The Aphids


(in alphabetical order)






































Nasonovia Mordvilko

Aphidinae: Macrosiphini

About 45 species with enlarged spiracular openings, cylindrical siphunculi, and apterae with sensoria on ANT III.  About two‑thirds of the species are nearctic; nearly half are associated with the related families Saxifragaceae and Grossulariaceae, host‑alternating species having Ribes as primary hosts.  The Old World subgenus Nasonovia s.str. is largely associated with Compositae and particularly Hieracium.  There are species on Ranunculaceae in both Old and New Worlds and species‑groups on Polemoniaceae and Orobanchaceae in North America.  Heie (1979) revised the world fauna, and Heie (1994) and Blackman (2010) reviewed the species in western Europe. Kadyrbekov (1995b, 2005a) reviewed the species in Kazakhstan, and synonymised subgen. Eokakimia Heie with subgen. Aconitaphis Ivanoskaya.

Nasonovia (Aconitaphis) alativica Kadyrbekov  (Fig.6g)  Apterae are dingy green or pale brown, with red eyes, antennae and legs dingy yellow with apices of segments dark brown, siphunculi and cauda dark brown; BL 2.5-3.2 mm. On flowers and flower stems of Aconitum spp., often forming large colonies, in Kazakhstan (Kadyrbekov 1995b). Subsequently it has also been recorded from Delphinium spp. Heteroecious, with Ribes heterotrichum as the primary host (Kadyrbekov 2014d); the primary host generations are undescribed. Very similar to, and possibly synonymous with, N. salebrosa.

Nasonovia (Kakimia) alpina (Gillette & Palmer)    Apterae are pale green, sometimes with dark intersegmental markings; BL 2.1-2.5 mm. On Orobanchaceae (Mimulus, Orthocarpus, Pedicularis) in western North America. Alate viviparae and males are produced in late August, suggesting host alternation, but attempted transfers to Ribes were unsuccessful (Gillette & Palmer 1934). Heie (1979) suggested that it might be a mid- to late summer form of  N. cynosbati. 2n=10.

Nasonovia (Ranakimia) altaensis Stenseth    Apterae are pale green; BL 2.4-2.9 mm. Alatae have dusky to dark dorsal abdominal markings. On Thalictrum simplex ssp. boreale (= rariflorum) in northern Norway and northern Finland. Monoecious holocyclic, with oviparae and alate males in July-August (Heikinheimo 1991).

Nasonovia (Kakimia) aquilegiae (Essig)    Apterae are pale or with a dark dorsal patch; BL 1.6-3.0 mm. On Aquilegia spp., widely distributed in North America. Monoecious holocyclic with oviparae and alate males in October-November. Heie (1979) discussed the variable pigmentation, and Heie (1987) described geographical variation. 2n=10.

Nasonovia (Kakimia) arizonensis Heie   Colour of apterae when alive is unknown, immatures are dirty greenish; BL 1.9-2.4 mm. On an uncultivated Ribes sp. in Arizona, USA, curling and spotting leaves. Smaller specimens (BL 1.3-1.8 mm) from R. cereum in Utah (vii.1965, BMNH collection leg. G.F. Knowlton) may also be this species, and it is also recorded from that host in Idaho and Washington (

Nasonovia (Kakimia) borealis Heie    Colour of apterae in life is unknown, probably pale yellowish with antennae and tips of leg segments and siphunculi dark; BL 1.9-2.9 mm. On Heuchera richardsonii in Canada (Manitoba, Ontario).

Nasonovia (Kakimia) brachycyclica Holman    Apterae are bright green or pinkish, with shining brown-black dorsal sclerotisation; BL 2.2-3.0 mm. Alatae are mainly dark brown to black. On Ribes uva-crispa in Europe (Czech Republic, France, Germany). Monoecious holocyclic, with an abbreviated life cycle, oviparae and alate males being produced in early June (original description).

Nasonovia (Kakimia) brevipes (Börner)    Apterae are probably pale brown; BL c.1.8 mm. Börner (original description) believed that the host was Hieracium, which would be unusual for a Kakimia, and requires confirmation. We  have included it in the key to apterae on Saxifraga, as this is a more likely host in the type locality (Heie 1979). Described from Austrian Alps, and an alata trapped in the Netherlands is possibly this species (Heie 1994).

