SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order)
An Old World genus of about 11 species mostly host-alternating between galls on Pistacia and the roots of Poaceae, and sometimes Cyperaceae, the life cycle taking two years (as in fig. 2). One species (mimeuri) is known to be monoecious on Pistacia. The taxonomy is difficult, particularly around G. utricularia (see below). Davatchi (1958) reviewed the morphs on Pistacia, and Brown & Blackman (1994) provided a key to emigrant alatae, and a partial key to sexuparae. Apterous exules and sexuparae have the anus displaced dorsally and an enlarged anal plate, which functions as a trophobiotic organ to retain honeydew for collection by ants. Emigrant alatae, however, have a quite different modification of the anal region, with the anal plate and cauda extended laterally to unite and form a sclerotised perianal ring, enclosing a membranous area with the anal aperture at its centre; why a different structure, presumably with the same function, has been developed in this morph is something of a mystery. The root-feeding generations are in accounts for north, central and southern Europe (Heie 1980b, Zwölfer 1958, Roberti 1939, respectively); and for India (Raychaudhuri 1980, A.K. Ghosh 1984).
Geoica anchusae Narzikulov Apterae are yellowish green or pale green, with greyish wax; BL c.2.9 mm. On roots of Anchusa italica in Tajikistan. It has subsequently been recorded in Kazakhstan from more likely hosts, the grasses Poa bulbosa and Eremopyrum orientale (Kadyrbekov 2014d).
Geoica harpazi Brown & Blackman Galls on Pistacia atlantica in Israel are not clearly identified, but probably similar in size and shape to those of G. utricularia; they may be the P. atlantica galls described by Wertheim (1954) as green, of rather irregular shape and with a wrinkled and uneven surface. Alate emigrants fly in October to found colonies on roots of Poaceae (Ammophila, Hordeum, Triticum). The appearance of the apterous exules in life is unknown; BL 1.7-2.6 mm. See Brown & Blackman (1994).
Geioca lucifuga (Zehntner) Apterae are pale yellowish or yellowish‑brown with a light covering of white wax; BL 1.9‑3.5 mm. Alatae have a yellowish abdomen with a broad transverse bar on each tergite, largest on the posterior segments. On roots of many genera and species of Poaceae and Cyperaceae, usually attended by ants. Recorded also from the roots of grape vine in Central Asia (Nevsky 1929a). Anholocyclic populations of this species occur world-wide except in cold temperate northern regions (Blackman & Eastop 2000, p.274-5). The holocycle has not been described, but Mordvilko (1935) suggested that the primary host might be Pistacia sinensis, and alatae (BL 2.1-2.2 mm) from galls on P. sinensis in Pakistan deposited first instars with chaetotaxy agreeing well with specimens of G. lucifuga from grass roots (Naumanne-Etienne & Remaudière 1995). Very similar alatae (but with BL 1.6-1.9 mm) have been collected from galls on P. khinjuk in Pakistan (BMNH collection, leg. M. Ghani). No information is available on the morphology of the galls in either case. 2n = 14 (Kulkarni 1984) or 18 (Blackman & Eastop 1994).
Geoica mimeuri (Gaumont) Monoecious holocyclic on Pistacia atlantica and (more rarely) P. terebinthus in Morocco. Galls are formed at the bases of leaflets, and resemble those of G. utricularia, but are smaller (Gaumont 1930). The alatae leaving the galls are sexuparae; Davatchi & Remaudiere (1957) found an alata containing one embryo with normal stylets, but this seems to be exceptional. Davatchi (1958) compared the sexuparae of G. mimeuri with those of G. setulosa and G. utricularia.
Geoica setulosa (Passerini) Galls on Pistacia khinjuk in Iran are formed at the bases of leaflets, and are like those of G. utricularia, but larger (Davatchi 1958); the gall on P. khinjuk in Israel illustrated by Koach & Wool (1977) is probably that of G. setulosa. They open in late August-October. Emigrant alatae (BL 1.8-2.3 mm, fig. 81I) found colonies on roots of grasses (e.g. Agrostis, Briza, Festuca, Holcus), attended by ants. Apterous exules are off‑white or pale greenish‑grey; BL 1.6-2.6 mm. In northern and central Europe, Spain (Pérez Hidalgo et al. 2007a), Italy, Iran, Turkey, Georgia, and introduced to USA (G.L. Miller et al. 2002). Where P. khinjuk does not occur G. setulosa is exclusively anholocyclic on grass roots, often overwintering in the nests of Lasius flavus. 2n = 20 and 24 (from grass roots).
Geoica sikkimensis Raychaudhuri, Pal & M.R. Ghosh Apterae are brownish; BL 1.9-2.1 mm. On roots of Polypogon fugax, and probably other grasses, in Sikkim and West Bengal, India.
Geoica swirskii Remaudière & Inbar Galls on Pistacia atlantica are of irregular shape, bulbous, c. 25 mm in diameter, arising from leaf stem between bases of leaflets. Apterous fundatrigeniae and emigrant alatae were collected from the galls in October (Remaudière et al. 2004). Only known from Jordan.
Geoica utricularia (Passerini) group Galls on Pistacia spp. in the Mediterranean area and south-west Asia are roughly globular, and are formed directly by the fundatrix feeding near the base of a leaflet, close to the main vein (fig. 126K, and Wertheim 1954). Colour, texture and shape seem to vary somewhat with the host species, but on P. palaestina and P. terebinthus they are smooth and yellowish with a pinkish tinge. Gall histology and its phylogenetic implications were studied by Álvarez (2011, 2012). Emigrant alatae (BL 1.7-2.6 mm, fig. 81H) emerge in July-October - the time of opening of the galls also seems to depend on the host species - and fly to colonise roots of Poaceae. Apterous exules on grass roots are off white, cream or yellowish white, lightly dusted with wax; BL 1.6‑3.0 mm (Plate 1i). Alatae have a pale yellowish‑green abdomen with dark transverse bars, largest on the more posterior tergites. Numerous species of grasses and cereals are colonised (e.g. Agrostis, Avena, Bromus, Deschampsia, Festuca, Hordeum, Lolium, Phleum, Poa, Triticum, Zea), and the colonies are invariably ant‑attended. All records are from Poaceae except for a colony found on Orobanche foetida in Tunisia (Boukhris-Bouhachem et al. 2011). A full account of the life history in Italy of G. utricularia s. str., which uses P. terebinthus as its primary host, was given by Roberti (1939, 1983). The considerable variation in morphology of galls under the name of G. utricularia, and of emigrant alatae and their embryos, has led to the recognition of other taxa closely related to G. utricularia but associated with other Pistacia spp.; G. muticae Mordvilko on P. mutica, G. rungsi Davatchi & Remaudiere on P. atlantica and G. wertheimae Brown & Blackman on P. palaestina (Brown & Blackman 1994). Koach & Wool (1977) also reported electrophoretic differences between samples from P. atlantica (perhaps G. rungsi) and P. palaestina (perhaps G. wertheimae), indicative of discrete populations on these two primary hosts in Israel. Wool & Koach (1976) studied the effects of genotype and environment on morphological variation in emigrant alatae of G. utricularia in Israel. The name G. utricularia is conventionally applied to anholocyclic populations of this group on grass roots throughout Europe and in North Africa (Morocco, Tunisia), the Middle East, Central Asia, North America, and China (as ssp. urunquiensis Qiao). However these populations vary considerably in chaetotaxy, particularly of R IV+V, abdominal tergite 8, cauda and anal plate, and it is possible that more than one species is involved, or that the name being applied is incorrect. Ivens et al. (2012a,b) used microsatellite DNA analysis to investigate clonal diversity of root-feeding populations identified as G. utricularia in the Netherlands. Remaudière & Remaudière (1997) list eight other available names for species in “the utricularia group”. Wool & Kurzfeld-Zexir (2008) noted preferential colonisation by G. wertheimae of barley compared with other cereals in laboratory rearing experiments. See also Blackman & Eastop (2000, p.275). 2n = 16, 17 and 18 (2n=18 for holocyclic populations on both P. atlantica (possibly rungsi) and P. palaestina (possibly wertheimae).
