 1999). This species could not be included in the key to aphids on Medicago on the basis of the published description.

Therioaphis dorycnii (Pintera) (Fig.40a) Apterous viviparae do not occur; BL of al. c. 1.7-1.8 mm. On Lotus (Dorycnium) spp. in eastern Europe (Czech Republic, Hungary) and Iran.. A record from Ukraine (Mamontova-Solukha 1963) may be referable to Th. litoralis. Pintera (1957a) described the ovipara.

Therioaphis hungarica Szelegiewicz Apterae are yellowish to orange with dark brown spots; BL 1.4-1.9 mm. Described from a reddish-flowered Ononis sp. in Hungary, and later collected on Ononis spinosa in Czech Republic (BMNH collection, leg. J. Holman). It was placed as a subspecies of Th. ononidis in Quednau 2003, but reinstated as a full species by Eastop & Blackman (2005).

Therioaphis (Pterocallidium) kermanica Remaudière Apterae are yellowish with black dorsal spots, BL 1.5-2.0 mm. On Astragalus pseudosquarrosus (= squarrosus) in Iran.

Therioaphis (Pterocallidium) khayami Remaudière (Fig.16f) Apterae are yellowish with extensive dark dorsal markings, and also usually with some dark submarginal sclerites ventrally on abdomen; BL 1.3-1.6 mm. On Astragalus sp. in Iran, and Afghanistan (BMNH collection, leg. R. van den Bosch).

Therioaphis kundurensis Quednau Apterae are yellow to yellowish green; BL 1.4-1.9 mm. On undersides of leaves of Caragana spp., attended by ants (Quednau 2003). In east Siberia.

Therioaphis (Rhizoberlesia) langloisi Remaudière & Leclant Apterae are golden yellow with brown transverse bands; BL 1.4-1.65 mm. In small colonies on undersides of older leaves of Coronilla glauca in the south of France and Spain (Mediterranean region), where it is anholocyclic (original description).

Therioaphis litoralis Hille Ris Lambers & van den Bosch ( = Therioaphis hillerislambersi Szelegiewicz; synonymy by Quednau, 2003) (Fig.40e) Apterae are bright yellowish to ochreous yellow; BL 1.4-1.8 mm. On Lotus and related genera (Anthyllis, Dorycnium). In Europe; southern France, Hungary, Czech Republic, Yugoslavia, Greece, Ukraine (Mamontova-Solukha 1963, as dorycnii) and Iran. An ovipara was found in October (original description).

Therioaphis loti Hille Ris Lambers & van den Bosch (Fig.40b) Apterae viviparae do not occur; BL of alata 1.4-1.7 mm. On Lotus (incl. Dorycnium) spp. in Israel, Iran and Turkey.

Therioaphis (Pterocallidium) luteola (Börner) (Fig.53b) Apterae are pale yellow or whitish; BL 1.7-2.0 mm. On Trifolium pratense in west, north, central and eastern Europe. Monoecious holocyclic with alate males (Heie 1982).

Therioaphis natricis Hille Ris Lambers & van den Bosch Colour of apterae in life is unrecorded; BL 1.25-2.2 mm. On Ononis spp. (especially natrix) in the Middle East (Israel, Lebanon, Iran). 2n=16.

Therioaphis obscura Hille Ris Lambers & van den Bosch (Fig.40d) Apterae are banded grey with black spots; BL 1.3-1.8 mm. On undersides of leaves of Lotus and Dorycnium spp. in France, Spain and Italy. There are also records from Ononis spp. Oviparae and alate males occur in October-November (original description), but it is also partially anholocyclic (Leclant 1966).

Therioaphis ononidis (Kaltenbach) Apterae are yellowish to orange with stiff black hairs with tubercular bases arising from large brown spots (see influentialpoints.com/Gallery); BL 1.8-2.2 mm. On reddish-flowered Ononis spp. in Europe, the Middle East and Central Asia. Records from North America and Japan, and from other genera of Leguminosae, are likely to be misidentifications of T. trifolii (Quednau 2003). Oviparae and alate males occur in October. 2n=16.

Therioaphis pteromaculata Quednau Only known from alatae, colour in life unknown; BL 1.8-2.0 mm. On Astracantha adscendens in Iran (Quednau 2003).

Therioaphis (Rhizoberlesia) riehmi (Börner) Alatae are yellow or orange with dark abdominal markings, the alatoid nymphs being paler yellow with four rows of dark dorsal spots (see influentialpoints.com); BL 2.0-2.7 mm. Apterae are very rare, and have not been described, but one is illustrated by Quednau (2003). On undersides of leaves of Melilotus spp., the preferred hosts, but there are also records from species of Medicago, Trigonella and Trifolium. In Europe, Algeria (Laamari et al. 2013), Middle East, India (Chowdhuri et al. 1969), Kazakhstan (Kadyrbekov 2009a), China (1 alata, BMNH collection. leg. L.A. Mound); introduced to North America, where it is widespread, and South America (Argentina: Ortego et al. 2004; Chile: Nieto Nafría et al. 2018). Monoecious holocyclic with oviparae and alate males in September (Hille Ris Lambers & van den Bosch 1964). 2n=16.

Therioaphis (Pterocallidium) subalba Börner (Fig.53a) Apterae are pale yellow, whitish anteriorly; BL 1.7-2.0 mm. On Trifolium spp. in northern, central and eastern Europe, and Turkey (Şenol et al. 2014); there are also records from Medicago sativa and Dorycnium (= Lotus) sp., and from Melilotus suaveolens in Korea (Quednau 2003).

Therioaphis tenera (Aizenberg) Apterous viviparae seem not to occur. Alatae are yellowish, the dorsal abdomen with variably developed dusky spots, and the siphunculi are pale (see influentialpoints.com); BL 1.7-2.2 mm. On Caragana spp., living dispersed on undersides of leaflets, mostly on lower branches. In Europe (Finland, Germany, Hungary, Czech Republic, Russia, Ukraine), Central Asia (Kadyrbekov 2017a) and introduced to Quebec, Canada (Quednau 2003). The record from east Siberia is referable to Th. kundurensis. A short-haired population on C. frutex in Ukraine was distinguished as a subspecies, T. tenera ssp. frutex Bozhko, and a related, more pigmented form is described from China as T. beijingensis (q.v.). Oviparae and alate males were described from Finland (Heikinheimo 1997); fundatrices are present in June and oviparae in late August-September. 2n=6.

Therioaphis (Pterocallidium) trifolii (Monell) Yellow Clover Aphid, Spotted Alfalfa Aphid, Spotted Clover Aphid Plate 4c (Fig.16g, Fig.40f,h) Apterae are pale yellow, greenish white to almost white, rather shiny, with rows of small light- or dark-brown pigmented raised spots (see aphids of Karnataka website); BL 1.4-2.2 mm. On many plants of Leguminosae/Fabaceae in genera Astragalus, Lotus, Medicago, Melilotus, Onobrychis, Ononis and Trifolium. Originally palaearctic; Europe, North Africa, Middle East, Central Asia, India, Pakistan, Japan (Sorin 1990c), and China (Zhang et al. 1999, as T. cana). Introduced to North and South America, South Africa, and Australia. Recognisable forms with more specific host-plant relations occur within the species, and are particularly evident where separate introductions have occurred to countries outside Europe. Some of these are regarded as distinct forms or subspecies, including the Spotted Alfalfa Aphid, T. trifolii ssp. maculata (Buckton), introduced to North and South America, Australia and New Zealand (for further details see Blackman & Eastop 2000 and 2007). T. trifolii ssp. ventromaculata Müller occurs on Astragalus onobrychus in Austria, and is also recorded from Poland and Czech Republic (Remaudière 1989c). T. trifolii ssp. albae Bozhko was described from Melilotus albus in Ukraine, and seems close to the nominate species (Quednau 2003). Monoecious holocyclic, with alate males, in cold temperate climates of northern and central Europe and more northerly USA, anholocyclic in warmer regions. 2n=16 (incl. ssp. maculata).

Thoracaphis van der Goot

Hormaphidinae: Nipponaphidini

Most species originally described in Thoracaphis have been removed to other genera, leaving the type species (arboris van der Goot) and a few others, mostly little known, which can hardly be said to form a homogeneous group. One species (kashifoliae) placed in a separate genus (Xenothoracaphis) in Blackman & Eastop (1994), is transferred to Thoracaphis following Remaudière & Remaudière (1997). Probably they were associated in the past with Distylium as primary host, as are other Nipponaphidini, but now only known from parthenogenetic generations on what were the original secondary hosts (Quercus, Lindera).

Thoracaphis arboris van der Goot Apterae are aleyrodiform, flattened, broadly oval, bluish-black with wax frosting on sides of body only; BL 1.3-1.6 mm. Immatures are dull greyish-green with rather more wax. On undersides of leaves of Quercus sp(p). in Java. Alatae collected in May and August (Noordam 1991). The life cycle is unknown; presumably anholocyclic.

Thoracaphis flava Takahashi Apterae are aleyrodiform, flattened, broadly oval, only slightly sclerotized, pale yellowish-brown; BL c. 0.5 mm. On undersides of leaves of Quercus sp. in Malaya (Takahashi 1950), and recently reported from Turkey (Şenol et al. 2014a), although this requires further confirmation. Other morphs and life cycle are unknown; presumed to be anholocyclic. This species clearly does not really belong in Thoracaphis; possibly it requires a new genus.

Thoracaphis kashifoliae (Uye) Apterae black, strongly sclerotized, covered with powder; BL c. 1.5 mm. Alatae are black with blackish subcosta, pterostigma and veins in forewing. On upper sides of leaves of Quercus glauca and Q. acuta in Japan. Anholocyclic, with alatae appearing in November (Takahashi 1958b; as Xenothoracaphis kashiwae). Kurosu et al. (2016) reported the occurrence of “defensive nymphs”; first instars of the alate morph with fore- and mid-legs enlarged to a varying extent, and observed to attack lepidopteran larvae.

Thoracaphis kumaoni Chakrabarti & Debnath Apterae are aleyrodiform, flattened, light to dark brown; BL 1.4-1.5 mm. On Quercus incana in northwest Himalaya, India (Chakrabarti & Debnath 2011). Other morphs and biology are unknown.

Thoracaphis linderae Shinji Apterae are aleyrodiform, flattened, broadly oval, green or brown becoming sooty black with age; BL 1.5-2.0 mm. On twigs of Lindera spp. in Japan (Shinji 1941). Alatae were collected in May (Shinji 1926). The life cycle is uncertain; Shinji (1926) reported overwintering as eggs on Lindera, but this was not confirmed by Takahashi (1958a) in his redescription of the species.

Thoracaphis quercifoliae A.K. Ghosh Apterae are flattened, subcircular, brown to dark brown sclerotic; BL 1.4-1.5 mm. On ?leaves of Quercus dealbata in Meghalaya, India (A.K. Ghosh 1988). Alatae (with secondary rhinaria distributed ANT III 20-22, IV 15-17, V 7-10) were collected in late March. The life cycle is unknown. This species perhaps belongs in Nipponaphis. An aphid collected on a plant of Lauraceae in China (Fujian) and used in DNA studies (Huang et al. 2012) was identified as this species.

Thoracaphis sp. on Cinnamomum Apterae are aleyrodiform, flattened oval, reddish-brown with creamy-white markings on prosoma and blackish abdominal plate. On bark of stems of Cinnamomum japonicum in Japan (Moritsu 1983; as ?cinnamomiae Shinji). This appears to be an undescribed species.

Thripsaphis Gillette

Calaphidinae: Saltusaphidini

About eight long-bodied, sedge-feeding aphids related to Subsaltusaphis but with pointed dorsal hairs, and often with wax pores. Empodial hairs are spatulate. Apterae often have rhinaria on ANT III, and alatae usually have extensive dark dorsal cross bands, sometimes merging into a solid patch. Quednau (2010) reviewed the world fauna, with keys and illustrations of all available morphs; his review made substantial changes to the nomenclature, giving Allaphis Mordvilko full generic status (q.v.), making Neobacillaphis Huculak a subgenus of Thripsaphis, and sinking several names in synonymy. Heie (1982) reviewed the species in north-west Europe, and Nieto Nafría & Mier Durante (1998) reviewed those on the Iberian peninsula.

Thripsaphis ballii (Gillette) Plate 4i (Fig.19f) Apterae are pale greyish green with extensive darker cross-bands and marginal areas, lightly wax-dusted; BL 1.9-2.5 mm. Alatae have 6-8 secondary rhinaria on ANT III, and dorsal abdomen with broad dark transverse bars that are sometimes partially fused across tergites 3-5. Described from Carex nebraskensis in western USA, producing oviparae and apterous males on this plant in October-November (Palmer 1952). Similar aphids also occur on other Carex spp. in eastern USA and Canada, Europe, Pakistan, east Siberia and China. Quednau (2010) recognised four subspecies, describing one as new. Unfortunately none of them have a clearly defined specific host relationship or geographical distribution. The nominate subspecies T ballii ssp. ballii is recorded from North America and Pakistan. T. ballii ssp. caespitosae Ossiannilsson, described from C. cespitosa in Sweden as paler, dirty yellowish in life and producing its sexuales in September (original description), and also recorded from Finland (Albrecht 2010) and Quebec, Canada, is considered by Quednau to have a holarctic distribution on various Carex spp. T. ballii ssp. longisetis Richards was described from Canada, and populations in east Siberia (Quednau & Shaposhnikov 1988) with similarly long hairs on the head and abdominal tergite 8 are also regarded as this subspecies. However there is intergrading with the nominate subspecies (Quednau 2010), and these hairs show almost continuous length variation in BMNH specimens. The fourth subspecies, T. ballii ssp. pennsylvanica Quednau from Carex sp. in Pennsylvanica, also has longer hairs, but has apterae with extensive dorsal sclerotisation (Quednau 2010). 2n=8 (for ssp. pennsylvanica).

Thripsaphis (Larvaphis) brevicornis (Ossiannilsson) Apterae are very slender, probably yellowish with brown markings; BL 2.7-3.0 mm. Alatae are apparently unknown. In leaf sheaths of Carex elata and C. lasiocarpa, often under water. In northern Europe (Sweden, Finland, Germany). Oviparae and apterous males occur in September-October (original description and Ossiannilsson 1959). The very short, 5-segmented antenna of this species are distinctive, and warrant the separate subgenus.

Thripsaphis caricis (Mordvilko) (= Thripsaphis gelrica Hille Ris Lambers; synonymy by Quednau 2010) Apterae are dark greyish brown, clothed with bluish grey wax powder; BL 2.5-2.8 mm. Alatae have 25-31 rhinaria on ANT III, and dorsal abdomen with broad dark bars separated between tergites. On Carex spp. (rostrata, vesicaria) in northern and central Europe (Heie 1982, as caricicola). [Quednau (2010) pointed out that the name caricicola Mordvilko 1921 should be treated as a synonym of Allaphis producta (Gillette).]

Thripsaphis polytricha Quednau Colour in life unknown, BL of apterae. c.2.7 mm. Alatae are unknown. On Carex sp. in Idaho, USA (Quednau 2010).