Nasonovia (Kakimia) carolinensis Heie    Apterae are greenish, dirty yellow, or slightly brownish; BL 1.8-2.5 mm. On Saxifraga michauxii (now Spatularia michauxii) in eastern USA (Appalachians, North Carolina). Probably monoecious holocyclic; oviparae in late September (original description).

Nasonovia (Kakimia) castelleiae (Sampson)    Colour of apterae is unrecorded; BL 1.9-2.5 mm. On Castilleia spp. in western USA (California, Oregon) and Canada (Alberta, British Columbia, Manitoba), and now also recorded from C. pavlovii in north-east Siberia (Chukotka; Stekolshchikov & Khruleva 2015). Oviparae and alate males occur in August-September (Heie 1979) in North American populations, but in July in Siberia.

Nasonovia (Kakimia) collomiae Palmer    Apterae are pale yellowish; BL 1.2-2.2 mm. On leaves of  Collomia linearis in western USA (Colorado, Utah, Washington).  Oviparae and alate males occur in October-November (Palmer 1952).

Nasonovia compositellae (Theobald)    Apterae are shining black dorsally (see influential points/Gallery); BL 1.6-2.5 (ssp. iberica 2.3-3.1) mm. On Hieracium spp. in Europe, feeding (ssp. nigra) in spring on the upper sides of the leaves which fold upwards to enclose the colonies, later feeding on stems and inflorescences. Currently N. compositellae is regarded as including three subspecies. (1) N. compositellae ssp. compositellae occurs in north-west Europe and Iceland and is monoecious holocyclic with oviparae and apterous males in August-September (Prior & Stroyan 1960). (2) N. compositellae ssp. nigra (Hille Ris Lambers) is widely distributed in Europe, and is also in Turkey, Iran and Kazakhstan (but records from North America are unsubstantiated). It reproduces parthenogenetically throughout the year in UK, overwintering on the basal rosettes of Hieracium, but sexual morphs have been reared from populations collected in the Alps (Prior & Stroyan 1960). (3) N. compositella ssp. iberica Heie occurs in Spain, the host plant again being probably a Hieracium sp.  2n=11 (heterozygous) for ssp. nigra (one UK sample)

Nasonovia (Capitosiphon) crenicorna Smith and Knowlton  (Fig.30b)  Apterae are light green with a faint dark spinal stripe; BL 3.5-5.1 mm. On Geranium richardsonii and G. viscosissimum in western USA (Palmer 1952, as Macrosiphum crenicornum, and Monoecious holocyclic, with oviparae and alate males in July-August.

Nasonovia (Kakimia) cynosbati (Oestlund)  Plate 21i   Apterae are pale green, usually with dark tips to antennal segments and siphunculi; BL 1.8-3.0 mm. Alatae have very variable dorsal abdominal pigmentation; pale green or (esp. in more northerly populations) with variably developed cross-bands or a Myzus-like patch. On wild and cultivated Ribes spp., and also recorded from Heuchera (Heie 1979, 1980a). Widely distributed in North America. Monoecious holocyclic, with oviparae and alate males in September-October. N. (K.) suguri Shinji is possibly this species. 2n=10.

Nasonovia (Kakimia) dasyphylli Stroyan    Apterae are pale green to yellowish green, sometimes tinged with red, and with a variably developed black dorsal abdominal patch; BL 1.3-3.0 mm. On Crassulaceae (Sedum, Aichryson, Sempervivum) and Saxifragaceae (Saxifraga).  In Europe, south to Spain, Italy (Barbagallo & Patti 1998) and Canary Isles (BMNH collection, on Aichryson dichotonum, leg. K.W.R. Zwart), and eastward to Caucasus. Monoecious holocyclic in eastern Europe with apterous males, but apparently anholocyclic in UK. Heie (1979) compared populations from different parts of Europe. 2n=12 (heterozygous; one sample from UK, collected by J.H. Martin).