Four or five nearctic species having the same ancient host association with Pistacia as Old World genera of Fordini. Four species (one un-named) form galls on Pistacia mexicana in Mexico, and are presumed to migrate to roots of various Compositae/Asteraceae, and also possibly to Ipomaea spp. Alatae have dark brown-black dorsal abdominal cross-bands. Muñoz Viveros & Remaudière (1999) provided descriptions and keys to galls on P. mexicana and all available morphs. The microscopic structure of the galls of three species was studied by Muñoz Viveros et al. (2014) and compared with other Fordini by Álvarez et al. (2016). As yet, however, it has not been possible to verify the complete life cycle of any species experimentally by transfer between primary and secondary hosts.
Geopemphigus blackmani Muñoz Viveros & Remaudière Galls on Pistacia mexicana in Mexico are large (maximum dimension up to 18 cm) lobulate, often at the apex of a branch. Emigrant alatae collected in June-August have dark dorsal abdominal cross-bands and dark wing veins; BL 1.6-2.2 mm. They are believed to migrate to roots of Compositae/Asteraceae (Tagetes, Dyssodia) where sexuparae believed to be of this species were found in September and December (Muñoz Viveros & Remaudière 1999). The presumed apterous exules are probably rather dark, with white wax; BL 1.6-1.9 mm. The alatae (sexuparae) have dark dorsal abdominal cross-bands and secondary rhinaria distributed ANT III 19-28, IV 2-4, V 0 (-1). The life cycle awaits experimental confirmation.
Geopemphigus floccosus (Moreira) Plate 2c Apterae according to the original description are pale dirty greenish yellow, with tufts of white wax; BL 1.9-2.2 mm. Alatae (virginoparae) have dark dorsal abdominal cross-bands, and secondary rhinaria distributed ANT III 14-20, IV 1-3, V 0-1, VI BASE 0. They occur on roots of Ipomaea (Convolvulaceae), and populations apparently of this species have also been found on Synedrella (Compositae/Asteraceae) and Dioscorea (Dioscoreaceae). In southern USA, Central America, the Caribbean and South America as far south as São Paulo, Brazil. Probably most populations are anholocyclic, as no alate sexuparae have been found, but there could perhaps be heteroecious holocyclic populations in regions where primary hosts are available. Emigrant alatae of “Geopemphigus sp. A”, described from galls on Pistacia mexicana in Mexico are very similar to alate virginoparae produced by root-feeding colonies of G. floccosus. The tubular galls of “Geopemphigus sp. A” are 15-35 mm long and 2.5-5.5 mm in diameter and very different from the galls of the three named species on P. mexicana. They produce emigrant alatae from June to August (Muñoz Viveros & Remaudière 1999). Sexuparae of sp. “A” collected from trunks of P. mexicana also agree closely in morphology with alate virginoparae of G. floccosus fom Ipomoea. Identity of G. floccosus and sp. “A” is therefore strongly suspected, but as yet attempts to make transfers between primary and secondary hosts have been unsuccessful.
Geopemphigus morral Muñoz Viveros & Remaudière Galls on Pistacia mexicana in Mexico are small (10-12 mm long and 5-8 mm deep), protruding from the underside of a leaflet. Emigrant alatae emerge in June to August. Possible secondary host populations and sexuparae have not yet been identified (Muñoz Viveros & Remaudière 1999).
Geopemphigus torsus Muñoz Viveros & Remaudière Galls on Pistacia mexicana in Mexico are elongate, spirally twisted, 3-8 cm long. Emigrant alatae collected in June have pigmentation similar to those of G blackmani; BL 1.6-2.1 mm. They are believed to migrate to form colonies on roots of Compositae/Asteraceae (Bidens odorata, Tagetes lunulata). Apterous exules on these plants suspected to be G. torsus are probably rather dark, with white wax; BL 1.6-2.7 mm. Alate sexuparae presumed to be this species are produced in September; they have dark dorsal abdominal cross-bands and secondary rhinaria distributed ANT III 17-28, IV 1-4, V 0-2, VI BASE 0 (-1). Sexuparae were also collected on trunks of P. mexicana in November (Muñoz Viveros & Remaudière 1999), but the life cycle still needs to be confirmed experimentally.
Two North American species forming galls on Ulmus. Closely related to Eriosoma, but without host alternation. The fundatrices have siphuncular pores, and the alate sexuparae produced in the galls have the secondary rhinaria on ANT III rather weakly developed and more widely-spaced than in alatae of Eriosoma (fig. 118A). Smith (1985) reviewed Georgiaphis as a subgenus of Eriosoma.
Georgiaphis gillettei Maxson & Hottes Forms leaf-curl galls on Ulmus americana and U. fulva (for illustration see Hottes & Frison 1931, as G. ulmi). Fundatrix and apterous fundatrigeniae are greenish blue, dusted with wax. Monoecious holocyclic on Ulmus. Alate sexuparae maturing in galls in June are greenish brown to greenish blue, wax-dusted; BL c. 1.5-1.6 mm) (Maxson & Hottes 1926). In eastern North America (Illinois, Iowa, North Carolina, Pennsylvania; BMNH collection). Synonymised with G. ulmi by Hottes & Frison (1931), but reinstated as a distinct species by Smith (1985). G. gillettei has R IV+V equal to or longer than HT II, whereas G. ulmi has R IV+V distinctly shorter than HT II.
Georgiaphis ulmi (Wilson) Gall is formed by curling one edge of the elm leaf into a spindle shape, 2-5 cm long. Galled leaves turn yellow, then red, then drop off (Wilson 1911). Fundatrix reddish-brown. Monoecious holocyclic, and apparently specific to Ulmus alata. Alate sexuparae, reddish brown with black head and thorax, are produced in second generation (Wilson 1911), leaving galls in May-June. In eastern North America.
One rather elongate-bodied Carex-feeding species found in both Asia and North America, perhaps most closely related to Byrsocryptoides, but with a much shorter rostrum.
Gharesia polunini Stroyan Apterae are greyish due to wax covering; BL 1.9-2.1 mm. Originally found on leaves of a plant of Cyperaceae, either Carex or Kobresia, in Pakistan. Subsequently recorded from California (Hille Ris Lambers 1966), and from British Columbia, Canada, where colonies in flocculent wax were found on stems and inflorescences of Carex mertensii (Foottit & Mackauer 1980). Alatae have narrowly transverse secondary rhinaria distributed III 10-12, IV 1-3, V 1-4, VI 1-2. Life cycle and origins unknown; possibly it is anholocyclic, and/or with a migration to Ulmaceae in its native region.