Thripsaphis sensoriata Hille Ris Lambers Only known from one alata, collected away from its host plant (presumed to be a Carex sp.) in Greenland.

Thripsaphis (Neobacillaphis) striata (Bozhko) Apterae are orange with a pair of broad brown longitudinal pleural stripes; BL c.1.4 mm. On Carex distans in southern Russia (Ciscaucasia).

Thripsaphis (Neobacillaphis) szelegiewiczi Huculak Plate 4f Apterae are brown with darker pleural and marginal longitudinal bands; BL 2.2-2.6 mm. Alatae have 9-13 rhinaria on ANT III. On Carex spp. in northern Poland. A population on Carex vulpina in the Ukraine is regarded as a subspecies, T. (N.) szelegiewiczii ssp. carpaticae (Mamontova), although possibly only a seasonal variant of T.( N.) striata (Quednau 2010).

Thripsaphis unciniae Quednau Colour in life unknown, BL of aptera c.1.9-2.1 mm. Alatae have 4-8 secondary rhinaria on ANT III. On Uncinia sp. and Carex sp. in Chile.

Tiliaphis Takahashi

Calaphidinae: Panaphidini

Four oriental species very similar to Eucallipterus, but with larger, pale siphunculi and more extensive forewing pigmentation. Accounts are available for Japan (Higuchi 1972), China (Zhang & Zhong 1982c, Qiao & Zhang 2003a) and east Siberia (Pashchenko 1988a; Quednau & Shaposhnikov 1988). Quednau (2003) keyed the species and illustrated all available morphs.

Tiliaphis coreana Quednau All viviparae are alate, greenish-yellow to yellow with brown-black sides to head and prothorax and longitudinal stripes on mesothorax, becoming more evident in late summer; BL 2.5-3.5 mm. Forewings have a deeply pigmented costal margin and a black zig-zag line between distal ends of veins. Immatures are green with 4 rows of black spots. In rows along veins on undersides of leaves, or as nymphs on upper sides of leaves, of Tilia spp. in Korea, China and eastern Siberia. Sexual morphs occur in September (Paik 1972, as Tiliaphis japonica; Quednau 1979, Quednau & Shaposhnikov 1988). 2n = 38 (Chen & Zhang 1985b).

[A DNA sequence assigned to T. shinae (Genbank accession no. GU978821) should be reassigned to T. coreana (Y. Lee et al. 2017).]

Tiliaphis pseudoshinae Quednau All viviparae are alate, whitish with brown-black markings on body and wings as in coreana, but without longitudinal stripes on mesothorax and with clear base to forewing; BL 2.0-2.6 mm. On Tilia sp. in Korea and eastern Siberia. Sexual morphs occur in September (Quednau 1979,Quednau & Shaposhnikov 1988).

Tiliaphis shinae (Shinji) All viviparae are alate, pale yellow to greenish-yellow with dark brown markings on body and wings as in pseudoshinae; BL 1.7-2.4mm. On undersides of leaves of Tilia spp. in Japan, China, Korea and eastern Siberia. Sexual morphs occur in early October (Takahashi 1961b, Higuchi 1972). 2n = 14 (Shinji 1931).

[A DNA sequence assigned to T. shinae (Genbank accession no. GU978821) should be reassigned to T. coreana (Y. Lee et al. 2017).]

Tiliaphis shinjii Higuchi All viviparae are alate, colour in life not recorded, dorsal dark markings as in coreana but with paired dark spinopleural patches on abdominal tergites; BL 2.0-2.7 mm. On Tilia spp. in Japan, Korea and eastern Siberia. It was recently reported to occur in Turkey (Şenol et al. 2014b), but the identification is suspect because this report refers to apterous rather than alate aphids. Sexual morphs were found in September (Higuchi 1972, Quednau & Shaposhnikov 1988).

Tiliphagus Smith

Eriosomatinae: Pemphigini

One North American species, differing from Prociphilus in having alatae with many more secondary rhinaria on the antennae.

Tiliphagus lycoposugus Smith (fig. 115A) Large leaf-nest galls are produced in spring by clumping and cupping terminal leaves so as to resemble a large, poorly-formed head of lettuce, 10-30 cm in diameter. On Tilia americana in eastern USA (Maryland, North Carolina). Fundatrices inside galls are brown, globose, BL 4.1-5.3 mm. Heteroecious holocyclic; emigrant alatae are reddish-brown, BL 2.6-3.3 mm, leaving gall in June-August to found colonies on the tuberiferous roots of Lycopus virginicus. Apterae on Lycopus roots are white to pale tan or flesh-coloured, with wax; BL 2.6-2.8 mm. Return migration of sexuparae to trunks of Tilia occurs in October-November (Smith 1965).

Tinocallis Matsumura

Calaphidinae: Therioaphidini

About 20 mostly small species usually associated with Ulmaceae, although species have also been described from Lythraceae (Lagerstroemia, Duabanga), and from Corylus, Dalbergia and Sapindus. Woody Leguminosae/Fabaceae seem to be recorded quite regularly as alternative or ‘casual’ hosts. All viviparae are alate and usually have paired spinal and marginal tubercular processes. Some species have conspicuous black markings on dorsal body and/or forewings. Problems with the taxonomy of Tinocallis may arise from seasonal variation; alatae in summer and autumn are often more pigmented than in spring and may have fewer secondary rhinaria. For example, specimens of Tinocallis platani collected in May have 17-25 (usually 20-23) rhinaria on ANT III, whereas specimens collected from the same trees in July-October have 7-21 (mostly 11-17) rhinaria on III; (V.F. Eastop, unpublished data). Quednau (2001) reviewed the world fauna, and regional accounts are available for Britain (Stroyan 1977), Fennoscandia and Denmark (Heie 1982), Italy (Patti & Barbagallo 1998), Europe (Kanturski et al. 2018), India (Chakrabarti 1988, A.K. Ghosh & Quednau 1990), eastern Russia (Pashchenko 1988a), China (Qiao et al. 2005b), Korea (Lee & Lee 2017) and Japan (Higuchi 1972). Quednau (2003) further revised the genus, illustrating all available morphs, erecting two new subgenera (Orientinocallis) and (Eotinocallis), and restoring three taxa treated as subgenera in Blackman & Eastop (1994), Pseudochromaphis, Quednaucallis and Sarucallis, to full generic rank.

Tinocallis (Orientinocallis) dalbergicola Quednau Alatae are blackish in life (J.H. Martin, pers. comm.); BL 1.9-2.2 mm. Only known from Hong Kong, on Dalbergia hancei (Quednau 2001). The sexual morphs are unknown.

Tinocallis (Orientinocallis) distincta Ghosh, Ghosh & Raychaudhuri Alatae are pale (colour unrecorded), with dark antennae, tibiae, dorsal abdominal tubercles, siphunculi and cauda; BL 1.8-2.2 mm. Described from an unidentified leguminous plant in West Bengal (M.R. Ghosh et al. 1971); Chakrabarti (1988) lists other records from West Bengal, mostly from undetermined plants, but including one sample collected on Duabanga sonneratioides. Oviparae and alate males occur in December-January, but alate viviparae are also present during winter months, so apparently in West Bengal there is only a partial holocycle.

Tinocallis (Orientinocallis) himalayensis Ghosh, Ghosh & Raychaudhuri Alatae are greyish or greenish, somewhat waxy (J.H. Martin, pers. comm.); BL 1.2-2.0mm. Forewings usually have dark spots at ends of veins and the branches of the media are brown-bordered distally. Described fom an unidentified leguminous plant (A.K. Ghosh et al. 1971e); specimens were subsequently collected in India (West Bengal, Sikkim) on Peltophorum fleragineum (Leguminosae) and Duabanga sonneratioides (Chakrabarti 1988). Also found on the undersides of leaves of Lagerstroemia macrocarpa (= speciosa) in Thailand (Danielsson & Robinson 1978, as T. khonkaensis), and on Lagerstroemia sp(p). in east and south-east Asia (Bangladesh, Thailand, Malaya, Singapore) (BMNH collection, leg. various collectors). The life cycle is unstudied; alate viviparae and apterous oviparae occur together in December-January in both northern India and Thailand, indicating that there is only a partial holocycle. Males are unrecorded.

Tinocallis insularis (Takahashi) Alatae are greenish-yellow with black-banded antennae; BL c. 1.6-1.7 mm. On undersides of leaves of Sapindus mukorossi in Taiwan (Takahashi 1927a) and Hong Kong (Martin & Lau 2011), and on Sapindus sp. in Japan (Higuchi 1972). The sexual morphs are unknown. Also recorded from Pterocarya stenoptera (Tao 1964), Aesculus chinensis (Tao 1999) and Magnolia sp. in Meghalaya, India (A.K. Ghosh & Raychaudhuri 1972b, as T. magnoliae), but these may all be vagrants.

Tinocallis latifoliae Lee & Lee Alatae are very pale yellow with small paired black sppots on ABD TERG 3-7, and appendages mainly pale except for contrastingly black joints of ANT III-VI; BL 1.2-1.5 mm. On Zelkova serrata var. latifolia in mountain areas of South Korea (Lee & Lee 2017). Closely related to T. zelkowae, its status as a separate species being supported by DNA barcoding (Y. Lee et al. 2017).

Tinocallis mushensis (Takahashi) Alatae are pale yellow, with three longitudinal thin brown stripes on head and pronotum; BL c. 2 mm. On young leaves of Zelkova formosana in Taiwan (Takahashi 1925). The sexual morphs are unknown.

Tinocallis (Sappocallis) nevskyi Remaudiere, Quednau & Heie Alatae are pale yellow or whitish, with very little pigmentation in spring and summer populations, except for a black spot at the apex of the hind femur (see influentialpoints.com/Gallery); BL 1.4-2.1 mm. Head and mesonotal lobes may be dark in autumn populations. On Ulmus spp. in central and south-west Asia (Afghanistan, Iran, Turkey, Georgia, Kazakhstan, Tajikistan, Uzbekistan, Pakistan, northern India). In recent times it has invaded and dispersed extensively in Europe; England (Hopkins 1997), Wales (Baker 2009), Belgium, Netherlands, France, Denmark, Germany, Sweden, Finland (Albrecht 2010), Poland, Hungary, Czech Republic, Switzerland, Italy (Patti & Barbagallo 1998). It is also recorded from Zelkova carpinifolia (= crenata), although Remaudière et al. (1988) thought that the occurrence of sexual morphs on this plant in Iran was ‘accidental’. Oviparae and alate males occur in November. Remaudière et al. (1988) discussed differences from T. saltans, with which it has been confused by some authors.

Tinocallis (Sappocallis) nikkoensis Higuchi Alatae are pale yellow with pale or dusky antennae and legs; BL c. 1.5mm. On Corylus spp. (heterophylla, sieboldiana) in Japan (Higuchi 1972). Biology and sexual morphs are unknown. [Paik’s (1965) record of T. zelkowae on Corylus sieboldiana in Korea should perhaps be referred to this species. T. sapporoensis Higuchi, which Blackman & Eastop (1994) suggested to be a midsummer population of T. takachihoensis, was made a synonym of T. nikkoensis by Quednau (2001).]

Tinocallis (Eotinocallis) platani (Kaltenbach) Plate l0b Alatae are yellow to greenish-white with extensive black-brown markings on head, thorax, abdomen and forewings, and black siphunculi; BL 2.0-2.2 mm. On undersides of leaves of Ulmus spp., especially U. laevis. Throughout Europe, across Asia to eastern Siberia (Pashchenko 1988a) and China (Qiao et al. 2005b), and introduced into western North America, where it occurs on U. americana. Oviparae and alate males occur on U. laevis in Europe in October. Olkowski et al. (1982) reported biocontrol with an introduced parasitoid in California. Patti & Barbagallo (1998) provided a recent account with colour photographs.

Tinocallis (Sappocallis) saltans (Nevsky) (figs 116B,D) Alatae are yellow to orange-yellow, with brown head and thorax even in early summer, with a more extensive distal black patch on the hind femur and more forewing pigmentation than in nevskyi; BL 1.4-2.2 mm (Nevsky 1929b, Quednau 1979). On Ulmus spp. in Spain (Núñez-Pérez et al. 1991), Italy (Patti & Barbagallo 1998), eastern Europe (Hungary, Romania) and central and east Asia (eastern Iran, Tajikistan, Kazakhstan, Uzbekistan, Urals, Afghanistan, Pakistan, Siberia, Korea and China). Also recorded from Zelkova serrata in Hungary (Ripka et al. 1998). Fundatrices collected in April and oviparae and alate males collected in October were described by Núñez-Pérez & Mier Durante (1993). Introduced into North America (Halbert & Pike 1990), Argentina (La Rossa et al. 1990) and Bolivia (cited by Naumann-Etienne & Remaudière 1995). [Some accounts under this name from north-west Europe (e.g. Heie 1982) and south-west Asia (Richards, 1967) should be referred to T. nevskyi (Remaudière et al., 1988).] 2n = 16 (in China; Chen & Zhang 1985b).

Tinocallis sophorae Zhang Colour of alatae in life is unrecorded; BL c. 1.7 mm. Described from Sophora japonica and Zelkova schneideriana in Jiangsu, China (Zhang & Zhong 1980c).

Tinocallis suzhouensis Zhang Alatae are dusted all over with white powder; BL c.2.2cm. On Zelkova schneideriana in China (Zhang & Zhong 1980c). Sexual morphs and life cycle are unknown.

Tinocallis (Sappocallis) takachihoensis Higuchi (fig. 51B) Alatae are pale yellow-green with shiny black head and thorax, black distal section of hind femur and base of hind tibia and black markings on the wings (see influentialpoints.com/Gallery); BL 1.8-2.0 mm. Recorded from Ulmus spp. in Japan (Higuchi 1972), China (Tao 1999) and eastern Siberia (Pashchenko 1988a, as T. ussuriensis), and also from Hemiptelea davidii in China (Zhang & Zhong 1980c, as T. hemipteleae). Introduced to Europe and North Africa, where it is recorded from Ulmus spp. in southern France (Quednau & Shaposhnikov 1988), Germany, Netherlands (Piron 2013), Denmark, Czech Republic and Poland (Kanturski et al. 2018a), Serbia (Petrović-Obradović et al.2018), Italy (Patti & Barbagallo 1998), Andorra (Mier Durante & Pérez Hidalgo 2002), Spain and the Balearics (Pérez Hidalgo et al. 2013), Malta (Mifsud et al. 2009), Algeria (Laamari et al. 2013), Greece (Papapanagiotou et al. 2012) and Turkey (Görür et al. 2011), and also introduced to USA (Maryland, first record 1996; Foottit et al. 2006). In England it has regularly been intercepted since 1997 on bonsai Zelkova serrata and Ulmus spp. imported from China, but was more recently collected outdoors on Ulmus glabra in Berkshire (Döring 2007). Oviparae and alate males occur in Sicily from mid-October (Patti & Barbagallo 1998) and an ovipara collected in France was illustrated by Quednau (2003). [T. nevskyi ssp. lianchengensis Zhang & Qiao, described from Ulmus sp. in China (Qiao & Zhang 1998b) is a synonym (Quednau 2001).] 2n = 16*.