Nasonovia (Kakimia) davidsoni Heie    Apterae are dark reddish brown with extensive dark dorsal sclerotisation and dark appendages; BL c.2.0-2.1 mm. On flowerstalks of Heuchera spp. in western California. Monoecious holocyclic, with an abbreviated life cycle, producing oviparae and alate males as early as May (Davidson 1915, as Macrosiphum heucherae).

Nasonovia (Kakimia) dzhetisuensis Kadyrbekov    Apterae are green; BL c. 2.8-2.9 mm. On Polemonium caucasicum and P. coeruleum, feeding singly on stems, in Kazakhstan. Monoecious holocyclic, with oviparae and alate males in early August (Kadyrbekov 1995b). This species has subsequently been recorded from north-west Russia (Nenets region) on P. boreale (Stekolshchikov 2017), feeding on fruits and undersides of transversely twisted leaves, with sexuales present in July.

Nasonovia (Kakimia) grossa Heie    Apterae have an extensive dark dorsal abdominal patch and black siphunculi; BL 2.9-3.2 mm. On Ribes spp. in the Rocky Mountain region, USA.

Nasonovia (Ranakimia) heiei Kadyrbekov    Body colour of apterae is unrecorded but probably dark, shiny, with red eyes, and appendages pale green darker at apices; BL 1.8-2.9 mm. On Aquilegia spp. in Kazakhstan, feeding on stems below flower heads, and also once collected from Delphinium elatum (Kadyrbekov 1995b).

Nasonovia (Kakimia) heucherae (Thomas)    Apterae are tan to light brown with variable dorsal sclerotisation, and appendages mainly dusky to dark brown; BL 1.8-2.4 mm. On Heuchera spp. in USA east of Rocky Mountains. Monoecious holocyclic, with an abbreviated life cycle, producing oviparae and alate males in May-June (Heie 1979). N. davidsoni in California is closely related.

Nasonovia (Kakimia) hottesi Heie    Apterae are unknown; BL of alatae 2.5-3.2 mm. Only known from alatae collected from Polemonium sp. (probably the true host; Heie 1979) and Antirrhinum sp. In Illinois, USA.

Nasonovia (Kakimia) houghtonensis (Troop)    Apterae are pale yellow-green to straw-yellow; BL 1.5-2.9 mm. Alatae have a pale abdomen with no dorsal markings. The species seems to be a complex of races or subspecies monoecious holocyclic on Ribes spp. in North America, partly separated by geographical, life cycle and/or host plant differences, with intermediate forms sometimes occurring (Heie 1979). N. houghtonensis ssp. cerei (Gillette & Palmer) is the most hairy and has the longest siphunculi, and may be the ancestral form for the group; it occurs in the Rocky Mountain region and produces alate males and oviparae in July-September. N. houghtonensis ssp. russellae Heie is also in the Rocky Mountain region, preferring different Ribes spp., with oviparae and alate males in July-August. N. houghtonensis s. str. attacks the Houghton variety of R. uva-crispa, is widely distributed in North America and has apterous males, producing its sexual morphs in September-October. N. houghtonensis ssp. occidentalis Heie occurs in north-western USA (life cycle is unknown), and N. houghtonensis ssp. similis Heie occurs widely in Canada, producing oviparae and apterous males in August-October.

Nasonovia jammuensis Verma    Apterae are pale brown with darker brown head and thorax; BL 2.1-3.2 mm. On undersides of leaves of Delphinium spp. in north-west India (Heie 1979), and in Pakistan it has been collected on Delphinium in summer and on Consolida in winter (Naumann-Etienne & Remaudière 1995). 2n=12 (Dutta & Gautam 1993).

Nasonovia (Kakimia) muesbecki Knowlton & Allen    Colour of apterae in life is unknown, probably pale with dusky markings; BL 2.4-2.8 mm. On Ribes sanguineum in western USA (California, Oregon, Washington). Apparently heteroecious holocyclic, migrating in May-June to an unknown secondary host, probably a species of Polemoniaceae, as it is closely related to N. takala (Heie 1979). Very similar aphids on Polemonium foliosissimum in Utah  have been described as a subspecies, N. (K.) muesbecki ssp. montana (Heie 1987)and aphids collected on Polemonium sp. in Oregon have also been identified as this subspecies (

Nasonovia nivalis (Börner)    Apterae are shining yellow-brown with brown segmental markings or cross-bars; BL 1.7-2.3 mm. On Hieracium spp. in Austria. Monoecious holocyclic with oviparae in autumn, males are unknown (Heie 1979).