One fern-feeding aphid in Central America with swollen siphunculi, antennal tubercles with inner faces divergent, and apterae having a humped black dorsal abdominal sclerite.
Gibbomyzus pteridophytorum Nieto Nafría, Perez Hidalgo, Martínez-Torres & Villalobos Muller Apterae are pale green or brownish green with appendages mainly pale brown, and with an extensive shiny black dorsal abdominal patch, raised to give the appearance of a large rounded hump; BL 1.5-1.9 mm. Alatae have a dark quadrate dorsal abdominal patch. On ferns (Blechnum, Pteridium and probably other fern genera) in mountainous regions of Central America (Costa Rica, central Mexico; Nieto Nafría et al. 2013b). The life cycle is unknown.
A genus for one North American species collected on Philadelphus, with a distinctive form of cauda and antennal sensoriation.
Glendenningia philadelphi MacGillvray Apterae are undescribed; for a photograph of the extremely globose fundatrix see aphidtrek.org. BL of alatae is 2.2-2.8 mm. On Philadelphus gordonianus (= lewisii) in western USA and British Columbia, Canada. The life cycle is unknown; the fact that all individuals were alatae in June suggests that migration occurs to an unknown secondary host, probably a moss. (The form of the cauda is similar to that of aphids that feed on mosses).
Five species, three palaearctic and two nearctic. associated with Alnus and Betula. Blackman (1989) reviewed and keyed the palaearctic species, which feed on aerial shoots, whereas the two North American species are apparently both subterranean (Richards 1968a). The palaearctic species have abbreviated life cycles, with sexual morphs appearing in July (Wieczorek et al. 2014). Parasitized by Aphidius aquilus (= sicarius), which also attacks some other Betula-feeding genera such as Betulaphis and Calaphis (Starý 1973).
Glyphina betulae (Kaltenbach) Plate 4c, d Apterae are dark green to almost black with a pale spinal stripe (see influentialpoints.com/Gallery); BL 1.2-2.0 mm. Immature stages are green. Monoecious holocyclic, in colonies on young shoots of Betula spp., especially B. pendula, and occasionally on Alnus spp. Ant-attended. Throughout Europe, and across Asia to Korea (S. Lee et al. 2001a), Japan and the Kuril islands; introduced and now widespread in North America. The life cycle is abbreviated; detailed descriptions of the oviparae and dwarfish apterous males, present in July-August, were provided by Wieczorek et al. (2014). The male reproductive system was studied by Wieczorek & Świątek (2009). 2n = 28 or 56; the significance of the karyotype variation is uncertain (Blackman 1989).
Glyphina jacutensis Mordvilko ( = schrankiana Börner) Apterae are brown; BL 1.7-2.0 mm. Immatures are pale brown. Monoecious holocyclic, in colonies on young growth of Alnus spp. (incana, glutinosa), ant-attended. In central and northern Europe. Oviparae and dwarfish apterous males appear in July-August; the life cycle and sexual morphs were described by Wieczorek et al. (2014). 2n=8.
Glyphina longiseta Richards Apterae are reddish brown with conspicuous white or grey paired patches on abdominal tergites 1 and 6, and dark brown to black appendages; BL 1.9-2.1 mm. On an unidentified Alnus sp. in Nova Scotia, Canada. The feeding site is not described but is by implication the roots or underground stems (Richards 1968a). Life cycle unknown.
Glyphina pseudoschrankiana Blackman Apterae are brown-black with variable white markings (see influentialpoints.com/Gallery); BL 1.5-1.8 mm. Immature stages are brown. Monoecious holocyclic, in colonies on young shoots , often on young trees, of Betula pubescens and related downy birches, attended by ants (Blackman 1989). In northern Europe, Central Asia (Kazakhstan: Kadyrbekov 2011b) and Japan. Apterous males and oviparae occur in July-August; the sexual morphs were described by Wieczorek et al. (2014). 2n=10 [the record of 2n=12 in the legend to Fig. 1 of Blackman & Spence (1996) is a misprint].
Glyphina setosa MacGillivray Apterae dark, reddish brown to almost black; BL 1.7-2.2 mm. Immature stages are light yellow green in early instars, tan to brownish in later instars. Ant-attended, on roots and underground stems of Betula papyrifera seedlings in New Brunswick, Canada (MacGillivray 1963). Apterous males and oviparae were collected in August-September.
One species in east and south-east Asia, characterised by lack of frontal horns and by apterae bearing spine-like dorsal hairs, and with yeast-like extracellular symbionts indicating an affinity with Cerataphis and Tuberaphis (Fukatsu et al. 1994). Noordam (1991) provided a generic diagnosis.
Glyphinaphis bambusae van der Goot Apterae are small pale olive green to reddish or greenish brown, somewhat shiny aphids scattered on young shoots or at leaf bases of bamboos (Bambusa spp., Dendrocalamus latiflorus, Indocalamus tessellatus, Phyllostachys sulphurea, Pleioblastus amarus, Schizostachyum brachycladum, Shibataea spp.). Immatures are spotted (Moritsu 1983, p.213). Alatae (BL c. 1.9 mm) have a bluish black abdomen (Noordam 1991). Jiang et al. (2012b) described a population on Bambusa sp. in China with short dorsal and marginal hairs (see also Qiao et al. 2018). The life cycle is not recorded. In India (Assam), China, Japan, Singapore, Java and Sulawesi.
Two palaearctic species forming galls on Populus, related to Pachypappa but the fundatrices have wax glands on head, thorax and abdomen, and emigrant alatae have antennae with large star-shaped primary rhinaria. Danielsson (1990a) reviewed the genus.
Gootiella alba Shaposhnikov Forming conical leaf-galls on Populus alba in Russia (Uralsk), Kazakhstan and Poland. Pale yellowish green apterae (BL 3.3-4.6 mm) live in dense wax-covered colonies on the lower side of the leaf-cone. Alatae are unknown, suggesting monoecy (Shaposhnikov 1952). The uncertainties of the life cycle were discussed by Danielsson (1990a). If there is host alternation then Aploneura juniperina Juchnevitch (1970), described from roots of Juniperus sabina in Kazakhstan, could be the secondary host form.
Gootiella tremulae Tullgren Forming large pale green to yellowish bag-like galls from single leaves of Populus tremula, often situated rather high up in old trees and therefore hard to find. The gall is formed from a longitudinally folded leaf which is paler than the normal leaves and becomes much enlarged with a thickened petiole (fig. 131C). Emigrant alatae are greyish green, clothed with wax, BL 3.2-4.0 mm, which in Sweden leave galls in early July to found colonies on roots of Juniperus spp. (Danielsson 1990b). Apterous exules are globular, grey, living in nests of white wax; BL 0.9-1.5 mm. It is also recorded from roots of J. communis in Britain (Carter & Danielsson 1993), and there is an earlier British record from Kent (possibly of vagrants), on Picea abies (Wood-Baker 1980). Anholocyclic overwintering on juniper roots is probably common. In UK, France (BMNH collection, leg. G. Remaudière), Denmark, Scandinavia and Poland. 2n=16.