Tinocallis (Sappocallis) ulmicola (Matsumura) (fig. 116C) Alatae are greenish-white, with black head and thorax, black apices of hind femora and bases of hind tibiae, and three large black patches on each forewing; BL 1.4-1.6 mm. On the undersides of leaves of Ulmus spp. in Japan, China, Korea and eastern Siberia (Quednau & Shaposhnikov 1988). Life cycle and sexual morphs are apparently unrecorded. 2n = 16.

Tinocallis ulmifolii (Monell) Alatae are pale yellow to greenish, reddish green or dark grey-green with white-bordered dusky longitudinal streaks on head and pronotum; abdomen with rows of white specks and dusky spots around hair-bases (see influentialpoints.com and aphid trek); BL 1.5-2.0mm. On the undersides of leaves of Ulmus spp. (especially U. americana), widely distributed in North America. Apterous oviparae and alate males occur in September-October (Hottes & Frison 1931, Palmer 1952, as Myzocallis ulmifolii); two variants of the oviparous morph were illustrated by Quednau (2003).

Tinocallis ulmiparvifoliae Matsumura (fig. 116A) Alatae are pale bluish-green, developing paired longitudinal white wax stripes on head and pronotum, a single, spinal white stripe on pterothorax and white wax spots on dorsal abdomen (Moritsu 1983). Tips of abdominal tubercles and distal ends of forewing veins are frequently dark. BL 1.6-2.7 mm. On Ulmus parvifolia in Japan, Korea, China and Taiwan. Introduced to Australia (Zeck 1933, as Myzocallis viridis), USA (California, Florida; see Foottit et al. 2006), England (on bonsai U. parvifolia and Zelkova serrata; Stroyan 1977 and BMNH collection, leg. R. Hammon), Italy (Patti & Barbagallo 1998) and Spain (Hidalgo & Nieto Nafría 2005). Oviparae have been recorded from Korea (Paik 1972) and China (in November; illustrated by Quednau 2003) but males are so far unrecorded. Hales (2020) studied populations in Australia. 2n = 16*.

Tinocallis viridis (Takahashi) (fig. 119A) Alatae are green, with green eyes, banded antennae, forewings with dark spots on pterostigma and at distal ends of veins, dark abdominal tubercles and dusky brown siphunculi; BL 2.2-2.7 mm. On Zelkova formosana in Taiwan, and on Z. schneideriana in China (Zhang & Zhong 1980c, as T. allozelkowae). Life cycle and sexual morphs are unknown. [This species has been confused in the literature with ulmiparvifoliae.]

Tinocallis (Eotinocallis) zelkovae Dzhibladze (fig. 119B) Alatae have a dark brown head, thorax, siphuncui and cauda, yellowish-green abdomen with extensive dark brown paired patches or cross-bands and maculate forewings with extensive pigmentation around veins; BL 1.4-1.8 mm. Described from Zelkova carpinifolia in Georgia and also collected on Zelkova sp. in Iran (BMNH collection, leg. R. van den Bosch). Monoecious holocyclic, with apterous oviparae and alate males (Dzhibladze 1957).

Tinocallis zelkowae (Takahashi) (fig. 119C) Alatae are shining pale yellow or greenish-yellow, with pale eyes, banded antennae, a fine brown mid-dorsal line on head and pronotum, pale yellow legs and siphunculi and a pair of small brown dorsal spots on each of abdominal tergites 3-7; BL 1.4-1.8 mm. On undersides of leaves of Zelkova serrata and related species in Japan (Takahashi 1919b), Korea and China, and introduced to England on bonsai Zelkova (Stroyan 1979; as T. nirecola), and to California, USA (Kono 1983). DNA barcoding studies (Y. Lee et al. 2017) revealed that specimens from Z. serrata var. latifolia in mountain areas of Korea were another, cryptic, species, now described as T. latifoliae (q.v.). Records from Ulmus may mostly be misidentifications of other species, although this aphid does seem to be recorded frequently, as immature stages as well as adults, on other plants; especially Betulaceae (Alnus, Carpinus) and Leguminosae (Robinia, Glycine max and ‘cultivated beans’). Monoecious holocyclic in Japan, with alate males and apterous oviparae in October-November; Inouye (1968) gives an account of the biology and descriptions of all morphs. 2n = 12*.

Tinocalloides Basu

Calaphidinae: Panaphidini

One species on Prunus in northern India and Nepal. A.K. Ghosh & Quednau (1990) gave a full account.

Tinocalloides montanus Basu All viviparae are alate, with brown head and thorax, pale yellow abdomen with two widely-separated rows of black spino-pleural patches; BL 2.1-2.6 mm. On undersides of leaves of Prunus spp., especially P. cerasus, in northern India (West Bengal, Himachal Pradesh) and Nepal. Holocyclic, with apterous oviparae and alate males in December-January. Sexuales occur on P. persica as well as on P. cerasus (Agarwala et al. 1984). 2n = 18 (Kurl 1981).

Titanosiphon Nevsky

Aphidinae: Macrosiphini

A genus for 4-5 palaearctic species mostly on Artemisia, similar to Uroleucon but with very long, robust siphunculi that lack polygonal reticulation. Pashchenko (1988a) keyed the east Asian species, and Kadyrbekov (2002c) keyed those of the western palaearctic region. The synonymies given here are discussed by Eastop & Blackman (2005)

Titanosiphon artemisiae (Koch) Plate 28i Apterae are very dark green to black; BL 1.7-2.5 mm. On thin stems of Artemisia spp., esp. campestris. Widely distributed in continental Europe, and also reported to occur in Iran (Rezwani et al. 1994). This species has been confused with T. minkiewiczi, so that it is not possible to apply all locality records to the correct species. Hille Ris Lambers’ (1947a) account applies to a mixture of the two species. Probably it is monoecious holocyclic.

Titanosiphon chondrillae Kadyrbekov Apterae are pale brown, or with pale brown head and thorax and dark green abdomen, with dark antennae, legs, siphunculi and cauda; BL 2.1-2.5 mm. On stems and upper leaves of Chondrilla sp. in east Kazakhstan (Kadyrbekov 2002c), but with a record also from Saussurea salsa (Kadyrbekov 2017a).

Titanosiphon dracunculi Nevsky ( = Titanosiphon baichengense Zhang) Apterae have a shiny orange head and thorax, mainly green abdomen, and mainly black antennae, legs and siphunculi; BL 2.3-3.2 mm. On flower stems of Artemisia dracunculus, occurring in large colonies attended by ants (Nevsky 1928a). In south-west Siberia, Central and eastern Asia. Oviparae and apterous males occur in September-October (Narzikulov & Umarov 1969).

Titanosiphon minkiewiczi Judenko (= T. benoisti Balachowsky, = T. kazakhstanicum Kadyrbekov) Apterae are very dark green to blackish; BL 1.8-2.8 mm. Living singly or in small colonies on thin stems and undersides of leaves of Artemisia spp., esp. campestris, in Europe and Central Asia (recorded from France, Germany, Poland, Lithuania, Belarus, Kazakhstan). Monoecious holocyclic, with oviparae and apterous males on A. campestris in September-October.

Titanosiphon neoartemisiae (Takahashi) (= T. bellicosum Nevsky, = T. zaisanicum Kadyrbekov; Eastop & Blackman 2005) Apterae are pale green to pale brown or reddish, with distal segments of antennae dusky, legs basally pale with darker femoral apices, tibiae and tarsi, and siphunculi dark with paler reddish bases; BL 1.7-2.8 mm. (However, specimens in the BMNH collection have very variable pigmentation, with tibiae sometimes dark but often quite pale.) Forming large ant-attended colonies on stems of Artemisia and Seriphidium spp., in south-west, central and eastern Asia (Narzikulov & Umarov 1969, as T. bellicosum, Pashchenko 1988a). Oviparae and alate males occur on A. scoparia in October-November in Central Asia (Nevsky 1928a, as T. bellicosum), but in Taiwan it is anholocyclic on A. capillaris (Takahashi 1927). Narzikulov & Umarov (1969) described specimens from Artemisia tenuisecta with particularly long siphunculi (c.0.7 × BL) as a subspecies, T. neoartemisiae ssp. hissaricum (as T. bellicosum ssp. hissaricum). Specimens with similarly long siphunculi were collected from A. campestris in Spain (BMNH collection, leg. D. Hollis), and had apterae with 11-13 secondary rhinaria on ANT III. 2n=8 (for specimens on A. dracunculus in Iran, leg. V.F.Eastop).

Toxoptera Koch

Aphidinae: Aphidini

NOTE: The name Toxoptera has been applied for many years to species resembling Aphis but having a stridulatory apparatus consisting of ventrolateral ridges on the abdomen and peg-like hairs on the hind tibia (Eastop 1952). Phylogenetic studies using both molecular and morphometric methods have now shown that this genus is paraphyletic, that all the species probably belong within Aphis, and that the three best known species (aurantii, citricidus and odinae) are not closely related (Kim & Lee 2008, Kim et al. 2010, Wang & Qiao 2009). Blackman et al. (2011) transferred T. odinae to Aphis. We have followed Lagos et al. (2014) and Aphid Species File in placing Toxoptera as a subgenus of Aphis, although it should probably eventually be synonymised with Aphis, further work being needed to establish the nearest relatives of each of the five or six remaining species.

Trama von Heyden

Lachninae: Tramini

Medium-sized to rather large pale brown or whitish aphids living mostly on roots of Compositae, in association with ants. Many of the nominal species described from Europe and Asia are little-known and not clearly distinguishable from the type species (troglodytes). T. rara is the only species in North America (formerly known there as T. oculata), and was probably introduced from Europe on Taraxacum roots. Subgenus Neotrama differs from Trama in presence of siphunculi and compound eyes, but DNA studies (Normark 1999) show a close relationship. Eastop (1953), Heinze (1962), Heie (1995), Nieto Nafría et al. (2002a), Blackman et al. (2019b) and Szelegiewicz (1978) provided accounts of European species. The unusual extent of karyotype variation in Trama was studied by Blackman et al. (2000).

Trama (Neotrama) afghanica Narzikulov Apterae are matt white; BL c. 3.1 mm. On roots of Euphorbia sp. in Afghanistan. Other morphs are unknown.

Trama (Neotrama) antennata Mordvilko Colour of apterae in life is unknown, probably whitish; BL c. 3.4-3.7 mm. On roots of Rumex sp. in Tajikistan. It was redescribed from the original material by Szelegiewicz (1982).

Trama (Neotrama) baroni (Hille Ris Lambers) Appearance in life is unknown, BL 4.2-4.9 mm. Described from roots of a Carduus sp. in Malta, in a nest of the ant Camponotus barbaricus, and apterae have also been collected in Lebanon and Turkey (BMNH collection). Other morphs are unknown.

Trama (Neotrama) bigrarta Zhang, Chen, Zhong & Li, in Zhang Apterae are dark yellow; BL unrecorded. On Sonchus oleraceus in Gansu Province, China (Zhang, 1999). Perhaps it is a synonym of T. caudata (or T. maritima).

Trama (Neotrama) caudata (del Guercio) Apterae are whitish, pale yellow or brownish; BL 2.5-3.3 mm. On roots of plants in composite genera of tribe Lactuceae (Cichorium, Hypochaeris, Picris, Lactuca, Leontodon, Sonchus, Taraxacum). Transfers between many of these genera are successful (Eastop 1953), so some of the other names currently listed in Neotrama (e.g. formicella, narzykulovi) are probably synonyms. In Europe, and also now recorded from Turkey (Özdemir et al. 2005), Iran (see Momeni Shahraki et al. 2019) and Central Asia (Kadyrbekov 2017a). Anholocyclic, no sexual morphs have yet been found. 2n= 9-12 (see Blackman et al. 2000).

Trama centaureae Börner Apterae are grey-green; BL c. 3 mm. On roots of Centaurea spp. in northern and central Europe (Heinze 1962, Heie 1995). Apart from its colour in life the characters distinguishing T. centaureae from T. troglodytes (short-based mesothoracic furca and absence of a frontal suture) show intraspecific variation, and the separate identity of this species needs further confirmation (Eastop 1953).

Trama (Neotrama) euphorbiae Juchnevitch & Kan Apterae are pale green or white; BL c.2.5 mm. On roots of Euphorbia lamprocarpa in Kazakhstan.

Trama (Neotrama) kulinitschae Narzikulov Apterae are milky white; BL c.4.7 mm. On roots of Ferula sp. in Central Asia (southern Tajikistan).

Trama (Neotrama) maritima (Eastop) Apterae are whitish, pale yellow or brownish; BL 2.6-3.9 mm. In ant-attended colonies on roots of Picris echioides and Sonchus asper in coastal regions of Europe (including the Azores and Madeira); also in Central Asia, if Protrama pamirica, described from Cichorium in Tajikistan (Narzikulov 1963), and also recorded from Kazakhstan (Kadyrbekov 2018a), is a synonym. It is apparently anholocyclic. 2n=10-14 (see Blackman et al. 2000).

Trama (Neotrama) nigrarta Zhang, Chen, Zhong & Li, in Zhang Apterae are dark yellow. On Sonchus arvensis in Gansu, China (Zhang 1999). Perhaps this is a synonym of T. caudata (or T. maritima).

Trama orobanches Mamontova Apterae have BL c. 3mm. On roots of Orobanche sp. in Ukraine (Mamontova 2012). Described as having apterae similar to T. rara but with a secondary rhinarium often present on ANT III (as well as those on IV-V), shorter hairs and smaller eyes.

Trama (Neotrama) penecaeca Stroyan Apterae are probably dirty greyish-white with brown head, antennae, legs, siphuncular cones and sclerites of posterior segments; BL 3.8-5.0 mm. On roots of Helianthus tuberosus in India (Jammu, Kashmir). Possibly monoecious holocyclic; an apterous male was described by Verma (1969).

Trama rara Mordvilko Apterae are whitish to olive-brown (see influentialpoints/Gallery); BL 2.5-3.5 mm. In ant-attended colonies on roots of Taraxacum spp., and sometimes on certain other Compositae, e.g. Helianthus. In Europe, Iran (Rezwani 2001), Kazakhstan (Kadyrbekov 2014e), Siberia, Japan and North America. 2n=12-14.

Trama (Neotrama) taraxaci Shinji Apterae are pale; BL c.1.5 mm. On (roots of ?) Taraxacum officinale in Japan. Possibly these are small or immature T. caudata, although that species is only certainly known from Europe.

Trama troglodytes von Heyden Plate 3g Apterae are white or yellowish white (see influential points/Gallery), or grey (older adults); BL 2.5-3.9 mm. They occur in ant-attended colonies on the roots of numerous Compositae (incl. Achillea, Artemisia, Centaurea, Cichorium, Cirsium, Cynara, Helianthus, Lactuca, Lapsana, Sonchus, Taraxacum). In Europe, west Siberia, south-west and Central Asia, and Japan. Various names have been applied to populations on different host plants, but there is no evidence from host transfers, karyotype or DNA of any specific host associations. Long thought to be entirely anholocyclic, but DNA evidence (Normark, 1999) and the discovery of oviparae and blind, apterous males in a population in southern England (Blackman et al. 2001) shows that sexual recombination occurs sometimes (but probably rarely) in this species. 2n=14-23 (see Blackman et al. 2000).