Nasonovia pilosellae (Börner)    Apterae are shining green with a series of paired dark dorsal markings to almost black (see; BL 1.2-2.5 mm. On Hieracium spp. living in spring inside upwardly rolled leaves, later moving onto stems and flowers (Hille Ris Lambers 1949). Throughout most of Europe, and in Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic, with oviparae and apterous males in October.

Nasonovia  (Kakimia) polemonii (Gillette and Palmer)    Apterae are pale to mid green, with distal parts of antennae and legs dark; BL 2.4-c.3.2 mm. On leaves and stems of Polemonium spp. in western USA (Colorado, Idaho, Utah). Apparently heteroecious, as Jensen has collected this species from Ribes sp(p). in Washington, Oregon and Arizona (

Nasonovia (Ranakimia) purpurascens (Oestlund)    Apterae are green, yellowish green or pale olive-green; BL 2.2-3.3 mm. Alatae have the dorsal abdomen pale or with indistinct markings. On Thalictrum spp. in North America and possibly also on Aquilegia sp (Leonard 1970). Monoecious holocyclic with an abbreviated life cycle, oviparae and alate males occurring in May-June.

Nasonovia (Ranakimia) ranunculi Heie    Apterae are rather pale, colour unknown; BL c. 2.8-2.9 mm. On Ranunculus sp. in Colorado, USA.

Nasonovia (Kakimia) ribifolii (Davidson)    Apterae are yellowish green with variable dark markings; BL c. 2.0 mm. On Ribes spp. in California, and also in New Mexico (Miller et al. 2016), curling and blistering leaves (the host plant in the original description was given as R. glutinosum, but corrected to R. malvaceum on the slide label). Monoecious holocyclic, with a very abbreviated life cycle; fundatrices are adult in March, and oviparae and alate males are produced in May (Heie 1979).

Nasonovia ribisnigri (Mosley)  Plate 21h   Apterae are shiny pale green to apple-green, sometimes reddish, with a dorsal pattern of dark brown markings when on secondary hosts (see; BL 1.3-2.7 mm. Alatae have a conspicuous pattern of black abdominal markings. In small colonies at shoot tips and in curled young leaves of Ribes spp. in spring, migrating to various plants in Compositae (Cichorium, Crepis, Hieracium, Lactuca, Lampsana), Brassicaceae (Sinapis, Sisymbrium), Orobanchaceae (Euphrasia), Plantaginaceae (Veronica) and Solanaceae (Nicotiana, Petunia). It seems more able to colonise plants with sticky hairs than most other aphids. In Europe, Middle East, Central Asia, Africa (Algeria, Burundi, Rwanda), North and South America, and more recently introduced to New Zealand (Stufkens & Teulon 2002) and Tasmania, from where it has now spread to mainland Australia (Dominiak et al. 2009). Differences between European populations of N. ribisnigri in ability to feed on resistant lettuce varieties were found by ten Broeke et al. (2013). Smith & Chaney (2007) studied natural control by syrphid predators in California. See also Heie (1979, 1994) and Blackman and Eastop (2000). 2n=12.

Nasonovia rostrata David & Hameed    Colour of apterae in life is unrecorded, probably rather pale; BL 1.6-2.1 mm. Originally described (both apterae and alatae) from Clerodendron infortunatum, and there have been subsequent collections from Compositae/Asteraceae (Adenocaulon, Conyza) and Acanthaceae (Strobilanthes). Heie (1979) redescribed the aptera.  On Strobilanthes spp. the aphids infest both undersides of leaves and inflorescences, and produce oviparae and alate males in October (Saha & Chakrabarti 1988a), so these are probably the true hosts. In northern India (Uttar Pradesh, Himachal Pradesh). 2n=12.