One species with large thoracic spiracles on Saussurea in east Asia, possibly related to Delphiniobium but with 5 hairs on first tarsal segments, and tapering siphunculi bearing hairs (Pashchenko 2000b).
Gredinia pilosotuba Pashchenko Apterae are black, shining: BL c.2.8 mm. Alatae are unknown. On shoot apices of Saussurea pulchella in east Siberia (Primorskii). Life cycle is unknown. Except for the large thoracic spiracles the description of this species, from two specimens, closely matches that of Macrosiphoniella lazoica, found on the same host.
About 50 east Asian species with a pattern of pale reticulation on the densely hairy siphunculi. About half the species fall into subgenus Trichosiphum, which have apterae in which this reticulation is confined to a region at the base of each siphunculus. Biology and host relations of the genus were discussed by A.K. Ghosh (1987). They live mainly on the shoots and young foliage of trees, particularly Fagaceae but also Moraceae, Betulaceae, Juglandaceae, and less commonly on members of evolutionarily more recent families such as Myrtaceae, Rosaceae and Rubiaceae. In the few species for which life cycle information is available the sexuales occur in spring or summer, both sexes are winged and eggs are somewhat flattened. Several of the most common and widespread species seem to be mainly or entirely anholocyclic. Raychaudhuri (1956) reviewed the world fauna, and more recent accounts are available for Japan (Sugimoto 2008), India (A.K. Ghosh & Agarwala 1993), China (G. Zhang & Zhong 1979b), Taiwan (Liao 1978, Tao 1962), Java (Noordam 1994) and the species introduced to North America (Halbert 2004). D. Zhang & Qiao (2007a) reviewed the Greenidea subgenus Paragreenidea. Q. Liu et al. (2013) studied the use of DNA barcoding for identification of Greenidea species and other Greenideinae in China; several of their samples were new host or distribution records for little-known species and their identity may need further verification. Q. Liu et al. (2015 used both mitochondrial and nuclear DNA to investigate the evolution of host relationships in Greenidea and related genera.
Greenidea (Trichosiphum) anonae (Pergande) Apterae are pear-shaped with yellow-brown head and thorax and dark brown abdomen; BL 1.3-1.8 mm. Siphunculi are black, curved outwards distally, 0.18-0.25 × BL in apterae and 0.5-0.6 × BL in alatae. On undersides of young leaves of Annonaceae (Annona, Artabotrys, Polyalthia) and also recorded from Antidesma, ?Baccauria, Pittosporum and Symplocos. A record from Quercus sp. in India (Chakrabarti & Raychaudhuri 1975) needs confirmation. It is sometimes ant-attended. A single alate male is known, collected in central India in February; probably most populations are anholocyclic. In India, Java, Malaya and Sumatra. Records from Japan seem to be erroneous (Sugimoto 2008). 2n=22 (Khuda-Bukhsh & Kar 1990).
Greenidea artocarpi (Westwood) Apterae are rather elongate pear-shaped, shiny brown or greenish with diffuse darker dorsal markings (see aphids of Karnataka website); BL 2.1-2.4 mm. Siphunculi are dark brown to black, long and slender; about 0.5 × BL in apterae and 0.8 × BL in alatae. On undersides of young leaves, usually near the main trunk, of Artocarpus spp. (heterophylla, incisa) in southern India and Sri Lanka (David 1956b, Raychaudhuri 1956). Records from south-east Asia should probably be referred to G. ficicola. The life cycle is unknown.
Greenidea ayyari Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae are pear-shaped, brown; BL 2.9-3.2 mm. Siphunculi are pale with darker apices, c. 0.3 × BL in apterae and 0.70-0.76 × BL in alatae (A.K. Ghosh & Agarwala 1993). Known only from one aptera and two immatures on a Quercus sp. in India (Manipur), and an ovipara on Lyonia sp. (the latter presumably a “casual” visit).
Greenidea brideliae Takahashi Apterae are elongate pear-shaped, yellowish green with dusky tips to appendages; BL 1.5-2.5 mm. Siphunculi are rather pale, darker at tips, about 0.6 × BL in apterae and 0.8 × BL in alatae. On Bridelia spp. (monoica, tomentosa) in Taiwan, Hong Kong and eastern China. Records from Japan seem to be erroneous (Sugimoto 2008). Biology is unknown.
Greenidea (Trichosiphum) bucktonis A.K. Ghosh, Basu & Raychaudhuri Apterae are pear-shaped, colour in life unrecorded; BL 1.6-2.5 mm. Siphunculi are 0.2-0.3 × BL in apterae and 0.5-0.7 × BL in alatae. On Syzygium and Eugenia spp. in northern India (Uttar Pradesh, Assam, Sikkim, Meghalaya, West Bengal). There are also records from Oreocnide (= Villebrunnea) integrifolia, Duabanga sonneratioides, Sapindus sp. and Prunus sp. (A.K. Ghosh & Agarwala 1993, Singh & Singh 2017c), but these are less likely to be true hosts. Biology is unknown.
Greenidea (Trichosiphum) camelliae Agarwala & Ghosh Apterae are pear-shaped,brown with yellowish head; BL 1.2-1.5 mm. Siphunculi are 0.25-0.31 × BL in apterae, alatae undescribed. On Camellia sp. in Meghalaya, India (A.K. Ghosh & Agarwala 1993). Very similar to G. (T.) bucktonis, and possibly small individuals of that species on a “casual” host.
Greenidea carpini Takahashi (= carpinicola Banerjee & Chakrabarti) Apterae are pear-shaped, blackish brown with mainly black appendages; BL 1.6-3.1 mm. Siphunculi are 0.19-0.27 × BL in apterae and about 0.6 × BL in alatae. Sugimoto (2008) provided a redescription of the aptera, ovipara and male and a first description of the alate vivipara, from all of which it seems clear that G. carpinicola is a synonym. On Carpinus spp. in Japan, and also in northern India (Uttar Pradesh, as carpinicola). Alate oviparae and males were collected in Japan in May (Takahashi 1963b). The male genitalia were described and illustrated by Wieczorek et al. (2012).
Greenidea (Paragreenidea) cayratiae Qiao & Zhang Apterae are pear-shaped, reddish brown; BL 2.5-2.8 mm. Siphunculi are dark, c.0.5 × BL in apterae and about c.0.8 × BL in alatae. On young leaves of Cayratia japonica, attended by ants. In Fujian Province, China (D. Zhang & Qiao 2007a).
Greenidea decaspermi Takahashi Apterae are pear-shaped, yellowish brown with dorsal abdomen dark brown to black; BL 1.7-2.0 mm. Siphunculi are brownish black with tips paler, 0.39-0.57 × BL in apterae and c.0.75 × BL in alatae (Raychaudhuri 1956). Described from Decaspermum fruticosum in Taiwan (unlikely to be the true host), and David et al. (1969) recorded it from Psidium guajava in India, but this may have been G. ficicola. There is also an Indian record from a Quercus sp. (A.K. Ghosh & Agarwala 1993).