Tramaforda Manheim

Eriosomatinae: Fordini

A genus for two species in the Middle East with emigrant alatae having 5 antennal segments and containing embryos with remarkably long hind legs. The sexuparae are also distinctive because of their large size, long rostrum and elongate hind tarsi. Barjadze et al. (2020) reported mitochondrial and microsatellite DNA differences between the two species.

Tramaforda koachi Barjadze, Foottit & Maw Forming flat galls on the margins of the leaflets of Pistacia atlantica, similar to those of T. wooli but without a serrated outer margin (see Fig.7 in Barjadze et al. 2020). The life cycle is similar to that of T. wooli, with migration of emigrant alatae in early October to an unknown secondary host. Only known from northern Israel and the Golan Heights (Barjadze et al. 2020) This is “Fordini sp. A” of Koach & Wool (1977) and Wool et al. (1994).

Tramaforda wooli Manheim Forming flat, green galls on Pistacia atlantica by folding the margin of a leaflet towards the upper surface. The outer edge of the gall is distinctly serrated (see Fig.7 in Barjadze et al. 2020). The number of aphids in each gall is remarkably small, at most only 25 (Manheim 2007). Emigrant alatae leave the gall in early October, flying to an unknown secondary host. Presumably there is a two-year cycle as in other Fordini, but the secondary host generations are undescribed, although they would be easily recognised by their long hind legs with elongate second tarsal segments. Sexuparae were collected on trunks of P. atlantica and P. palaestina in late May to early June (Wool et al. 1994, as “Fordini sp. B”). Only known from Israel, mostly in the south.

Tricaudatus Narzikulov

Aphidinae: Macrosiphini

One Asian species very like Liosomaphis, but with tubercular spinal processes on the last two abdominal segments, and the immatures have spinulose hind tibiae.

Tricaudatus polygoni (Narzikulov) Plate 16b (Fig.42q) Apterae are whitish, pale yellow or yellowish green; BL 1.6-1.7 mm. Alatae have a quadrate black dorsal abdominal patch. On undersides of leaves of Polygonum and Persicaria spp., or on leaves and twigs of Spiraea. Other recorded hosts (Berberis, Bidens, Prinsepia) require confirmation. In Central Asia, India (Himachal Pradesh) and Japan. Heteroecious holocyclic, with the sexual phase on Spiraea (Chakrabarti & Banerjee 1993a). 2n=8.

Trichaitophorus Takahashi

Chaitophorinae: Chaitophorini

Seven east Asian species, at least six of them associated with Acer. Related to Chaitophorus and Periphyllus, but apterae have fused head and pronotum, and long thick, hollow hairs around margin of body, and their siphunculi lack polygonal reticulation. DNA data is available for only one species (koyaensis) and indicates a close relationship with Periphyllus (Wieczorek et al. 2017b). Alatae where known are very different from apterae, with head and pronotum distinctly separated and dark dorsal abdominal sclerotisation. To judge from the work of Sugimoto (2013) on T. koyaensis (see below), there can be considerable seasonal intraspecific variation in the distribution and shape of the dorsal hairs in this genus. Chakrabarti & Mandal (1986) revised the genus and keyed the species then known, Sorin (2002) provided a key to species, and Qiao et al. (2004a) reviewed the species in China.

Trichaitophorus aceris Takahashi Apterae are dark green with the last two segments of the antennae dusky, tarsi dark and siphunculi black (Takahashi 1937a); BL 1.4-1.8 mm. On young leaves of Acer spp.; originally described from A. rubescens in Taiwan. Aphids identified as aceris have since been recorded from various Acer spp. in north-west India and Nepal (Chakrabarti & Mandal 1986). However, there must be some doubt about the identity of the Indian populations, as they are pale in life and have pale tarsi and siphunculi (A.K. Ghosh, 1980). Alatae of Indian populations differ greatly from apterae, having black-brown head and thorax, antennae and legs mainly dark, abdomen with a large black patch on abdominal tergites 4-6 and separate sclerites on other tergites, and Chaitophorus-like siphunculi with polygonal reticulation (Quednau & Chakrabarti 1976, as Periphyllus pusillus). Chakrabarti et al. (2019) distinguished alatae from an Acer sp. in Bhutan as a new species, T. acerifolius Chakrabarti, Das & Sarkar, but it needs to be confirmed that these are not the unknown alatae of another species. The populations identified as T. aceris in northwest India are monoecious holocyclic; apterous males and oviparae were described by Chakrabati & Mandal (1986), collected on Acer sp. in October-November.

Trichaitophorus aenigmatosus Pashchenko Apterae are pale green to green; BL c.1.05 mm. On Acer tegmentosum in eastern Siberia (Pashchenko 1988a). Other morphs and life cycle are unknown.

Trichaitophorus ginnalarus Qiao, Zhang & Zhang Colour of apterae in life is unrecorded; BL 1.3-1.6 mm. On Acer ginnale in Heilongjiang, China (Qiao et al. 2004a). Other morphs and life cycle are unknown.

Trichaitophorus japonicus Sorin Apterae are green, with black-brown tarsi and pale siphunculi; BL c. 1.6 mm. On Acer tschonskii at high elevations in Japan, feeding on upper sides of leaves along mid-rib and main veins (Sorin 1979b). Sugimoto (2020b) provided some further description.of the syntype apterae. Other morphs and life cycle are unknown.

Trichaitophorus kominecola Sugimoto Apterae are yellow in life; BL 0.9-1.8 mm. Monoccious holocyclic on Acer micranthum in Japan, colonising undersides of leaves. Fundatrices were collected (on twigs) in late April, and oviparae and apterous males were collected in late October. Sugimoto (2020b) described and illustrated all morphs, finding differences between summer and autumn forms of apterae and in the marginal hairs of first instar larvae produced by different morphs. This species appears to be specific to A. micranthum, often in montane regions.

Trichaitophorus koyaensis Takahashi (= Periphyllus montanus Sorin, = Trichaitophorus takahashii Sorin; synonymy by Sugimoto 2013) Apterae are yellow, yellowish green or brownish green with pale antennae, legs and siphunculi; BL 0.9-1.6 mm. On Acer rufinerve in Japan (Takahashi 1961b), and also recorded from A. crataegifolium and A capilllipes (Sugimoto 2013, 2020b). Alatae have black dorsal abdominal cross-bands. Sugimoto (2012) has provided detailed descriptions and photomicrographs of all morphs including fundatrices (collected in April), oviparae and apterous males (October to early December). There are two forms of apterous viviparae; the form occurring in late summer and autumn was described by Sorin (1979b) under the name Periphyllus montanus. There are also different forms of first instars (Sugimoto 2013), some of which have lamellate marginal hairs similar to those of aestivating nymphs of some species of Periphyllus (Sorin 2002, as T. takahashii).

Trichaitophorus recurvispinosus Hille Ris Lambers & Basu Apterae are flattened, pale glassy white with tips of antennae and legs dusky brown; BL 2.4-2.5 mm. Stated to live on undersides of rolled leaves of a plant that was unidentified at the time the description was published (Hille Ris Lambers & Basu 1966), but subsequently identified as Elaeocarpus sikkimensis (BMNH collection, leg. D. Hille Ris Lambers). In West Bengal, India. Other morphs and life cycle are unknown. [The host plant and leaf-rolling habit is unlikely for a chaitophorine and needs additional confirmation. Probably the true hosts are Acer spp., as listed by Chakrabarti & Mandal (1986) under T. aceris. However, the size, colour in life, length of antennae and shape of marginal hairs make it unlikely that the synonymy with T. aceris proposed by these authors is correct.]

Trichosiphonaphis Takahashi

Aphidinae: Macrosiphini

About 12 palaearctic, mainly east Asian Myzus-like species with hairs on the siphunculi, typically host alternating between Lonicera and Polygonum. In Trichosiphonaphis s.str. (3 spp.) the siphunculi have a distinct apical flange, whereas in subgenus Xenomyzus (9 spp.) the siphunculi are flangeless Remaudière et al. (1992b) provided a review of the genus and a key to species.

Trichosiphonaphis (Xenomyzus) alpestris (Hille Ris Lambers) Apterae are brown, BL 1.7-2.2 mm. In ant-attended colonies on twigs of Lonicera xylosteum in alpine Europe (Austria, France, Switzerland). Monoecious holocyclic, with oviparae and apterous males in September (original description).

Trichosiphonaphis (Xenomyzus) cornuta Miyazaki (Fig.42e) Apterae are yellow, BL 1.2-1.3 mm. Alatae have secondary rhinaria distributed III 70-73, IV 33-41, V 18-30. On Polygonum thunbergii in Japan.

Trichosiphonaphis (Xenomyzus) corticis (Aizenberg) (Fig.38k) Apterae are dirty greenish brown; BL 1.80-2.50 mm. In ant-attended colonies on twigs of Lonicera spp. in north-eastern and central Europe (Russia, Byelorussia, Latvia, Finland, Poland, Romania, Germany, Czech Republic, Austria, Switzerland), and also in east Siberia (Pashchenko 1988a). Stekolshchikov & Buga (2006b) provided descriptions of apterous males and oviparae collected on L. xylosteum in October.

Trichosiphonaphis gerberae A.K. Ghosh & Raychaudhuri Apterae are pale yellowish; BL 1.9-2.2 mm. Alatae have secondary rhinaria distributed III 30-32, IV 15-17, V 10-11. Described from Gerbera macrophylla, an unlikely host, and later records are from Polygonum or Persicaria spp. (Raychaudhuri 1980). The life cycle is unknown.

Trichosiphonaphis lonicerae (Uye) (Fig.38h,i) Apterae are shiny greenish black or blackish brown, with paler appendages; BL 2.0-2.2 mm. Alatae have secondary rhinaria distributed III 32-55, IV 14-27, V 4-12. On twigs of Lonicera spp. in Japan, Korea and possibly also in Hong Kong (Martin & Lau 2011). The life cycle is unclear; it is found on Lonicera from April to November (Remaudière et al. 1992b), but possibly there is at least a partial migration to Polygonum.

Trichosiphonaphis (Xenomyzus) polygoni (van der Goot) (Fig.42d,f) Apterae are yellow to dark green, sometimes almost black; BL 1.3-2.2 mm. Alatae have secondary rhinaria distributed III 20-41, IV 8-21, V 1-11. On Persicaria and Polygonum spp. in east Asia (east Siberia, Japan, China, Taiwan, India, Pakistan, Java), and introduced to Africa (Burundi, Kenya; Remaudière at al. 1994), USA (Smith & Denmark 1982), Central America (Panama, Remaudière at al. 1992; Costa Rica, Zamora Mejías et al. 2012), and Australia (Brumley 2020). Miyazaki (1971) found it on the roots of the host. The life cycle is unknown. 2n=12 (Chen & Zhang 1985a, as T. ishimikawae).

Trichosiphonaphis (Xenomyzus) polygonifoliae (Shinji) Plate 20c (Fig.38j, Fig.42g) Apterae are greenish brown; BL 1.8-3.0 mm. In colonies on twigs of Lonicera spp. in spring, migrating in June-July to roots of Persicaria and Polygonum spp. (Hori 1938b, as Aulacorthum lonicerae). Spring migrant alatae have secondary rhinaria distributed III 16-46, IV 0-16, V 0. Gynoparae (with many more secondary rhinaria distributed III 80-100, IV40-63, V 0-9) and males return to Lonicera in October. The ovipara was illustrated by Petrović-Obradović et al. (2010). Native to east Asia (Japan, China, Korea, east Siberia), and introduced to Europe (France – Remaudière et al. 1992b, Italy – Coceano & Petrović-Obradović 2006,, Serbia – Petrović-Obradović et al. 2010, UK – Martin 2000) and Turkey (Ozdemir et al. 2005). 2n=12.

Trichosiphonaphis polygoniformosana (Takahashi) Plate 20d (Fig.42i) Apterae are dull blackish brown to black, with dirty yellow siphunculi and cauda; BL 1.8-2.2 mm. On Polygonum and Persicaria spp. in China, Japan, Taiwan, Korea, Thailand and east Siberia. Possibly with host alternation to Lonicera, but the life cycle is unclear. The original description included apterae from L. japonica in Taiwan, and apterae from this host in Japan were also mentioned, but these specimens are not mentioned in later publications (e.g. Takahashi 1965c), and may have been T. lonicerae Uye. Specimens from Takahashi in the BMNH collection (leg. F.V. Theobald) labelled “Lonicera japonica, Japan” (no date) are another, very small, apparently undescribed species.

Trichosiphonaphis (Xenomyzus) tade (Shinji) (Fig.42h) Apterae are dark yellowish brown to muddy green with dark antennae and siphunculi; BL c. 1.5-1.6 mm. On stalks of Polygonum thunbergii in Japan (Miyazaki 1971, Moritsu 1983). 2n=12.

Tripartita Yeh

Hormaphidinae: Nipponaphidini

A genus for one species in Taiwan with alate migrants from galls on Distylium having 3- or 4-segmented antennae, the flagellum being undivided or indistinctly divided and bearing numerous secondary rhinaria.

Tripartita formosana Yeh Galls on Distylium racemosum in Taiwan are green, spindle-like, 9-19 mm long and 4-9 mm wide, arising from the upper sides of leaves, with openings on the undersides, and with several galls on a single leaf (Yeh & Ko 2017). Galls of this species were also found on Distyliopsis dunnii. Developing galls contained fundatrices from October to May, producing alatae (BL 1.5-2.3 mm) from February to June, which presumably migrated to an unknown secondary host. First instars believed to be sterile soldiers were also observed, walking on the surfaces of galls and leaves.

Tshernovaia Holman & Szelegiewicz

Aphidinae: Macrosiphini

Two Asian species related to Macrosiphoniella but with a characteristic caudal structure, and an apparent association with both Campanulaceae and Artemisia. Contrary to Lee et al. (2002c), this structure is a modification of the actual cauda, rather than a process on abdominal tergite 8; the protruding structure beneath it is an extension of the anal plate. It is possible that this peculiar cauda has some function in relation to ant attendance.

Tshernovaia adenophorae Holman & Szelegiewicz Plate 28c (Fig.7a) Apterae are dull black with a paler transverse bar anterior to base of cauda; BL 2.0-2.8 mm. In ant-attended colonies on stems and undersides of leaves of Adenophora spp., and also found on Artemisia annua (Lee et al. 2002c). In Russia (Transbaikalia), Mongolia and Korea..

Tshernovaia spirocaudicula Ivanoskaya Apterae are yellowish, slightly waxy, with black antennae, legs, siphunculi and cauda; BL c.2.2 mm. On Artemisia sp., and also collected from stems of “bluebells” (Campanulaceae). In Siberia.

Tsugaphis Takahashi

Hormaphidinae: Hormaphidini

One or two highly specialized conifer-feeding species. Its relationship to other Hormaphidinae is uncertain, but a cladistic analysis (Chen et al. 2014) indicates a sub-basal position in a clade with Hormaphidini. Only apterous viviparae are known.

Tsugaphis sorini Takahashi Apterae are pale greenish, including antennae and legs; BL c. 0.9 mm. On undersides of leaves of Tsuga sieboldii in Japan (Takahashi 1957). Other morphs, biology and life cycle are unknown.