Nasonovia (Aconitaphis) salebrosa (Ivanoskaya)    Apterae are shining pale green with dark brown head, antennae, legs, cauda and dorsal markings, red eyes and pale siphunculi; BL c. 3.7 mm. On Aconitum and Delphinium spp. in Siberia (Ivanoskaya & Ostanin 1971) and Kazakhstan (Kadyrbekov 1995b).

Nasonovia (Kakimia) sampsoni Heie    Apterae are pale green, BL 1.8-2.5 mm. Living between the flowers of Castilleia spp. in western North America (California, Alberta).

Nasonovia (Ranakimia) saurotarbagataica Kadyrbekov    Apterae are bright yellow; BL 2.0-2.5 mm. Alatae are unknown. On flowers and flower stalks of Thalictrum spp. (collinum, minor, simplex) in east Kazakhstan (Kadyrbekov 2005a).

Nasonovia (Kakimia) saxifragae (Doncaster & Stroyan)    Colour of apterae in life is unknown, presumably often very dark dorsally, and with dark appendages; BL c. 2.0-2.1. On Saxifraga spp. in Greenland, Iceland, Jan Mayen, Norway (Heikinheimo 1990) and northern Britain (where a paler form occurs, see Heie 1987). Monoecious holocyclic, with oviparae and apterous males in August-September (Heie 1979).

Nasonovia (Kakimia) smithi Heie    Apterae are yellow tan, usually brownish or greenish, with extensive dark dorsal sclerotisation and mainly dark appendages; BL 1.3-2.5 mm. On Heuchera villosa in western mountains of North Carolina. Monoecious holocyclic, with oviparae and alate males in September-October (original description).

Nasonovia (Kakimia) stroyani Heie    Colour of apterae in life is unknown, probably pale yellow or pale green with antennae dark distally, siphunculi dusky-dark distally and cauda dusky-dark; BL 2.3-2.8 mm. Alatae have dark cross-bands on the dorsal abdomen, fused into a patch on tergites 3-5. On leaves and upper parts of stems of  Tellima grandiflora and Boykinia occidentalis (= elata) in western USA and Canada. The Nasonovia sp. near cynosbati observed by Jensen (2000: 451) on Tolmiea menziesii and two other species of Saxifragaceae is also presumably this aphid. The life cycle is unclear; on Tolmiea it may overwinter parthenogenetically (Jensen 2000), but immature apterous males have been collected in March (original description), and there are five males in the BMNH collected in early June in British Columbia (Victoria; leg. E.O. Essig). Previously this species has been confused with N. cynosbati; Heie (original description) compared these two closely-related species.

Nasonovia (Kakimia) takala (Hottes)    Apterae are yellowish green; BL 2.3-2.9 mm. On leaves of  Ipomopsis aggregata in western USA (California, Colorado, Idaho). Monoecious holocyclic, with oviparae and alate males in September.

Nasonovia (Kakimia) tiarellae Heie    Apterae are unknown. BL of alatae is 1.7-2.4 mm. On Heuchera americana and Tiarella cordifolia in North Carolina, USA. Monoecious holocyclic, with an abbreviated life cycle, oviparae and alate males appearing in April-June.

Nasonovia (Kakimia) vannesii Stenseth    Apterae are dark green; BL c. 3.0-3.1 mm. Alatae have a black dorsal abdominal patch and cross-bands. On Ribes spp. in Norway, Finland and north-west Russia, but suspected to be of North American origin (Heie 1979), and there are similar specimens from Ribes divaricatum in California in the BMNH collection (leg. D. Hille Ris Lambers). Probably heteroecious holocyclic; immatures and an alate male provisionally assigned to this species were found by Heikinheimo (1990) on Saxifraga groenlandica (= S. cespitosa) in Finland in late August.

Nasonovia (Kakimia) vockerothi (Richards)    Apterae are pale, colour not recorded; BL 2.1-2.7 mm. Alatae have dark dorsal cross-bands, partially fused. On Saxifraga tricuspidata in north-eastern Canada.