Greenidea (Trichosiphum) eugeniae Takahashi Apterae pear-shaped , dark brown to black in life with paler head and spinal region, pale antennae and legs and dark siphunculi; BL c.1.6 mm. Siphunculi are c. 0.25 × BL in aptera. Y. Lee et al. (2016) have provided morphometric data and excellent illustrations of apterae. The alata is undescribed. On Syzygium zeylanicum (= Eugenia spicata) in Sumatra (Takahashi 1941b) and on unidentified species of Myrtaceae in Laos (Y. Lee et al. 2016).
Greenidea ficicola Takahashi Apterae are pear-shaped, yellowish brown to dark brown (see aphids of Karnataka website); BL 1.7-2.5 mm. Siphunculi are yellowish brown to dark brown, curved outwards distally, 0.29-0.42 × BL in apterae and 0.55-0.76 × BL in alatae. On Ficus spp., on undersides of young leaves and on shoots, or concentrated on the fruits. Also recorded in India from Psidium guajava, Duabanga sonneratioides and Litchi chinensis (A.K. Ghosh & Agarwala 1993), and from Java on Streblus elongatus (Noordam 1994). In east and south-east Asia, Australia, Africa (Burundi), and in recent years spreading to South America (Peru, Rubín de Celis et al. 2006; Brazil, Sousa-Silva et al. 2005; Argentina, La Rossa et al. 2011), Mexico (Salas-Araiza et al. 2011), southern USA (Florida; Halbert 2004), Hawaii (Nagamine & Garcia 2012), southern Europe (Spain, Pérez Hidalgo et al. 2009c; Italy, Barbagallo et al. 2005a; Malta, Mifsud 2008; Madeira, Aguiar et al. 2013) and North Africa (Ben Halima-Kamel 2009; Laamari et al. 2013). Bella et al. (2009) discussed its rapid spread around the world. Introduced populations are recorded mainly from ornamental Ficus spp. Its biology is little known; anholocyclic in Taiwan (Takahashi 1923), and no sexual morphs are known. Sometimes ant-attended. New parasitoids, including a new braconid genus (Indoephedrus) have been described from this aphid (Samanta et al. 1993, as G. neoficicola, and Samanta 1985). 2n=22.
Greenidea (Trichosiphum) flacourtiae van der Goot Apterae are pear-shaped, with pale brown head and shiny brown abdomen; BL 1.4-2.4 mm. Siphunculi are dark brown to black, 0.33-0.38 × BL in apterae, 0.73-0.90 × BL in alatae. On youngest parts of developing shoots of Distylium stellare in Java. Other morphs and biology are unknown (Noordam 1994). However, specimens collected from Castanopsis sp. in Huangdong province, China have been identified as this species and used for DNA studies (Q. Liu et al. 2013).
Greenidea (Trichosiphum) fulva Noordam Apterae are pear-shaped, yellow, pale brown or dirty whitish, with antennae and legs pale except at extremities; BL 1.6-2.1 mm. Immatures are whitish with abdomen very pale green). Siphunculi of apterae are pale brown or orange brown with black tips, 0.39-0.47 × BL, siphunculi of alatae are black and 0.56-0.82 × BL. On Bridelia monoica in Java, using the long narrow R IV+V to feed in the interstices between developing shoots (Noordam 1994). The life cycle is unknown.
Greenidea (Trichosiphum) haldari Maity & Chakrabarti Apterae are pear-shaped, blackish, with siphunculi pale to brown, short, stout and outwardly curved, only 0.14-0.16 × BL, which is 2.0-2.4 mm. On Quercus incana in Western Himalaya, India, in stipules and on undersides of young leaves, attended by ants. Also recorded from Bhutan (Chakrabarti & Das 2014). The alate morph was described by Chakrabarti & Debnath (2009).
Greenidea (Trichosiphum) heeri Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae are pear-shaped, shiny yellowish brown with darker brown dorsal abdominal markings; BL (1.3-) 1.6-2.5 mm. Siphunculi are brown, 0.2-0.3 × BL in apterae, 0.66-0.72 × BL in alatae. On undersides of younger leaves and new stem growth of Myrtaceae, especially Syzygium spp., and also recorded from Engelhardtia spicata (possibly a “reserve” host). In east and south-east Asia, as far west as Pakistan, where Naumann-Etienne & Remaudière (1995) recorded alate oviparae (as G. formosana) collected in spring at Islamabad on Syzygium cumini. This taxon, originally described as a subspecies of G. formosana, has been synonymised with G. psidii (q.v.) by some authorities, but is here treated as a distinct species with a shorter RIV+V, and possibly a different host association, pending further investigation. Agarwala (1983) recorded observations of a population identified as G. formosana (= psidii) ssp. heeri on guava in Bhutan, but Psidium is not listed as a host of heeri in A.K. Ghosh & Agarwala (1993), so the identity is in doubt. G. nigricans Noordam, dscribed from Syzygium spp. in Java (Noordam 1994) may be small specimens of G. heeri. Kurl (1986) recorded chromosome numbers of 7, 8 and 9 for various populations identified as heeri.
Greenidea (Trichosiphum) isensis Sorin & Agarwala Apterae are pear-shaped, reddish brown, with blackish brown spots or patches on antennae and legs; BL 2.7-3.2 mm. Siphunculi are dark brown to blackish brown, 0.44-0.53 × BL in apterae and 0.65-0.69 × BL in alatae. On Quercus gilva in Japan (Sorin & Agarwala 2002).
Greenidea kheemothicola Lee & Lee Apterae are elongate-bodied, shiny pale yellow to greenish yellow; BL 2.5-2.9 mm. SIPH pale with darker tips, 0.63-0.77 × BL. Alatae are unknown. On undersides of leaves of Glochidion fagifolium (= G. hohenackeri) in Laos (Y. Lee et al. 2016).
Greenidea (Trichosiphum) kuwanai (Pergande) Apterae are broadly pear-shaped, with head, prothorax and appendages shiny mid- to dark brown; BL 2.2-2.6 mm. Siphunculi are blackish brown, 0.20-0.36 × BL in apterae and 0.43-0.60 × BL in alatae. Immatures have a blue-grey bloom dorsally. On young shoots and leaves of Quercus spp., with possibly valid records also from Lithocarpus spp. and Castanea spp. Records from Castanopsis (= Shiia) are likely to be G. okajimai and, according to Takahashi (1962b), Raychaudhuri's (1956) description is partly that species. In east Siberia, Japan, Korea and Taiwan. Alate oviparae occur in Japan in November-December (Takahashi 1923), but males have not been recorded. G. brachyunguis, described from Quercus sp. in Sikkim, India (Chatterjee, Mondal & Raychaudhuri 1981), and G. hangnigra, described from Quercus sp. in China (Chang & Zhong 1979b) are very similar to and likely to be synonymous with G. kuwanai. 2n=20 (Blackman 1986).
Greenidea longicornis M. R. Ghosh, A.K. Ghosh & Raychaudhuri Apterae are elongate, pale yellow; BL 2.5-3.0 mm. Siphunculi are pale with black apices, 0.71-0.87 × BL in apterae, c.1.1× BL in alatae. Recorded from Engelhardtia sp. in West Bengal and from Cassia sp. in Meghalaya, India, but neither of these may be the true host.