Tsugaphis sorini ssp. piceicola Shaposhnikov & Gabrid (fig. 65B) Apterae are pale green, translucent, older specimens becoming dark brownish-green; BL c. 0.9 mm. On Picea spp., especially pungens, schrenkiana, living in yellow depressions on upper and undersides of needles, which eventually become twisted and brittle, and drop prematurely. In Kyrgyzstan. Anholocyclic, overwintering as hibernating apterae (Shaposhnikov & Gabrid 1987). Differences from sorini s. str. are very small, and the two could be synonyms.

Tubaphis Hille Ris Lambers

Aphidinae: Macrosiphini

Two palaearctic species associated with Ranunculaceae, resembling Myzus but with a distinctive constriction at the base of the cauda. Alatae are without a dark dorsal abdominal patch, and have numerous secondary rhinaria on ANT III, and also some in a row on IV or IV-V.

Tubaphis clematophila (Takahashi) (Fig. 23a,b; Fig.45e) Apterae are green, with apices of antennal segments brownish; BL c.1.8-2.0 mm. Alatae have secondary rhinaria distributed III 32-36, IV 11-14, V 2-9. On Clematis spp. and Ranunculus sp. in Japan, Korea, China (Su et al. 2011) and east Siberia, and it has also been collected on Clematis connata in Pakistan (Naumann-Etienne & Remaudière 1995). 2n=12.

Tubaphis ranunculina (Walker) Plate 17f (Fig.45f,g) Apterae are yellowish with pale appendages; BL 1.2-1.9 mm. Alatae have secondary rhinaria distributed III 23-38, IV 0-10, V 0-6. On undersides of leaves of Ranunculus spp. throughout Europe, and also recorded from western Siberia (Altai), India (West Bengal), east Siberia (Kamchatka) and Japan (for detailed distribution see Stekolshchikov & Buga 2014). Monoecious holocyclic, sexual morphs in October (Blackman 2010); detailed descriptions of the ovipara and alate male are provided by Stekolshchikov & Buga (2014).

Tuberaphis Takahashi

Hormaphidinae: Cerataphidini

About 14 species in east and south-east Asia, typically with host alternation between often elaborately-branched galls on Styrax, and secondary hosts in Loranthaceae or Santalaceae. It is one of three genera of Cerataphidini in which the typical intracellular aphid symbionts (Buchnera) have been replaced by extracellular yeast-like symbionts, the others being Cerataphis (except C. bambusifoliae) and Glyphinaphis (Fukatsu et al. 1994, Xu et al. 2018). Determinations of the life cycles and/or symbiont types of several species (e.g. Aoki & Kurosu 1993, Kurosu et al. 1994, Aoki et al. 1995, Stern et al. 1997) have resulted in transfers to this genus of species described from Styrax in the genus Astegopteryx, and from Loranthaceae, originally placed in Rappardiella. A cladistic study (Chen et al. 2014) placed Tuberaphis close to Cerataphis, and this is confirmed by molecular work (e.g. Xu et al. 2018). Noordam (1991) redescribed and keyed the species in Java (as Rappardiella), and Jiang et al. (2012a) and Qiao et al. (2018) reviewed and keyed the Chinese species. Fukatsu et al. (1994) compared symbionts and gall structure with other Cerataphidini. Aoki & Kurosu (2010) reviewed life cycles, gall formation and soldier behaviour.

Tuberaphis breviseta A.K. Ghosh (= Neotuberaphis indica of Blackman & Eastop 1994) Apterae are dark brown, BL 1.3-1.5 mm, living in white wax in marginal leaf-folds on Eugenia sp. in Nagaland, India. Alatae have a pale brown abdomen and dark wing veins (Raha & Raychaudhuri 1981, as Neotuberaphis indica). A.K. Ghosh (1988) provided a redescription of type specimens. The life cycle is unknown.

Tuberaphis cerina (Noordam) Apterae are yellowish white tinged with reddish brown, with a dull transparent coat of wax; BL 1.1-1.3 mm. On Dendrophthoë pentandra in Java, living pressed close to surfaces of rolled leaves, attended by ants.

Tuberaphis coreana Takahashi (= taiwana of Blackman & Eastop 1994) (Fig.60b) Galls are large (about 14cm in diameter), pale greenish-yellow and coral-like, with small circular distal openings, and arise from the stem of Styrax formosanum in Taiwan (fig. 124G). Alate emigrants (BL c.1.75 mm) were collected in August (Takahashi 1934b). The secondary hosts, assuming the synonymy with T. taiwana is correct, are Viscum spp., on which apterae of the presumed secondary host generations have been collected in Japan and Korea. The colour of these apterae in life is unrecorded, but presumably they have a wax fringe; BL 1.7-2.0 mm. The synonymy is fairly well-supported by molecular data (Stern et al. 1997) but secondary host populations are not known from Taiwan, and some further confirmation seems necessary (see Aoki & Kurosu 1993, as taiwana).

[Tao (1969, as Astegopteryx) synonymised this species with A. vandermeermohri (Hille Ris Lambers), but this is untenable because the gall described for that species is quite different, and there are also differences in the soldier morph and first instar exule (Aoki & Kurosu 1993).]

Tuberaphis cymigalla (Qiao & Zhang) Galls are green, branched, arising from twigs, with many openings, on Distylium racemosum in Fujian, China (Qiao & Zhang 1998a, as Rappardiella). Only a single aptera from the gall is described, and the host association is an unlikely one requiring further confirmation of the generic placement of the aphid, and its host.

Tuberaphis dendrotrophe Qiao, Jiang & Chen (in Jiang et al. 2012a) Apterae are broadly oval, brown, with margin of body covered with white wax; BL 1.5-1.8 mm. Alatae are undescribed. On Dendrotrophe frutescens in Hong Kong (Tuberaphis sp. in Martin & Lau 2011). Jiang et al. (2012a) provided descriptions of adult apterae and all larval instars (but most of the measurements in Table 1 of their paper are about 33% too high).

Tuberaphis leeuweni (Takahashi) Galls are formed from flower buds of Styrax subpaniculatus in Sumatra. They are slender, green, pod-like, downy, 5-8 cm long, with a distal opening (fig. 124D). Alatae, BL about 1.6 mm, were collected from them in June by Takahashi (1936, as Astegopteryx), and in September by Aoki et al. (1995) The September-collected alatae were found to be sexuparae, indicating an absence of host alternation. Soldier-like second instars occur in the galls (Aoki et al. 1995).

Tuberaphis loranthi (van der Goot) (Fig.39a-d) Apterae are orange-brown, with a narrow wax fringe, and a segmentally-divided dusting of mealy wax; BL 1.5-2.1 mm. On old leaves and in dense colonies on woody stems of mistletoes (Scurrula spp.; records from Loranthus should probably also be referred to this host genus). In Taiwan, India (A.K. Ghosh 1988), Java (Noordam 1991) and Papua-New Guinea (BMNH collection, leg. E.J. Brough). The life cycle is unknown.

Tuberaphis macrosoleni Noordam Apterae are brown, with antennae and legs paler, and with a dorsal layer of flaky wax, becoming thicker and woolly on margins; BL 1.3-1.6 mm. In rolled-leaf galls on Macrosolen cochinchinensis in Java (Noordam 1991).

Tuberaphis owadai Kurosu & Aoki Galls on Styrax tonkinensis in northern Vietnam are silvery grey, coral-like, 9-24 cm across, similar to those of T. coreana and T. sumatrana on other Styrax spp. (Kurosu & Aoki 2003). Colony sizes in the galls were estimated at 50,000 to 180,000 individuals. Alatae, BL 1.5-1.9 mm, produced in the galls in September, were found to be sexuparae, so the species lives on Styrax without host alternation. Numerous second instar sterile soldiers were present in the galls. Jiang et al. (2012a) provided a redescription including immature stages from galls on Styrax japonicus in Yunnan, China.

Tuberaphis scurrulae Noordam (Fig.39e) Apterae are velvety black or dull brownish black, with a flat narrow marginal wax fringe; BL 1.7-2.0 mm. On twigs of Scurrula sp. in Java (Noordam 1991).

Tuberaphis styraci (Matsumura) (= Hamiltonaphis styraci of Blackman & Eastop 1994) Large, coral-like galls are formed on the twigs of Styrax obassia in Japan. The galls take two years to develop (Aoki & Kurosu 1990, as Astegopteryx). Colonies in the galls may reach a size of 15,000 to 20,000 individuals, of which more than half may be sterile second instar soldiers. The soldiers not only defend the colony but also clean the gall, by pushing out honeydew droplets and exuviae (Aoki & Kurosu 1989, as Astegopteryx). Shibao et al. (2004, 2016) studied the controlling mechanism for soldier production, and Kutsukake et al. (2004) identified a soldier-specific venomous protease presumably used in colony defense. There is no host alternation. Alate sexuparae (BL c. 1.4 mm) appear in late July or August of the second year and deposit yellowish sexuales on the undersides of the Styrax leaves. Oviparae lay overwintering eggs probably in fissures in the bark (Aoki & Kurosu 2010).

Tuberaphis sumatrana (Hille Ris Lambers) Galls are coral-like, silvery grey, about 10 cm in diameter when fully developed, with small openings on distal lobes, arising from the stem via a short thick branching tube (fig. 124F). They are found on Styrax spp. (serrulatus var. mollissimus, subdenticulatum) in Sumatra. Alate emigrants, BL about 1.5-1.6 mm, were collected in February (Hille Ris Lambers 1931a, as Astegopteryx).

Tuberaphis takenouchii (Takahashi) The ‘broccoli-head-like’ gall of this species on Styrax japonicus (fig. 124I) was redescribed by Aoki & Usuba (1989; as Aleurodaphis), who found that it contained sterile second instar soldiers. Migration occurs to Loranthaceae, including Scurrula sp. This host alternation and synonymy was proposed by Kurosu et al. (1994) and supported by the molecular studies of Stern et al. (1997). For a description of the alate emigrant see Takahashi (1934a, as Astegopteryx). Apterae on Scurrula are dark grape-coloured, usually with little wax, BL 1.2-1.4 mm, and inhabit rolled leaves (Noordam 1991, as Rappardiella plicator). In Taiwan, Sulawesi and Java. To judge from a paratype slide in the BMNH collection, T. loranthicola A.K. Ghosh 1988, originally described from Taxillus sutchuenensis in China (Tseng & Tao 1938, as Astegopteryx loranthi) is probably a synonym.

Tuberaphis viscisucta Zhang Colour of apterae in life is unrecorded, not wax-powdered; BL c. 1.3 mm. On Viscum album in Yunnan, China, causing downward curling of edges of leaves. This is likely to be a synonym of T. takenouchii.

Tuberaphis xinglongensis (Zhang) Apterae were described from an unidentified plant in Hainan, China (G. Zhang & Zhong 1982b, as Astegopteryx). Alatae have numerous secondary rhinaria, distributed ANT III 33-39, IV 13-17, V BASE 7-10. This species was subsequently recorded from an unidentified species of palm, and then from Cocos nucifera. Its generic placement in Tuberaphis by Jiang et al. (2012a) was confirmed by Chen et al. (2014), although the host association is an anomalous one for a member of this genus.

Tuberculatus Mordvilko

Calaphidinae: Myzocallidini

About 60 species of oak- and chestnut-feeding aphids with one or more tubercular abdominal spinal processes. All viviparae and males are alate. The genus includes some well-defined subgenera of limited distribution (Tuberculoides in western palaearctic, Orientuberculoides, Nippocallis and Acanthocallis in east Asia, Pacificallis in western nearctic, and Toltecallis in Mexico). Richards (1968c) and Hille Ris Lambers (1974) reviewed the world fauna, and Quednau (1999) revised the subgeneric classification and provided a key to species and illustrations of all available morphs. Quednau (2003, pp. 294-6) keyed the currently known oviparae of four subgenera. Regional accounts are available for Britain (Stroyan 1977), north-west Europe (Heie 1982), Iberian peninsula (Nieto Nafría & Mier Durante 1998), India (Chakrabarti 1988; A.K. Ghosh & Quednau 1990), China (Qiao et al. 2005b), eastern Siberia (Pashchenko 1988a) and Japan (Higuchi 1970). W. Zhang & G. Zhang (1991) did a cladistic analysis of 23 species in China. Yao (2011) constructed phylogenetic trees based on mtDNA sequences for 20 species of Tuberculatus in Japan, including some undescribed species, and showed that wing-loading was greater in ant-attended species. Y. Lee et al. (2017) found variation in DNA barcoding sequences indicating the existence of cryptic species in several subgenera. Quednau (1992) keyed the species of subgenus Pacificallis, and Qiao & Zhang (2002a) reviewed Chinese species of the subgenus Nippocallis.

Tuberculatus (Acanthocallis) acuminatus Zhang, Zhang & Zhong Alatae are whitish with dark head and siphunculi; BL c. 1.3 mm. On Quercus acutissima in China (G. Zhang et al. 1990b). Sexual morphs unknown.

Tuberculatus (Tuberculoides) albosiphonatus Hille Ris Lambers Alatae are pale green, with black-banded antennae, a sharply-bordered dark brown spot on the pterostigma, and pale siphunculi; BL 1.1-1.7 mm (Hille Ris Lambers 1974a). On Quercus infectoria (incl. var. boissieri) in Iraq (Kurdistan) and Israel (BMNH collection, leg. V.F. Eastop). Sexual morphs are unknown.

Tuberculatus (Tuberculoides) annulatus (Hartig) Alatae are yellowish, greyish-green or pink to purple in summer, with black-banded antennae, black tarsi, and dark distal two thirds or more of siphunculi (see influentialpoints.com/Gallery); BL 1.7-2.2 mm. On undersides of leaves of Quercus spp., especially Q. robur, less commonly Q. petraea. Throughout Europe, Iran (Rezwani et al. 1994), Siberia (Pashchenko, 1988b), Kazakhstan (Kadyrbekov 2017a), north-west China (Qiao et al. 2005b) and introduced on European oaks to Australia, New Zealand, North and South America. Alate males and apterous oviparae occur in October (northern hemisphere). Rohitha & Penman (1984) studied development under laboratory conditions, Kennedy (1986b) studied the role of tarsi in host selection, Heimbach (1986) studied population dynamics in relation to honeydew production, and Dahlsten et al. (1990) reported specific parasitoids in Germany. [DNA barcode data now indicate that two species may be confused under the name T. annulatus, differing in colour, COI sequence and morphological features, and both present in both Europe and North America (Skvarla et al. 2017; Y. Lee et al. 2017).] 2n = 14.

Tuberculatus (Tuberculoides) borealis Krzywiec Alatae are pale blue-green, yellow mottled with green, or yellow (see influentialpoints.com/Gallery), with blackish-banded antennae and siphunculi only apically dark (rarely over more than distal half); BL 1.9-2.3 mm. On Quercus robur, more rarely on Q. petraea or robur × petraea hybrids, in Europe (UK, Belgium, Denmark, Norway, Sweden, Italy, Poland, western Russia, Turkey), east to Iran (BMNH collection, leg. S.H. Hodjat), and introduced to North America (British Columbia; Quednau 1999). Sexual morphs occur in October-November (Krzywiec 1971, Quednau 1999).