Nasonovia (Aconitaphis) wahinkae (Hottes)  (Fig.6d,e,f)  Apterae are shining green-bronze to dark green or orange-yellow (see; BL 2.3-3.6 mm. On Aconitum columbianum and Delphinium occidentale in western USA (Rocky Mountain region). Monoecious holocyclic; oviparae and alate males were collected on D. occidentale in August-September in Utah (Heie 1979). Populations on cultivated Delphinium in Canada (Manitoba) and Alaska differ in chaetotaxy and rhinarial number, and are regarded by Heie (1979) as a subspecies, N. (A.) wahinkae ssp. robinsoni (Richards). A population in north-east Siberia (Chukotka), producing sexual morphs in July on pedicles, flowers and undersides of slightly swollen leaves of D. brachycentrum ssp. maydellianum, has been identified as this subspecies (Stekolshchikov & Khruleva 2015).

Nasonovia werderi Lampel    Colour of apterae in life is unknown, BL 2.3-2.4 mm. On Aquilegia alpina in Switzerland, feeding on flower stems and inflorescences in July.

Nasonovia williamsi (Smith and Parron)    Apterae are shining yellowish green to amber-yellow (for photo see; BL 1.5-2.3 mm. On undersides of leaves of Potentilla spp., causing tight leaf-curl. In western USA (California, Idaho, Utah, Oregon, Nebraska) and Canada (Nebraska, Manitoba).

Nearctaphis Shaposhnikov

Aphidinae: Macrosiphini

About 12 North American species having short siphunculi ornamented with closely-spaced rows of small spinules or nodules and a short, usually triangular, cauda. Probably most of them alternate between Pomoideae as primary hosts and Leguminosae or Orobanchaceae as secondary hosts, but host alternation has only been confirmed by experimental transfers for two species. Most species are poorly known; it is quite likely that those Nearctaphis species recorded from one species of primary host may also utilise other species or genera of Pomoideae. Hille Ris Lambers (1970a) revised the genus, and Robinson (1984) briefly reviewed the genus and provided a key to species.

Nearctaphis argentinaeradicis Gillette & Palmer    Apterae are pale; BL 1.4-1.7 mm. On roots of Potentilla anserina in Colorado, USA. Monoecious holocyclic, with oviparae in late August (Palmer 1952). Hille Ris Lambers (1974) redescribed this species.

Nearctaphis bakeri (Cowen)  Plate 9e (Fig.40i)   Spring colonies infest tips of twigs, young leaves and blossom buds of woody Pomoideae in spring in North America (Crataegus, Cydonia, Malus, Pyrus).  Apterae in spring populations are pale green to yellow green, sometimes pinkish anteriorly; BL 1.4-2.3 mm.  Alatae have a black dorsal abdominal patch.  Heteroecious holocyclic, producing abundant alatae in the third and subsequent generations which migrate to Leguminosae (e.g. Medicago, Melilotus, Trifolium, Trigonella), and sometimes on plants in other families (Capsella, Castilleja, Valeriana, Veronica). It is an important pest on Trifolium pratense (Blackman & Eastop 2000, p.311).  Apterae on secondary hosts are dark green to salmon pink with variably developed dorsal dark spots or patches (see Smith (1923) gave a detailed account of this species.  Native to and widely distributed in North America, and introduced into Europe, the Middle East, Central Asia, India and Japan. Introduced populations all seem to be anholocyclic on secondary hosts.  2n=12.

Nearctaphis californica Hille Ris Lambers   Colour of apterae in life is unknown, probably dark; BL 2.2-2.9 mm. On Melilotus albus and “either Trifolium or Medicago”, in California, USA, and probably migrating to Sorbus as primary host, as apterae collected in British Columbia, Canada on Sorbus aucuparia appear to be this species (Robinson 1984).  Appearance on Sorbus in life is also unknown; adult apterae are probably rather dark with broad dark dorsal abdominal cross-bands, BL 1.5-2.0 mm (specimens from Sorbus, BMNH collection, leg. Cho-kai Chan).  2n=12. 

Nearctaphis clydesmithi Hille Ris Lambers   Apterae and alatae on Crataegus in spring and early summer are dark brown to black, somewhat shiny; BL 1.6-2.1 mm.  Apparently heteroecious, but secondary host generations are unknown.  Oviparae and alate males were collected on C. crus-galli in October (Hille Ris Lambers 1970a).  In eastern USA (North Carolina, Maryland, Virginia, Wisconsin).