Greenidea longirostris A.N. Basu Apterae are pear-shaped, whitish brown or yellowish brown; BL 2.3-2.7 mm. Siphunculi are pale brown, 0.43-0.56 × BL in apterae, 0.60-0.75 × BL in alatae (A.N. Basu 1970). On Schima wallichii in northern India (Manipur, Meghalaya, West Bengal). Very similar to G. schimae on the same host, but with a somewhat longer R IV+V; its separate species status needs to be verified.
Greenidea longisetosa Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae mainly brown, pear-shaped; BL 2.8-3.1 mm. Siphunculi dark brown to black with pale apices, 0.23-0.26 × BL. Alatae are unknown. Described from an unidentified plant in West Bengal, India, with a subsequent record from Ixora sp. (see Singh & Singh 2017c), but this is unlikely to be the normal host plant..
Greenidea maculata Noordam Apterae are elongate pear-shaped, head and border of body orange-brown, with a large brown central dorsal abdominal patch; BL c. 2.3 mm. Siphunculi are black except at base, about 0.8 × BL (known from only one specimen). On a young shoot with developing leaves of Syzygium antisepticum in Java (Noordam 1994).
Greenidea magna Noordam Apterae are elongate pear-shaped, pale green, green or brownish, the head paler, with a brown spot on the posterior abdomen; BL 2.4-3.4 mm. Femora are whitish, contrasting with dark tibiae and tarsi. Siphunculi are pale brown or brown with black apices, 0.37-0.57 × BL in apterae, darker and 0.83-0.95 × BL in alatae. On undersides of leaves and young shoots of Castanopsis spp. and Castanea crenata in Java (Noordam 1994).
Greenidea mangiferae Takahashi Apterae are pear-shaped, yellowish brown to dark brown; BL c. 2.0 mm. SIPH are black, very long; almost 0.5 × BL in apterae, and 0.75 × BL in alatae. On Mangifera indica and Euphoria longana (= Dimocarpus longan) in Taiwan (Takahashi 1925, 1931), and China. Biology is unknown. 2n=20 (Chen and Zhang 1985).
Greenidea (Trichosiphum) manii Ghosh, Basu & Raychaudhuri Apterae pale yellow; BL 2.4-2.5 mm. SIPH are pale, c.0.3 × BL in apterae, alatae are undescribed. Described from an unidentified plant in India (Meghalaya), and there are subsequent records from Alnus nepalensis and Quercus sp. (Singh & Singh 2017c). It could possibly be a synonym of G. (T.) myricae, which is longer haired but otherwise very similar.
Greenidea mushana Takahashi Only the alate vivipara of this species is described, distinguished by having more numerous secondary rhinaria on ANT III (about 55) than any other known Greenidea species. Alata is blackish brown with long black SIPH, about 0.7 × BL. On young shoots of Quercus sp. in Taiwan (Takahashi 1925).
Greenidea (Trichosiphum) myricae Takahashi Apterae are pear-shaped, yellowish green or green with tips of siphunculi dark brown; BL 1.9-2.5 mm. Siphunculi are 0.34-0.46 × BL in apterae and 0.72-0.83 × BL in alatae. Described from specimens feeding on a young leaf of Myrica rubra in Taiwan (Takahashi 1925), and collected several times from this plant in Japan (Sugimoto 2008), indicating that it is a true host. However, aphids collected on Alnus formosana two years before the original description were later identified by Takahashi as this species (Raychaudhuri 1956; specimens now in BMNH collection), Tao (1999) also records it from Alnus cremastogyne, and in Japan it has been collected from Quercus phillyraeoides (Sugimoto 2008).
Greenidea (Trichosiphum) nigra Maki Apterae are pear-shaped, brown to dark brown-black; BL 1.6-3.6 mm. Siphunculi are black, about 0.27-0.37 × BL in apterae and 0.58-0.68 × BL in alatae. On evergreen Quercus spp. and Lithocarpus spp. in Taiwan (Liao 1978) and Japan. Sugimoto 2008) gave a full redescrition of this species from Japan including the male and ovipara, and compared it with G. kuwanai, with which it has often been confused in the literature. Takahashi (1937b) described specimens with finer hairs as a "variety", G. nigra kanzanensis.
Greenidea (Trichosiphum) nipponica (Suenaga) Apterae are very broadly pear-shaped, with abdomen dorsoventrally flattened, almost circular, black suffused with red posteriorly (Moritsu 1983); BL 1.9-2.5 mm. SIPH are dark, 0.22-0.32 × BL in apterae and 0.47-0.62 × BL in alatae. On young growth of Quercus spp. in Japan and Korea, regularly attended by ants. In Japan, alate oviparae and males are produced in autumn and winter (Takahashi & Sorin 1959, Sugimoto 2008), and fundatrices were found in April (Sugimoto). Also recorded from Pakistan, but there the sexual morphs were found on three species of Quercus in summer, and also black sclerotic apterous viviparae were collected on buds at 2700m in winter, persisting mixed with pale-coloured spring colonies until May (Naumann-Etienne & Remaudière 1995). G. (T.) kumaoni Chakrabarti & Raychaudhuri, on Quercus sp. in Uttar Pradesh, India (Chakrabarti & Raychaudhuri 1978) is apparently very similar if not synonymous, differing only in that some of the dorsal hairs are blunt or bifurcate. 2n=18 (Blackman 1986).
Greenidea (Trichosiphum) okajimai Suenaga Apterae are broadly pear-shaped, dark brown dorsally with yellow-brown head, margins of body and legs (Moritsu 1983); BL 2.0-2.8 mm. Siphunculi are brown or blackish, about 0.24-0.37 × BL in aptera and 0.63-0.79 × BL in alata. On Castanopsis (= Shiia) cuspidata and C. sieboldii in Japan, with one collection from Quercus glauca var. amamiana (Sugimoto 2008). The life cycle is abbreviated; alate oviparae and males start appearing in late May, in the second generation after the fundatrix (Takahashi & Sorin 1959, Sugimoto 2008). There is a record from West Bengal (A.K. Ghosh et al. 1971e), but this needs to be substantiated as it is not mentioned by A.K. Ghosh & Agarwala (1993). G. prunicola Ghosh, Banerjee & Raychaudhuri, described from Prunus sp. in Sikkim (A.K. Ghosh et al. 1971c), is probably vagrant okajimai, although specimens collected on Castanopsis sp. and Cyclobalanopsis sp. in China and used in DNA barcoding and sequencing studies were identified as this species (Q. Liu et al. 2013, 2015). G. kunmingensis Chang, described from Castanopsis spp. in China (Chang & Zhong 1979b), is likely to be a synonym, as also is G. castanopsidis Noordam, described from Java (Noordam (1994). The latter name has also been applied to specimens collected from unidentified hosts in China and used for DNA barcoding and sequencing (Q. Liu et al. 2013, 2015).
Greenidea (Trichosiphum) pallidipes Noordam Apterae are broadly pear-shaped, head and margins of body pale brown, central area of abdomen shiny dark brown, and legs pale brown except for apices of tarsi; BL 1.3-2.2 mm. Siphunculi are brown, darker distally, 0.25-0.38 × BL in aptera and c.0.8 × BL in alata. Described from Java on Castanopsis spp., Castanea crenata and Quercus glauca, living on young parts of developing shoots and sometimes also on the bark of older branches (Noordam 1994). The description does not distinguish this insect adequately from the Indian species G. quercicola, and it may therefore be a synonym.