Tuberculatus (Pacificallis) californicus (Baker) Alatae are pale yellowish-green (Baker 1917a) with banded antennae and brown-black spots at bases of all tibiae; BL 1.7-2.5 mm. On Quercus lobata in western USA (California, Colorado, Oregon, Utah). Sexual morphs occur in November (BMNH collection, leg. R. van den Bosch).

Tuberculatus (Orientuberculoides) capitatus (Essig and Kuwana) Alatae are pale green, greenish-yellow or whitish-yellow, antennae ringed with black, legs pale; BL 2.4-2.6 mm. Under leaves of Quercus spp. in east Asia (China, Taiwan, Korea, Japan). Ant-attended (Yao 2011). Oviparae and males were illustrated by Moritsu (1983). A single alata trapped in Korea was described as a subspecies, T. capitatus intermedius (Hille Ris Lambers 1974a); Quednau (1979) redescribed this form from specimens collected in Korea on Castanea crenata, and (Quednau 1999) also recorded it from Q. variabilis. 2n = 14.

[Note that the illustrations of T. capitatus by Quednau (1999, p. 231, figs 872-887) should be substituted by those in Quednau (2003, p. 301, figs 2451-2464).]

Tuberculatus (Nippocallis) castanocallis (Zhang & Zhong) Alatae squat-bodied with thickly-bordered wing veins, femora dark distally, conspicuous large dark marginal abdominal tubercles and dark siphunculi; BL c. 2.2 mm. On upper sides of the leaves of Castanea bungeana and C. seguinii in China (Liaoning, Hebei, Shandong, Zheijiang, Hunan, Guangxi, Yunnan; Qiao & Zhang 2002a, Qiao et al. 2005b).

Tuberculatus (Nippocallis) cereus (Zhang & Zhong) Alatae are squat-bodied with very thickly-bordered forewing veins, the pigment extending between Cu_1b and the branches of the media, and dark marginal abdominal tubercles and siphunculi; BL c. 1.9 mm. On Castanea bungeana in Shandong, China (Qiao & Zhang 2002a).

Tuberculatus (Nippocallis) ceroerythros Qiao & Zhang Alatae are pale red in life, coated with white powder, with thickly-bordered wing veins; BL c. 2.3 mm. On Castanea bungeana and C. seguinii in China (Guangxi, Jiangxi; Qiao & Zhang 2002a, Qiao et al. 2005b). Sexual morphs and life cycle are unknown.

Tuberculatus (Pacificallis) chrysolepidis Quednau Colour of alatae in life is unknown; BL 1.9-2.9 mm. On Quercus chrysolepis in California, USA (Quednau 1992). Sexual morphs and life cycle are unknown.

Tuberculatus (Pacificallis) columbiae Richards Colour in life was not observed; BL 2.0-2.5 mm. On Quercus garryana, recorded from Oregon (BMNH collection, leg. D. Hille Ris Lambers) and British Columbia (Richards 1965). Sexual morphs have not been described.

Tuberculatus cornutus Richards Colour in life is not recorded; BL c. 1.9mm. On an unidentified Quercus sp. in Turkey. Sexual morphs have not been described, but the ovipara, observed in October (Richards 1969a), is illustrated by Quednau (1999).

Tuberculatus (Tuberculoides) eggleri Börner Alatae are pale green or yellow-green with orange thorax, banded antennae and siphunculi pale, or only dark apically; BL 1.7-2.6 mm. On upper or undersides of leaves of Quercus spp. (especially pubescens; also recorded from aegilops, canariensis, cerris, etruscus, frainetto, ilex, mirbeckii, petraea). In southern, central and eastern Europe, North Africa (Algeria), and eastward to Turkey and Moldavia. Monoecious holocyclic on Q. pubescens, but it may overwinter as viviparae in mild winters. Lampel (1974) discussed seasonal variation and differences from T. borealis. [T. africanus Hille Ris Lambers, described from Quercus mirbeckii (canariensis) in Algeria (Hille Ris Lambers 1974a), is the early summer form of T. eggleri (Quednau 1999).]

Tuberculatus (Tuberculoides) etruscus Barbagallo & Binazzi Alatae are yellowish-green, with head and thorax slightly brownish, and siphunculi darker towards apices; BL 1.5-2.9 mm. On Quercus frainetto in Italy (Barbagallo & Binazzi 1991). Sexual morphs and life cycle are unknown.

Tuberculatus (Orientuberculoides) fangi (Tseng & Tao) Alatae are pale green with banded antennae; BL 1.8-2.9 mm. On undersides of leaves of Quercus spp. (acutissima, variabilis) in China, Korea and Japan (Hille Ris Lambers 1974a, Quednau 1999). Sexual morphs have not been recorded.

Tuberculatus (Orientuberculoides) fuscotuberculatus Zhang, Zhang & Zhong Alatae are yellowish green to green with banded antennae, dark longitudinal stripes on tibiae, tubercles on abdominal tergites 1-2 or 1-3 often dark and arising from dark sclerites, and siphunculi dark at least distally; BL 2.6-3.2 mm. On Quercus acutissima in China (G. Zhang et al. 1990b), and also recorded from Japan, and Korea, where two vagrant oviparae were collected in November (Quednau & Lee 2001; described and illustrated by Quednau 2003, p. 291).

Tuberculatus (Toltecallis) garciamartelli Remaudière & Quednau Alatae are whitish, with banded antennae and a pair of irregular pale green submarginal stripes on the abdomen; BL 1.4-1.9 mm. On Quercus microphylla in Mexico (Remaudière & Quednau 1983). Sexual morphs are unknown.

Tuberculatus (Acanthocallis) grisipunctatus Zhang, Zhang & Zhong Alatae are green in life; BL 1.7-2.3 mm. Recorded from various Quercus spp. in China (Zhang et al. 1990b, Quednau 1999), although according to Qiao et al. (2005b) it occurs only on Q. dentata. Sexual morphs are unknown.

Tuberculatus (Orientuberculoides) higuchii Hille Ris Lambers Alatae are pale yellow (Quednau & Shaposhnikov 1988); BL 1.8-2.7 mm. On Quercus spp. (dentata, mongolica, serrata) in Japan, Korea, China and eastern Siberia. In Japan, populations with thicker and blunter abdominal spinal processes and shorter PT were described by Hille Ris Lambers (1974) as a subspecies, T. higuchii breviunguis, which might be seasonal variants. It is not clear whether “higuchii-a” and “higuchii-b” of Yao (2011) correspond to these two named taxa. Hoever, DNA barcoding studies have now indicated that the name T. higuchii may be being applied to as many as four cryptic species, with minor morphological differences betwen them (Y. Lee et al. 2017). Sexual morphs are unknown.

Tuberculatus (Acanthotuberculatus) indicus L.K. Ghosh Alatae are reddish to greenish-yellow with pale eyes, banded antennae, mainly dark hind femora and bases of hind tibiae and long black spinal processes on abdominal tergites 2 and 3, contrasting with long pale processes on thorax and abdominal tergite 1 (see Moritsu 1983; as T. fulviabdominalis); BL 2.0-3.2 mm. On Quercus spp. in India, Korea, China and Japan, attended by ants (Yao 2011). The life cycle is variable; alate viviparae were collected in December-January in Manipur, India (Chakrabarti 1988), but oviparae and males were collected in South Korea in October-November (Quednau & Lee 2001). The ovipara is illustrated by Quednau (1999). This is the species referred to in Japanese and Korean literature (e.g. Higuchi 1972, Paik 1972) as T. fulviabdominalis (Shinji) – see Hille Ris Lambers (1974). However, DNA barcoding studies have now indicated that the name T. indicus may be being applied to more than one species (Y. Lee et al. 2017).

Tuberculatus (Tuberculoides) inferus Barbagallo Alatae are yellowish green, lightly dusted with wax, with banded antennae and dark-tipped spinal processes on abdominal tergite 3; BL 1.3-2.2 mm. Described from Quercus pubescens (sensu lato) in Sicily, inhabiting only the undersides of the most basal leaves on young plants (Barbagallo 1990). Also known from Quercus sp. in Turkey (BMNH collection, leg. D. Hille Ris Lambers). Life cycle and sexual morphs are unknown.

Tuberculatus (Acanthotuberculatus) japonicus Higuchi Alatae are yellow to green or brown, with pale eyes, mesothorax yellowish-green or brown dorsally and dark brown ventrally, and dark green-black spinal processes on ABD TERG 2 and 3 (Quednau & Shaposhnikov 1988); BL 2.5-2.9 mm. On Quercus spp. (dentata, mongolica), attended by ants, in Korea, Japan and eastern Siberia, and also recorded from China (Qiao et al. 2005b). Sexual morphs were collected on Q. dentata (possibly the preferred host) in Korea in early November (Quednau & Lee 2001).

Tuberculatus (Orientuberculoides) kashiwae (Matsumura) Alatae are pale yellow to pale greenish-yellow, sometimes yellowish-white or nearly white (Quednau & Shaposhnikov 1988); BL 1.4-1.9 mm. On undersides of leaves of Quercus spp. (dentata, mongolica) in Japan, China, Korea and eastern Siberia. DNA barcoding studies (Y. Lee et al. 2017) indicate that there may be two cryptic species under this name, the distribution and host specificity of which needs to be clarified. Oviparae and a male were collected in Japan in the second half of September (Quednau 2003), and Paik (1972) records oviparae in Korea in early November. 2n = 14.

Tuberculatus (Pacificallis) kiowanicus Hottes Alatae are pale yellow, yellowish green or green, with head and thorax light brown, orange or pinkish, often dusky on sides; BL 1.6-2.6 mm. On undersides of leaves of Quercus gambelii and Q. gunnisonii in Colorado and Utah, USA. Sexual morphs were collected on Q. gambelii in October (BMNH collection, leg. G.F. Knowlton) and the male was described by Hottes (1949b, as Myzocallis tonkawa). Palmer’s (1952) account under the name M. maureri applies to this species.

Tuberculatus (Orientuberculoides) konaracola (Shinji) Alatae are pale green or pale yellow-green to almost white, with banded antennae, brown marginal stripes on pronotum, dark Cu_1a and Cu_1b in forewing, dark tibiae, brown-black spinal processes on abdominal tergite 3 and dark distal halves of siphunculi; BL 1.7-2.8 mm. On Quercus spp. (serrata, dentata) in Japan (Shinji 1941); redescribed by Hille Ris Lambers (1974) from alatae trapped in Korea. Holocyclic in Japan; Shinji (1941) described the sexual morphs, and Quednau (1999) illustrated an ovipara from Q. dentata in Japan in November.

Tuberculatus (Orientuberculoides) kunugi (Shinji) According to the original description, alatae are green with black eyes, banded antennae and with long spinal processes on thorax and abdominal tergites 1-3 that are all concolorous with body (Shinji, 1924); BL not recorded. On Quercus acutissima and Q. serrata in Japan (Higuchi & Miyazaki 1969); no specimens are known, and it could not be included in the key.

Tuberculatus (Nippocallis) kuricola (Matsumura) Alatae are rather squat-bodied, pale green (Essig & Kuwana 1918) or red (R.L. Blackman, unpublished observations), covered with white wax; BL 1.2-1.9 mm. Forewings are patterned with very broad bands of fuscous that follow the veins. Brachypterae occur commonly (Takahashi 1924). On undersides of leaves, along the mid-ribs, of Castanea spp., especially C. crenata, attended by ants (Yao 2011). Records from Quercus require confirmation. In Japan, China, Taiwan and Korea, and introduced to Brazil (Bergmann et al. 2002), Spain (Manilla et al. 2001) and Madeira (Ilharco 1984). DNA analysis (Y. Lee et al. 2017) has indicated that the name might be being applied to two species with minor morphological differences; it is not known whether one of these could be M. kuricola var. cantonensis of Takahashi (1936a). Monoecious holocyclic in Japan; sexuales occur in October-November (Takahashi 1924, Higuchi 1972). Sakata (1999) studied interactions between T. kuricola, Lachnus tropicalis and the ant Lasius niger. 2n=14 (Shibata’s record of 2n=12 in both male and female cells must be erroneous.)

Tuberculatus (Toltecallis) leptosiphon Quednau Colour of alata in life unknown; BL c.1.7 mm. On Quercus microphylla in Mexico (Quednau 1999). Sexual morphs are unknown.

Tuberculatus (Acanthocallis) macrotuberculatus (Essig & Kuwana) Alatae are dark green or mid-brown with blackish cross-bands on abdominal tergites 1 and 2 bearing dark (sometimes pale-tipped) processes and forewing veins dark-bordered; BL 2.0-3.2 mm. Immatures are pale green, sometimes darker posteriorly, often with a whitish spinal band. On upper and undersides of leaves of Quercus dentata, attended by ants. In Japan, and probably more widely distributed, as previously synonymised and confused in the literature with T. quercicola (e.g. Quednau 1999). Watanabe et al. (2015) described two forms of this species from central and eastern Hokkaido. 2n=16 (Blackman 1986, as T. quercicola).

Tuberculatus maculipennis Hille Ris Lambers Alatae are greyish due to wax dusting, with pale antennae and legs, dark head and thorax, blackish spinal and marginal processes, and maculate forewings; BL 1.0-2.0mm. On Quercus spp. (infectoria, ?calliprinos, ?robur) in south-west Asia (Turkey, Lebanon, Iraq) and also recorded from Q. pubescens in Italy (Patti & Barbagallo 1997). Sexual morphs were collected in Turkey in October-November (Hille Ris Lambers, 1974).

Tuberculatus (Nippocallis) margituberculatus (Zhang & Zhong) Alatae are squat-bodied with dark head and thorax, wing-veins very thickly bordered with pigment extending between Cu_1a and branches of media (but leaving basal parts of these veins unbordered), very conspicuous large black marginal tubercles on abdominal tergite 4, and dark siphunculi; BL c. 1.9 mm. On Castanea bungeana in China (see Qiao & Zhang 2002a), and in Hong Kong it was found on C. mollissima (Martin & Lau 2011). Records from Quercus spp. (Qiao et al. 2005b) require further confirmation. Quednau (1999) illustrated an ovipara collected on Castanea sp. in October.

Tuberculatus (Pacificallis) maureri (Swain) Alatae are light green to apple green with red eyes, dusky-dark tibiae and forewing with a conspicuously dark-bordered Cu_1b vein; BL 1.4-2.2 mm. On Quercus agrifolia and Q. kellogii in California, USA. Sexual morphs were found in early November (BMNH collection, leg. D. Hille Ris Lambers).

Tuberculatus (Tuberculoides) maximus Hille Ris Lambers Colour in life is unknown, probably pale, with banded antennae, forewings with dark triangular spots at distal ends of Cu_1a and branches of media, and a rather dark cauda; BL 2.0-3.1 mm. On Quercus spp. (macranthera, persica) in southwest Asia (Iran, Turkey, Armenia), and there is also a record from Greece (Tsitsipis et al. 2007). Hille Ris Lambers (1974) discussed seasonal and geographic variation; sexual morphs tentatively ascribed to this species were collected on Q. macranthera in Iran in November (an ovipara is illustrated by Quednau 1999).

Tuberculatus (Toltecallis) mexicanus Remaudière & Quednau Alatae are very pale green to whitish with a pair of dark lateral stripes on prothorax and a pair of darker green lateral spots on each of abdominal tergites 1, 3 and 6; BL 1.3-2.4 mm. On Quercus spp. (peduncularis, rugosa) in Mexico (Remaudière & Quednau 1983), and subsequently recorded from Panama (Quirós et al. 2009). Apparently anholocyclic.

Tuberculatus (Tuberculoides) moerickei Hille Ris Lambers Plate 9b Alatae are pale yellow with banded antennae and distally dark siphunculi; BL 1.4-2.6mm. On Quercus spp. (especially infectoria) in Mediterranean (Cyprus) and southwest Asia (Iraq, Israel, Lebanon, Turkey). A sample from Q. ?pedunculata (= robur) in Turkey was described as a subspecies, T. moerickei galatensis (Hille Ris Lambers, 1974), but was considered to be merely a variant population by Quednau (1999). 2n = 14.

Tuberculatus (Tuberculoides) naganoe (Shinji) According to original description, alatae are light green with red eyes, banded antennae and black tibiae, and with pale finger-like spinal processes on head, thorax and each abdominal tergite; BL c. 1.6mm. On Quercus spp. in Japan (Shinji 1941). No specimens are known and it could not be included in the key on the basis of the available information. It was considered a nomen dubium by Remaudière & Remaudière (1997).

Tuberculatus (Tuberculoides) neglectus Krzywiec Alatae are pale yellow, with darker yellow mesothorax, eyes usually whitish yellow, antennae banded, siphunculi dark except at base (see influentialpoints.com/Gallery); BL 1.4-2.2 mm. On undersides of leaves of Quercus petraea, more rarely on Q. robur or hybrids between the two. In northern and north-west Europe (Britain, Ireland, Sweden, Denmark, Germany, northern Poland). Sexual morphs occur in November (Krzywiec 1965; Heie 1982).

Tuberculatus (Orientuberculoides) nervatus Chakrabarti & Raychaudhuri Colour of alata in life unrecorded; antennae banded, thorax probably dark with postscutum black, forewing veins slightly bordered with fuscous, the long, finger-like spinal processes on abdominal tergite 1-3 are dusky to dark, and siphunculi are dark towards apices. BL c. 3.6mm. On Quercus griffithii (see Quednau 1999) in India (Meghalaya, Manipur, Nagaland; Chakrabarti & Raychaudhuri 1976, Chakrabarti 1988). Life cycle and sexual morphs are unknown.

Tuberculatus (Acanthocallis) nigrosiphonaceus (Zhang & Zhang) Colour of alatae in life unknown, but with dark head and thorax, dusky wings with dark veins, dark transverse bars on abdomen and dark siphunculi; BL c. 2.3 mm. On Quercus liaotungensis in China (Beijing; W. Zhang & G. Zhang 1994, as Myzocallis), and oviparae were collected in Japan in late September on Quercus mongolica var. grosseserrata (described and illustrated by Quednau 2003, p. 292-3). Quednau (1999) transferred this species to Tuberculatus (Acanthocallis).

Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers Alatae are pale greenish-yellow (Quednau & Shaposhnikov 1988), in midsummer the tibiae and finger-like processes on abdominal tergite 3 may be dark (Quednau 1999); BL 2.4-3.2 mm. On undersides of leaves of Quercus spp. (dentata, mongolica) in India (Meghalaya), Bhutan (Chakrabarti & Das 2014), Korea, China, eastern Siberia and Japan (Yao & Katagiri 2011). A.K. Ghosh & Quednau (1990) provided a redescription. Sexual morphs and life cycle are unknown.

Tuberculatus (Tuberculoides) pallescens Hille Ris Lambers Alatae are bright green, with banded antennae including short blackish-brown sections near middle of ANT III; BL 2.5-2.5 mm (Hille Ris Lambers 1974a). On Quercus spp. (calliprinos, infectoria, ?robur) in south-west Asia (Iraq, Lebanon, Turkey), and also recorded from Q. pubescens in Sicily (Barbagallo & Stroyan 1982). Sexual morphs were collected on Q. calliprinos in Turkey in November.

Tuberculatus (Pacificallis) pallidus (Davidson) Alatae are pale green; BL 1.5-2.2 mm. On leaves of a wide range of Quercus spp. (including non-native oaks) in western USA. Partly anholocyclic; alatae were collected in December-January in southern California. Separation from T. quercifolii, which is holocyclic on native Californian oaks is difficult and needs further investigation.

Tuberculatus (Orientuberculoides) paranaracola Hille Ris Lambers Alatae are greenish yellow or yellowish green to pale green (with ‘dark green reflecting whitish’), with banded antennae and dark tibiae (Quednau & Shaposhnikov 1988); BL 1.6-2.3 mm. On undersides of leaves of Quercus mongolica (incl. var. grosseserrata) in Japan, China, Korea and eastern Siberia. Hille Ris Lambers (1974) distinguished a subspecies, T. paranaracola hemitrichus, from yellow traps in Korea, and this subspecies is also recorded from Q. acutissima and Quercus sp. in China (Chen & Zhang 1985b, Quednau 1999). The life cycle and sexual morphs are unknown. 2n = 14 (for ssp. hemitrichus).

Tuberculatus (Pacificallis) pasaniae (Davidson) Alatae are pale green mottled with darker green and with ‘many delicate pruinose markings’ (Davidson 1915); eyes dark red, antennae ringed with black, spinal process whitish; BL 2.0-2.6 mm. On undersides of leaves of Lithocarpus densiflora in California and Oregon, USA. An ovipara collected on Lithocarpus in November was illustrated by Quednau (1999), but the occurrence of alate viviparae in November to mid-February indicates at least partial anholocycly.

Tuberculatus (Pacificallis) passalus Quednau Colour of alatae in life is unknown; BL 1.3-2.2 mm. On Quercus dumosa in California USA. Oviparae were collected in late September-December, but it is apparently partially anholocyclic, as alate viviparae occur in December and March (Quednau 1992).

Tuberculatus (Acanthotuberculatus) pilosulus Quednau Colour of alatae in life is unknown; BL 1.6-2.5 mm. On Quercus serrata and Quercus sp. in Japan, attended by ants (Yao 2011). Oviparae were collected in the second half of November (Quednau 1996).

Tuberculatus (Nippotuberculatus) pilosus (Takahashi) Alatae are yellowish brown with blackish eyes, with forewing veins having broad brown borders; BL 1.9-2.8 mm. Described from Quercus sp. in Taiwan (Takahashi 1929) and redescribed from Japan on Q. phillyraeoides (Higuchi 1970), but there are some discrepancies between the two descriptions which warrant further investigation. Yao (2011) identified a population on Lithocarpus edulis in Japan as this species. Sexual morphs and life cycle are unknown.

Tuberculatus querceus (Kaltenbach) Alatae are dirty greenish to straw-coloured, covered with fine powdery white wax, with a dark bifurcate spinal process (fig. 94A), a dark patch near end of hind femur and dark siphunculi; BL 1.4-2.4 mm. On undersides of leaves of Quercus robur in Europe and east to Iran, Turkey and the Caucasus (Heie, 1982), Central Asia (Kadyrbekov 2017a), and introduced to British Columbia, Canada (Foottit et al. 1999), and to South America (Argentina, Chile; Caballero et al. 2000, Ortego et al. 2006). DNA barcoding has raised the possibility that specimens from Europe (Poland) and Canada may be different cryptic species (Y. Lee et al. 2017). Sexuales occur in October.

Tuberculatus (Acanthocallis) quercicola (Matsumura) Alatae are green with brown head and thorax, banded antennae, dark hind femora, dusky wings with bordered wing veins, and dark bars on abdominal tergites 1 and 2; BL 1.8-2.7 mm. On upper and undersides of leaves of Quercus mongolica (incl. ssp. crispula), attended by ants. Records from other Quercus spp. may be incidental; those from Q. dentata should probably be referred to T. macrotuberculatus (Watanabe et al. 2015). In Japan, Korea, China and eastern Siberia. Quednau & Shaposhnikov (1988) noted some differences in life between populations on Q. dentata and Q. mongolica (but their aphids on Q. dentata were probably T. macrotuberculatus, q.v.). Yao & Akimoto (2002), Yao (2011) and Yao & Katagiri (2011) studied the effects of ant attendance on composition of the honeydew and wing development. Yao (2012) described the life cycle and studied seasonal changes in the flight apparatus in male and alate females. 2n = 16 (Chen & Zhang 1985b).

[Also note:

1. T. pappus Zhang, Zhang & Zhong 1990 is probably a synonym; oviparae collected in Japan on Q. mongolica var. grosseserrata (= ssp. crispula) in late September, described under that name by Quednau (2003, p. 293-4), and illustrated along with a male (2003, p. 299), should probably be referred to T. quercicola.

2. DNA barcoding of specimens from Korea has revealed that aphids from Q. aliena identified as T. quercicola may be another distinct, cryptic, host-specific species (Y. Lee et al. 2017)].

Tuberculatus (Pacificallis) quercifolii (Davidson) Alatae are pale green with olive green head and mesothorax (Davidson, 1919); BL 1.2-2.4 mm. On leaves of Quercus douglasii in California, USA. Apparently holocyclic; fundatrices occur in April (BMNH collection, leg. D. Hille Ris Lambers), and an ovipara collected in early December was illustrated by Quednau (1999). Hille Ris Lambers (1974) probably had a mixture of T. quercifolii and T. pallidus (see Quednau 1992).

Tuberculatus (Orientuberculoides) querciformosanus (Takahashi) Alatae are yellow or yellowish white with yellow eyes and antennae ringed with black; BL 1.5-2.5 mm. On Quercus spp. (aliena, dentata, mongolica, variabilis) in Japan, China, Taiwan, Korea and eastern Siberia. Sexual morphs and life cycle are unrecorded.

Tuberculatus (Acanthotuberculatus) radisectuae Zhang, Zhang & Zhong Alatae are brown and have wings dusky with dark veins, black distal parts of hind femora, and tubercles on abdominal tergites 2 and 3 black and arising from dark transverse bars (but those on abdominal tergite 1 pale); BL c. 2.6 mm. On Quercus fabri in China (Zhang et al. 1990b, as T. japonicus ssp. radisectuae). 2n = 14 (Chen & Zhang 1985b).

Tuberculatus (Tuberculoides) remaudierei Nieto Nafría Alatae are pale green with red eyes; BL 1.8-3.0 mm. On Quercus pyreniaca in Spain, and there is also a record from Greece (Tsitsipis et al. 2007). Sexual morphs occur in October (Nieto Nafría & Mier Durante 1978).

Tuberculatus (Toltecallis) spiculatus Richards Alatae are white to very pale green; BL 1.9-2.4 mm. On Quercus rugosa and Q. microphylla in Mexico, and also recorded from Q. arizonica in Arizona, USA (Quednau 1999). A subspecies, T. spiculatus rebecae, was described from Q. rugosa in Mexico (Remaudière & Quednau 1983). Sexual morphs are unknown.

Tuberculatus (Arakawana) stigmatus (Matsumura) Alatae are shining dark brown to bluish-black with banded antennae, pale fore legs and black hind legs (see Moritsu 1983); BL 2.0-2.6 mm. Immatures are pinkish blue or pinkish brown, with longitudinal rows of black spots. On Quercus spp. in Japan, Korea, China and eastern Siberia. DNA analysis has indicated that the name may be being applied to two species with minor morphological differences (Y. Lee et al. 2017). Sexual morphs occur in October in Korea (Paik 1972).

Tuberculatus (Orientuberculoides) yokoyamai Takahashi Alatae are pale greenish yellow or pale yellow with whitish eyes and banded antennae (see Moritsu 1983); BL 1.6-2.4mm. On Quercus spp. (crispula, dentata, mongolica, serrata) in Japan, China, Korea and eastern Siberia. DNA barcoding has indicated that the name may be being applied to two species with minor morphological differences (Y. Lee et al. 2017). An ovipara was collected on Q. dentata in South Korea in November (Quednau & Lee 2001; described and illustrated by Quednau 2003, p. 292). 2n=14.

Tuberoaphis Tseng & Tao

Aphidinae: Macrosiphini

One or two species associated with Hydrangea, having long dorsal hair-bearing processes, and eyes without distinct ocular tubercles. The life cycle is unknown; possibly there is a host alternation to ferns, as in the related genus Macromyzus.

Tuberoaphis hydrangeae Tseng & Tao Apterae are yellow, with apices of appendages dark brown; BL c.1.1-1.2 mm. On leaves of Hydrangea aspera in Szechuan, China.

Tuberoaphis hydrangeae ssp. digitata Hille Ris Lambers & A.N. Basu Plate 22d (Fig.32a) Apterae are pale glassy white to pale yellowish green, with apices of appendages darker; BL 1.4-1.7 mm. On undersides of leaves of Hydrangea robusta in West Bengal, India, producing large numbers of alatae in late May and probably migrating to an unknown secondary host (original description). Matsumuraja indica A.K. Ghosh, M.R. Ghosh & Raychaudhuri, described from (?) Rubus sp., is a synonym, but Rubus is an unlikely secondary host.

Tuberocephalus Shinji

Aphidinae: Macrosiphini

Fifteen or more species of Myzus-like aphids in east Asia, often with hair-bearing siphunculi, galling the leaves of Prunus in spring. Where the life cycle is known there is host alternation to Anthemidae (Artemisia, Chrysanthemum), where they are small aphids on growing points or undersides of leaves (Tuberocephalus s.str.) or on subterranean shoots or roots (subg. Trichosiphoniella). The generations on secondary hosts look very different from the spring forms on Prunus, and this has contributed to the taxonomic confusion in the group. Accounts are available from Japan (Miyazaki 1971), China (Su et al. 2010) and eastern Russia (Pashchenko 1988a). Moritsu & Tokumoto (1972) gave an account of Japanese Tuberocephalus (under Myzus) on flowering cherries in which they included several unnamed species. Sorin & Remaudière (1998) reviewed the genus and provided new information about morphs and life cycles.

Tuberocephalus artemisiae Shinji Spring colonies are in large, bag-shaped galls produced on the upper sides of the leaves of Prunus donarium “var. spontanea subvar. speciosa” (= P. jamasukara var. speciosa?) and P. yedoensis. The galls are yellowish green becoming bright red on upper side when mature. Fundatrices are dark green, BL c. 2.2 mm, and all their progeny are alatae (BL c.1.6 mm, with greenish-yellow abdomen without a central patch), which migrate to Artemisia spp. Apterae on Artemisia are pale yellow to dull yellow, body rather elongate, with pale appendages; BL 1.1-1.5 mm. They form colonies on the undersides of the leaves, heavy infestations curling leaves and causing plants to become sooty with mould (Miyazaki 1971). Gynoparae and males return to Prunus in October-November (Sorin & Remaudière 1998). It is recorded from Japan, China and east Siberia.