Greenidea (Paragreenidea) parthenocissi Saha & Chakrabarti Apterae are greenish, BL 2.1-2.7 mm. On undersides of leaves of Parthenocissus semicordata in north-east India (Uttar Pradesh). Alate oviparae and alate males were collected in September-October (original description).
Greenidea photiniphaga Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae are brownish; BL c.2.5 mm. Siphunculi pale, darker at apices, c.0.5 × BL in aptera, alata unknown. On a Photinia sp. (probably not its true host) in West Bengal, India (Raychudhuri et al. 1973).
Greenidea (Trichosiphum) prinicola Sugimoto Apterae are pear-shaped, pale brown to brown; BL 1.9-2.7 mm. Siphunculi of aptera are brown, 0.23-0.29 × BL, of alatae mainly black, 0.47-0.56 × BL. On Quercus spp. of the section Prinus (Q. dentata, Q. serrata) in Japan. Fundatrices were collected in April-May, and alate oviparae and males in October-November (Sugimoto 2008). This species was previously confused in the literature with G. nipponica.
Greenidea (Trichosiphum) psidii van der Goot (= formosana Maki) Apterae are pear-shaped, shiny dark reddish or yellowish brown; BL 1.5-2.4 mm. Siphunculi are yellowish brown, darker at base and apex, curved outwards distally, 0.26-0.42 × BL in apterae and 0.66-0.77 × BL in alatae. On young shoots and undersides of young leaves of Psidium guajava and other Myrtaceae (Rhodomyrtus, Eugenia, Melaleuca, Plinia). Halbert (2004) established the priority of the name psidii over the name formosana that has been widely applied to this aphid. The taxonomy of the psidii group requires clarification. Possibly G. psidii s. str. restricts its feeding to Myrtaceae, but similar aphids occur on other plants such as Ficus and Engelhardtia. Records of G. psidii on Ficus spp. in Taiwan (Liao 1978) are possibly G. fici Takahashi, described from eastern China (Takahashi 1937e). G. guangzhouensis Chang, also described from Ficus in China and very similar to G. psidii, may also be G. fici. G. fici is not, however, adequately differentiated from psidii in Takahashi's description. Aphids identified as G. psidii occur in India, Pakistan, Bangladesh. Nepal, eastern China, Taiwan, Japan, Philippines, Loochoo, Java and Sumatra, and have been introduced to Hawaii (1993), California (1998; see Halbert 2004), Brazil (Lazzari et al. 2006), Costa Rica (Pérez Hidalgo et al. 2009a), Mexico (Salas-Araiza et al. 2011), Panama and Venezuela (Cermeli et al. 2012). Reports of G. ficicola on Psidium in Colombia (R.D. David et al. 2009) should also perhaps be referred to this species. Sexual morphs are unrecorded; parthenogenetic morphs occur all-year-round in Taiwan (Takahashi 1923). Takahashi (1924a) described the immature stages. Indian populations on guava in the BMNH collection tend to have a shorter R IV+V than those in south-east Asia (1.6-2.0 × HT II, as opposed to 1.9-2.2 × HT II). G. psidii-like aphids on Syzygium spp. and Engelhardtia spicata in India with an even shorter R IV+V (1.2-1.75 × HT II) are here regarded as a distinct species, G. heeri. Chromosome number is 2n=18 according to Kulkarni & Kacker (1979).
Greenidea (Trichosiphum) quercicola Basu, Ghosh & Raychaudhuri Apterae are pear-shaped, colour in life unrecorded; BL c. 1.9 mm. Siphunculi of aptera are probably rather pale, about 0.24 × BL, of alata c. 0.63 × BL. On Quercus dealbata (= floribunda?) and Q. serrata in Meghalaya, India (R.C. Basu et al. 1975). Other morphs and biology are unknown.
Greenidea quercifoliae Takahashi Apterae are pear-shaped, greenish yellow, slightly dusky on dorsal abdomen, with legs yellowish brown except for dark apices (Takahashi 1921); BL c. 1.8 mm. Siphunculi are blackish, about 0.45 × BL in aptera and 0.73 × BL in alata. On young leaves of Quercus spp. in Taiwan; apparently rather uncommon, and biology is unknown.
Greenidea querciphaga Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae are pear-shaped, brown, probably with a darker abdominal dorsal patch; BL 3.6-3.8 mm. Siphunculi are long and pale with darker tips, 0.38-0.42 × BL in aptera and c. 0.76 × BL in alata. On an unidentified Quercus sp. in West Bengal, India (Raychaudhuri et al. 1973, and also recorded from Nepal (A.K. Ghosh & Agarwala 1993). Biology and sexuales are unknown.
Greenidea rappardi Raychaudhuri Apterae are shining brown, with paler brown head, thorax and band across between siphunculi; BL 1.6-1.9 mm. Siphunculi are 0.36-0.43 × BL in aptera, about 0.73 × BL in alata. On young shoots, undersides of young leaves and flower-buds of Scurrula and Loranthus sp. in Java (Noordam 1994).
Greenidea schimae Takahashi Apterae are elongate pear-shaped, head pale brown, body dull green with a somewhat reddish hue; BL 2.3-3.5 mm. Siphunculi are 0.44-0.55 × BL and yellowish brown in aptera, about 0.77-0.82 × BL and dark brown in alata (Noordam, 1994). On young leaves of Schima spp. in Taiwan and Java (Raychaudhuri 1956).
Greenidea (Trichosiphum) siamensis Takahashi Apterae are greenish black, with black siphunculi; BL 1.3-1.7 mm. from Rhodomyrtus sp. (?) in Thailand (as G. anonae ssp. siamensis), and given full species status by Robinson (1972).
Greenidea (Trichosiphum) sikkimensis Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae are pear-shaped, dark brown; BL 1.5-2.1 mm. Siphunculi of aptera are short and stumpy, only 0.15-0.25 × BL, of alata c. 0.4 × BL (A.K. Ghosh & Agarwala 1993). On an unidentified Quercus sp. and an unidentified Castanopsis sp. in India (Sikkim, Meghalaya).
Greenidea sutepensis Takahashi Apterae are elongate pear-shaped, pale; BL 2.2-2.4 mm. Siphunculi are pale brownish, about 0.55 × BL in aptera, alata are not known. Described from an unidentified tree, possibly a Dipterocarpus sp., in Thailand (Takahashi 1941, Raychaudhuri 1956).
Greenidea (Paragreenidea) symplocosis A.K. Ghosh, Basu & Raychaudhuri Apterae are elongate pear-shaped, pale (colour in life unrecorded); BL 2.1-3.3 mm. Siphunculi are long and pale, c. 0.6-0.75 × BL in aptera, c. 0.78 × BL in alata. On Symplocos spicata (= laurina), but with a record also (vagrants?) from Eurya sp. (A.K. Ghosh & Agarwala 1993). In India (Meghalaya, Sikkim, West Bengal). Some specimens collected in Taiwan on Quercus glandulifera (= serrata) and used for barcoding were identified as this species. Other morphs and life cycle are unknown (A.K. Ghosh et al. 1969a). The differences between this species and the preceding two described by Takahashi (schimae, sutepensis) are not very clear.