Tuberocephalus (Trichosiphoniella) higansakurae (Monzen) Spring colonies in galls caused by upward rolling and thickening of edges of distal parts of leaves of cherry trees, the sausage-shaped leaf-roll becoming yellowish-green or red. Dark green fundatrices give rise to reddish-yellow apterae (BL 1.5-1.9 mm) and alatae (BL c. 1.8-1.9 mm). Alatae have a dusky yellow abdomen with a dark central dorsal patch and fly to an unknown secondary host. Greenish-white apterae of a Tuberocephalus that could be this species were collected on Gnaphalium (BMNH collection, leg. R.L. Blackman), but this needs experimental verification. On Prunus spp. (cherries) in Japan, Korea and China; however, it seems that Zhang & Zhong (1976) in China may have had T. misakurae. Monzen (1929) recorded alatae (gynoparae) returning to P. subhirtella in October-November, but did not describe the sexual morphs.

Tuberocephalus (Trichosiphoniella) jinxiensis Zhang & Zhong Described from apterae only (BL 1.4-1.8 mm), collected on Prunus humilis in China in June (Zhang & Zhong 1976). Barrel-shaped galls are formed on the undersides of the leaves along lateral veins. The life cycle is unknown. It seems very similar to T. higansakurae, but with shorter siphunculi.

Tuberocephalus lazikouensis Zhang, Chen, Zhong & Li, in Zhang Apterae are green; BL c.1.7 mm. On undersides of leaves of Artemisia argyri in Gansu province, China (Zhang 1999), and subsequently in Shaanxi and Yunnan provinces (Su et al. 2010). Other morphs and life cycle are unknown.

Tuberocephalus (Trichosiphoniella) liaoningensis Zhang & Zhong Galls on Prunus tomentosa in Japan are formed from curled leaves and are multi-coloured yellow and green with some reddish purple when mature (Sorin & Remaudière 1998). Fundatrices are deep blackish green, producing blackish-brown apterae (BL 1.7-1.9 mm), and emigrant alatae that migrate to Artemisia vulgaris var. indica (in Japan). The colonies on Artemisia are found on subterranean young shoots. This species was originally described from China (Zhang & Zhong 1976), where it is a common species on Prunus pseudocerasus, with heavy infestations curling and withering young leaves in spring, and it is also recorded from P. salicina (G. Zhang 1999) and Amygdalus triloba (Su et al. 2010). 2n=12 (Chen & Zhang 1985a).

Tuberocephalus (Trichosiphoniella) misakurae Moritsu & Hamasaki Spring colonies live in bag- and sausage-shaped galls formed by rolling and thickening of edges of distal parts of leaves of cherry trees, as in the closely-related T. higansakurae. It is recorded from Prunus pauciflora, P. pseudocerasus and P. takenakae in Japan (Moritsu & Hamasaki 1983) and from P. pseudocerasus in China (BMNH collection, leg. V.F. Eastop). The fundatrix is deep green or green; the colour of its apterous progeny is not recorded, BL 1.5-2.0 mm. Alatae have a yellowish-green abdomen with a dark central dorsal patch and migrate in May to form colonies on the thin lateral roots of Chrysanthemum. Apterae on the secondary host are greenish brown, BL 0.9-1.2 mm. Gynoparae, males and oviparae were collected in Japan in October-early December (Sorin & Remaudière 1998). 2n=12.

Tuberocephalus (Trichosiphoniella) momonis (Matsumura) Spring colonies roll the edges of the leaves of Prunus persica and turn them red. Fundatrices are dark green, broadly oval, with BL0 2.1-2.2 mm (Su et al. 2010). Apterous fundatrigeniae are yellowish-brown to dark brown with a greenish tinge (Matsumura 1917); BL 1.4-2.0 mm. Alatae have a pale yellowish-brown abdomen without a distinct dorsal patch and are produced in late May to June (Su et al. 2010 and BMNH collection, leg. V.F. Eastop), but colonies persist on peach into summer and autumn, so it seems likely that there is no host alternation. Su et al. (2010) described apterae collected in September, which were dark green with BL 1.4-1.8 mm, and also provided descriptions of alate gynoparae, alate males and oviparae collected in October. In Japan, China, Taiwan, Korea and Mongolia. This species has been confused in the literature with T. higansakurae, T. misakurae and T. sakurae. The true momonis seems to be specific to peach. 2n=12 (Chen & Zhang 1985a).

Tuberocephalus naumanni Sorin & Remaudière Mature galls formed on Prunus cornuta are shiny, thick-walled, pale green to white, 3-4 mm wide and 20-25 mm long. They contain up to three generations of pale to bright green aphids, all the third generation being emigrant alatae (BL 1.8-2.1 mm). Migration occurs in May to an unknown secondary host. Only known from Pakistan (Sorin & Remaudière 1998).

Tuberocephalus pakistanicus Remaudière & Sorin Mature galls on Prunus cornuta are sausage-shaped, papery, pitted, 2.5-3.0 mm wide and 15-30 mm long, dark green to red with reddish speckles and spots sometimes coalesced into a red patch on the upper side of the gall. All the progeny of the fundatrix are reddish-brown emigrant alatae, BL 1.6-2.1 mm, which emerge in early May and fly to an unknown secondary host (Sorin & Remaudière 1998). Gynoparae have been trapped in October (Naumann-Etienne & Remaudière 1995). It is only known from Pakistan.

Tuberocephalus (Trichosiphoniella) sakurae (Matsumura) Plate 17g Spring colonies cause terminal leaves of Prunus (Cerasus) spp. to become extensively curled, twisted and yellowed or reddened (see Moritsu 1983). Apterae on Prunus are deep green to black (fundatrices paler), with black siphunculi; BL 1.5-1.8 mm. Immatures are bright greenish yellow. Recorded from P. cerasus, P. jamasakura, P. lannesiana, P. maximowiczii, P. sachalinensis, P. serrulata, P. triloba and P. yedoensis, and from Japan, China, Korea and eastern Siberia (including Kuril Islands; as Sorbaphis kurilensis Ivanoskaya-Shubina 1966). It has recently been recorded from P. serrulata in USA (New York, Connecticut; Skvarla et al. 2017). Alatae are without a distinct dorsal abdominal patch. They migrate to the roots and subterranean young shoots of Artemisia spp. (Sorin & Remaudière 1998). Apterae on the secondary host are deep bluish green to black, BL c.1.6-1.7 mm, with immatures greenish yellow. Alate males and gynoparae are produced in October in Japan and were described by Sorin & Remaudière (1998). [N.B. Tao’s (1966) sakurae was probably Myzus yamatonis, and Miyazaki’s (1971) description is partly of misakurae.]

Tuberocephalus sasakii (Matsumura) Conspicuous, brownish, elongate pouch-like galls are produced in Japan on the upper sides of leaves of Prunus spp. between two veins (see Moritsu 1983). Progeny of the fundatrix are all alate, with bright yellow abdomen; BL 1.2-1.8 mm. Alatae are without a distinct dorsal abdominal patch. They leave the gall in June through an opening on the underside of the leaf and fly to found colonies on Artemisia spp. (Monzen 1929, Sorin & Remaudière 1998). Apterae on Artemisia are whitish or greenish, dorso-ventrally flattened, BL 0.8-1.3 mm, and live adpressed to the undersides of leaves The return migration occurs in October (Monzen 1929). In Japan, Korea, the Kuril islands (Pashchenko 1988a), Taiwan, China and India. It is possibly anholocyclic in West Bengal as it is common on Artemisia but not known to occur there on Prunus (Ghosh 1973). The record from Java (Noordam 2004) applies to T. tsengi (q.v.).

Tuberocephalus (Trichosiphoniella) tianmushanensis Zhang (= T. higansakurae ssp. hainnevilleae Remaudière & Sorin; synonymy by Su et al. 2010) Galls on Prunus are of two types; bag-shaped on the edge of tender leaves formed by the fundatrices, and leaf-rolls similar to those described for the nominal form, bright reddish yellow when mature and inhabited by their apterous progeny. Fundatrices are matt brown, BL 1.7-2.0 mm, giving rise to very dark apterae (BL 1.5-1.9 mm) and alatae (1.6-2.0 mm). Emigrant alatae have a black dorsal abdominal patch. On Prunus spp. (cherry trees) in China and Japan; also found in France on seedlings of P. subhirtella var. pendula imported from Japan (Remaudière & Sorin 1993, as T. higansakurae ssp. hainnevilleae), and it has possibly become established on the same host in northern Italy (Pellizzari & Frigimelica 2014). Emigrant alatae colonised subterranean young shoots of Artemisia in transfer experiments in Japan (Sorin & Remaudière 1998, as T. higansakurae ssp. hainnevilleae). Gynoparae, oviparae and males were collected on Prunus in mid- to late November. A redescription of the type specimens (from Zhejiang province, China) was provided by Su et al. (2010), identifying the host as P. pseudocerasus.

Tuberocephalus tsengi Tao Apterae are yellowish green with dark red eyes, black tips to antennae, legs and siphunculi, and a pale cauda (see influentialpoints.com/Gallery); BL 1.2-1.9 mm. On young growth of Artemisia spp. in China, Indonesia and West Malaysia, and recently found (on A. vulgaris) in southern England (Blackman et al. 2020). Presumably there is a primary host generation forming galls somewhere on Prunus, but this has still to be recognised, and its full distribution is uncertain because of confusion with T sasakii, with which it was erroneously synonymised. [In Japan there is an undescribed Tuberocephalus species on Gnaphalium sp., with shape of antennal tubercles and ANT I, reduced eyes and sculpturing of dorsal cuticle very like T. tsengi (BMNH collection, leg. R. Blackman, RLB 2171). The apterae are greenish white in life, and compared with T. tsengi it has shorter antennae, longer R IV+V and shorter cauda.]

Tuberocephalus usubai Sorin The gall is undescribed and only the emigrant alata is known, BL c.2 mm. On Prunus buergeriana in Japan (Sorin 2011). The alata is very similar to that of T. naumanni, but has more rhinaria on ANT III (7-12 compared with 3-8 in naumanni) and a much longer R IV+V (c. 1.2 × HT II, compared with 0.7-0.8 × HT II in naumanni).

Tuberocephalus (Trichosiphoniella) uwamizusakurae Sorin & Remaudière Fundatrices are dark green in life, inhabiting elongate pouch-like galls along the margins of the upper sides of the leaves of Prunus grayana in Japan. When mature the galls are brownish green and about 10 mm wide and 50-60 mm long. Spring migrant alatae (BL 1.7-2.0 mm) are produced in the second generation and migrate in April-May to found colonies on the curled lateral roots of Artemisia vulgaris var. indica. Apterae on Artemisia roots are probably dark green to black, BL 1.0-1.4 mm. An ovipara was collected on P. grayana in late November (Sorin & Remaudière 1998).

Tuberolachnus Mordvilko

Lachninae: Tuberolachnini

Three species with a single large conical spinal process or tubercle on abdominal tergite 4. A.K. Ghosh (1982b) provided an account of the two species then known. It seems probable that the presence of a dorsal tubercle is not a reliable character on which to base a genus, and that some other Asian Salix-feeding species currently placed in Lachnus (salicis, tatakaensis) may be more closely related to T. salignus than are the two species of Tuberolachniella.

Tuberolachnus (Tuberolachniella) macrotuberculatus Yang, Qiao & Zhang Apterae have a blackish-brown head and yellowish-brown thorax and abdomen, the abdomen bearing a large bladder-like tubercle; BL 5.1-5.3 mm. On Eriobotrya japonica in Sichuan, China (Yang et al. 2005a). Other morphs and biology are unknown.

Tuberolachnus salignus (Gmelin) Plate 15c, d Apterae are mid-brown to dark brown with rows of dark spots and a large dark brown tubercle in the centre of the dorsum (see influentialpoints.com/Gallery); BL 5.0-5.8 mm. Alatae have the forewing membrane unpigmented. On stems and branches of numerous Salix spp. and also very occasionally recorded from Populus. Large colonies build up in late summer and attract numerous honeydew-feeding insects. Virtually cosmopolitan in distribution, wherever willows grow or are planted; only recently (2013) reported to occur in New Zealand (Gunawardana et al. 2014) and Australia (2014; Dominiak &Worsley 2018). Apparently anholocyclic everywhere; no sexual morphs are known. Mittler (1957, 1958a, b) used this species in classic studies of aphid feeding and nutrition, and Hargreaves & Llewellyn (1978) studied its ecological energetics. Effects of temperature on development and fecundity on various willow species were studied by Collins & Leather (2001) and Özder et al. (2007), and selection between willow varieties was studied by Aradottir et al. (2009). Genetic diversity of Chinese populations was studied by F. Fang et al. (2017). In Japan, T. salignus is parasitised by a possibly specific parasitoid, Pauesia salignae, which might indicate that this is where it is endemic. 2n = 20.

Tuberolachnus (Tuberolachniella) sclerata Hille Ris Lambers & Basu Apterae are blackish with a conspicuous reddish-brown to black broad-based tubercle on the dorsum; BL 4.0-5.2 mm. In large colonies on undersides of leaves, petioles and young shoots of Eriobotrya petiolata in West Bengal (at 2100 m); on leaves they form rows along the mid-rib and main veins (Hille Ris Lambers & Basu, 1966). Specimens of this or a very similar species have also been collected on E. japonica in Indonesia (BMNH collection, leg. D. Noordam). Alatae and other morphs are undescribed, and the life cycle is unknown.

Tumoranuraphis Zhang, Chen, Zhong & Li

Aphidinae: Macrosiphini

A genus for one large Asian species with 5 hairs on first tarsal segments, clavate siphunculi and numerous rhinaria on antennal segment IV of alatae.

Tumoranuraphis indica (Chakrabarti & Maity) Apterae of BL about 4.6-4.9 mm, pale greyish covered with powdery wax, with brown, clavate siphunculi (fig. 88C). Alatae have black head and thorax and blackish-brown dorsal, abdominal markings. Fundatrices are greenish with dark dorsal spots. It feeds along the mid-ribs on upper sides of leaves of Prunus cornuta in late May-June, causing the leaf laminae to fold upwards and come together above the aphids. In Uttar Pradesh, India (Chakrabarti & Maity 1984), and recorded from north-west China on P. pseudocerasus (G. Zhang 1999, as T. cerasicola). Heteroecious holocyclic; alatae leave Prunus in late June to fly to an unknown secondary host (Medda et al. 1986). Oviparae were collected in China (Eastop & Blackman 2005).

Turanoleucon Kadyrbekov

Aphidinae: Macrosiphini

Two species on Asteraceae: Cardueae described from the desert zone of Kazakhstan, related to Macrosiphoniella but with an elongate R IV+V, a well-developed median frontal tubercle and 5 hairs on first tarsal segments (Kadyrbekov 2002a). The two species are very similar and the reported differences might be due to nutrition; Goodarzifar et al. (2016) provide comparative data.

Turanoleucon mitjaevi Kadyrbekov Apterae are black, BL 2.2-3.2 mm. On stems of Cousinia spp. in deserts of south and east Kazakhstan.

Turanoleucon jashenkoi Kadyrbekov Apterae are black, BL 2.3-4.0 mm. On stems of Echinops spp. in deserts of south and east Kazakhstan, and recorded also from Iran by Goodarzifar et al. (2016), who provide a redescription.