Greenidea (Paragreenidea) viticola Takahashi Apterae are brownish yellow, sometimes dusky on dorsum, with dark antennae and siphunculi; BL 2.6-3.0 mm. On Vitis (= Ampelopsis) cantoniensis in Taiwan.
About 12 species in east and south-east Asia described from members of taxonomically diverse plant families. Host plants of several species, including two from the Philippines (see Raychaudhuri 1972), are unknown. Greenideoida are rather elongate aphids distinguished from other Greenideinae by the short last rostral segment with no clear division between R IV and R V. The siphunculi have no reticulation and are curved abruptly outwards near their apices. Alatae where known are distinctive, having forewings with an elongate, dark pterostigma and the media only once-branched, and hind wings much reduced without oblique veins (except in subgenus Neogreenideoida). Raychaudhuri (1956) revised the species then known, and there have been more recent revisions of the species in India (A.K. Ghosh & Agarwala 1993 and Java (Noordam 1994). Subgenus Pentatrichosiphum was reviewed (as a full genus) by D. Zhang & Qiao (2007b).
Greenideoida bhalukpongensis A.K. Ghosh, Banerjee & Raychaudhuri Apterae are elongate-bodied, dark brown; BL 1.4-2.1 mm. Siphunculi are dark brown, curved outwards, 0.27-0.53 × BL in aptera and c. 0.6 × BL in alata. Described from Uvaria sp. and an unidentified plant, but subsequently recorded from Mesua ferrea by A.K. Ghosh & Agarwala (1993), who summarise the differences from G. ceyloniae. In India (Arunachal Pradesh, Assam, Tripura). The life cycle is unknown.
Greenideoida ceyloniae van der Goot Apterae are elongate spindle-shaped, of variable colour, yellow, greenish yellow or dark green (Takahashi 1950; as mesuae); BL 1.8-2.2 mm. Alatae are yellowish with a black head and thorax and a large black dorsal abdominal patch. Siphunculi are very long, dusky to dark but paler at apices; 0.52-0.76 × BL in aptera and 0.7-0.9 × BL in alata. On young leaves and branches of Mesua ferrea in India, Sri Lanka and Malaya. Alate oviparae and alate males occur in March in India (A.K. Ghosh 1987). Agarwala & Roy (1987) studied seasonal changes in the reproductive activity of the apterous viviparae.
Greenideoida elongata van der Goot Apterae are very elongate-bodied, whitish to greenish white; BL 1.6-2.3 mm. Alatae have black head and thorax and a large grey-black patch on the bright green abdomen. Siphunculi of aptera are pale brown with darker apices, 0.50-0.74 × BL; of alata 0.67-0.87 × BL. On Bridelia tomentosa, feeding on lower sides of leaves along the midrib, usually attended by ants. In Java, Malaya and Sumatra (Raychaudhuri 1956). An alate ovipara was collected in June in Malaya, otherwise the life cycle is unknown.
Greenideoida fransseni Noordam Apterae are elongate-bodied, with whitish or colourless head and appendages (antennae ringed with black), and pale yellow body; BL 1.5-1.9 mm. Siphunculi of aptera are 0.41-0.47 × BL, alatae are undescribed. On Bridelia tomentosa, living separately on the leaves (Noordam 1994). In Java, collected in 1932, and unknown since.
Greenideoida hannae van der Goot Apterae are elongate-bodied, whitish or pale greenish white; BL c. 2.35 mm. Siphunculi are rather short; about 0.3 × BL. On young succulent shoots of a shrubby Streblus sp. in Java (van der Goot 1917), unknown since the original collection. Other morphs and biology are unknown. Raychaudhuri (1956, 1972) expressed doubts about whether this species belongs in Greenideoida.
Greenideoida (Pentatrichosiphum) lambersi Basu Apterae are elongate-bodied, pale glassy white; BL 2.0-2.5 mm. Siphunculi of aptera are brown, 0.24-0.30 × BL. Found on lower sides of leaves of Hymenodictyon sp. at an altitude of 2000 m in West Bengal, India (A.N. Basu 1964). Raychaudhuri & Chatterjee (1977), and also A.K. Ghosh & Agarwala (1993), subsequently described a possible alata of this species collected on Premna sp., but alatae with the same collection data were ascribed to G. noonadanae by A.K. Ghosh et al. (1971e). The life cycle is unknown.
Greenideoida (Pentatrichosiphum) lutea (Basu) Apterae are elongate-bodied, probably pale; BL 2.7-3.2 mm. Siphunculi of apterae are pale brown with darker apices, 0.44-0.57 × BL, of alatae dark brown, 0.55-0.65 × BL. On undersides of leaves of Litsea spp., and also recorded from Lindera sp., and (presumably vagrants) from Cestrum fasciculatum and Buddleja sp. In India (Arunachal Pradesh, Meghalaya, Sikkim, West Bengal), Bhutan and Indonesia (A.K. Ghosh & Agarwala 1993), and also recorded from Litsea cubeba in China (Hainan; Q. Liu et al. 2013). The biology and life cycle are unknown.
Greenideoida noonadanae Heie Only the alate morph is known, described from the Philippines, host and appearance in life unknown (Heie 1967). Two blackish alatae collected on Premna sp. in West Bengal were identified as this species by A.K. Ghosh et al. (1971e), although material with the same collection data has subsequently been described as the alata of G. lambersi, see above.
Greenideoida (Neogreenideoida) philippensis Raychaudhuri Apterae are elongate-bodied, colour in life unrecorded, probably pale; BL c. 2.3 mm. Siphunculi are pale and about 0.7 × BL in aptera, dark and about 0.9 × BL in alata. On "Crotorylon celebicum" (presumably Cratoxylum celebicum) in the Philippines (Raychaudhuri 1956). The biology and life cycle are unknown.
A genus for one North American species with distinctive wax plates in the fundatrix (fig. 47B), oval secondary rhinaria in alatae and first instar exules with thickened hind femora.
Grylloprociphilus imbricator (Fitch) (= frosti Smith & Pepper) Host-alternating between Fagus grandifolia and roots of Taxodium distichum in eastern USA. Fundatrices develop on undersides of leaves of Fagus, moving to petioles and stems on maturity to form large colonies with flocculent wax, which can persist through the summer. Aoki et al. (2001) demonstrated that the slender long-legged 4th instar nymphs defended the colony by attacking predators. In North Carolina, alate emigrants become adult on Fagus from April to November. Apterous exules occur on roots of Taxodium throughout the year, indicating partial anholocycly. Sexuparae fly from November-February (Smith & Denmark 1984, Denmark & Smith 1986).
A genus for one North American species on Lonicera related to the palaearctic genera Hyadaphis and Semiaphis, but with siphunculi reduced to pores, a rounded cauda, and a full set of marginal abdominal tubercles.
Gypsoaphis oestlundi Hottes Plate 13f (Fig.38b) Apterae are green, wax-covered,: BL 1.5-3.3 mm. Alatae have secondary rhinaria on ANT III and IV. On young shoots and leaves of Lonicera spp. in North America (Hottes & Frison 1931). The life cycle is unknown. 2n=4.