Nearctaphis crataegifoliae (Fitch)  Apterae in spring colonies on Crataegus are greenish or reddish, BL 1.6-2.5 mm, twisting and curling the leaves and turning them dark purple.  Heteroecious holocyclic, migrating in May-June to stem bases, runners and roots of Leguminosae (Trifolium, Melilotus, Lathyrus).  Apterae on the secondary hosts are olive-yellow or pink to brilliant red; BL 1.7-2.3 mm. Patch (1915b) described the life cycle (as Aphis brevis).  In central and eastern North America.  Aphids with a similar biology in western USA (California, Colorado, Utah) have much darker pigmentation of head, posterior abdomen and siphunculi, and were described (Hille Ris Lambers 1970a) as a subspecies, N. crataegifoliae occidentalis.

Nearctaphis hottesi Hille Ris Lambers    Apterae are greyish green with a very irregular dorsal abdominal pattern of  “twisted lines in olive to reddish brown”; BL c.1.4-1.8 mm.  In flower heads of  Castilleja linariaefoliae in Colorado, USA. Probably migrating to an unknown primary host; alate males were collected on Castilleja in September.

Nearctaphis kachena (Hottes)    Apterae are dark green, mottled with dark brownish black,with yellowish green siphunculi; BL 1.5-2.0 mm. In flower heads of Castilleia spp. in western USA.

Nearctaphis nigrescens Hille Ris Lambers    Colour of apterae in life is unknown, probably dark green to black; BL c. 2.7-2.8 mm. On Melilotus albus in Utah, USA. The primary host is probably Crataegus sp., as two alate gynoparae (BL 2.2-2.6 mm) collected in September on Crataegus sp. in Utah were assigned to this taxon by Hille Ris Lambers (1970c), and it has also been recorded from Crataegus in spring in Washington (A. Jensen, N. nigrescens was described as a subspecies of  N. californica, but the differences appear to warrant full species status.

Nearctaphis sclerosa (Richards)  Apterae in spring colonies are dark brown; BL about 1.9 mm.  It is apparently heteroecious holocyclic between Crataegus douglasii and Lathyrus nutalli in British Columbia, but with an early return to the primary host; gynoparae and oviparae were collected on C. douglasii in early August (Richards 1968c, as Roepkea sclerosa).  It is also recorded from Colorado (Smith & Parron 1978).

Nearctaphis (Amelanchieria) sensoriata (Gillette & Bragg)  Apterae in life are brown to blackish; BL 2.0-2.2 mm.  On leaves and twigs of Amelanchier sp(p). in western North America.  Alatae black, with a large solid dorsal abdominal sclerite.  Probably heteroecious holocyclic with unknown native Leguminosae as secondary hosts; an aptera was collected on Medicago hispida, a non-indigenous leguminous plant, in California (Hille Ris Lambers, 1970a). Autumn populations on Amelanchier comprise alate gynoparae, oviparae and alate males (Hille Ris Lambers 1970a).  However, populations persist on Amelanchier into summer, and host alternation needs to be confirmed.

Nearctaphis vera (Shaposhnikov)    Apterae are undescribed; alatae are dirty green, BL c.2.7 mm. Alatae have secondary rhinaria distributed III 42-44, IV 25-27, V 10-13. On Oxytropis sordida in northern Russia (Khibiny Mountains, Kola Peninsula; Heie 1992, as Mamontova vera) and also recorded from Finland (Albrecht 2010).

Nearctaphis yohoensis Bradley   Apterae in life are dark brown, BL 1.7-2.2 mm, in dense ant-attended colonies on new growth and flowers of Sorbus scopulina in British Columbia (Bradley 1965b).  Also recorded from Montana, Manitoba, Idaho and Alaska. The Alaska collection was from Cotoneaster acutifolia in early July (Robinson 1984), and apterae of what appears to be the species were collected at the same time from Pedicularis (Orobanchaceae). It is unclear whether host alternation was occurring between Cotoneaster and Pedicularis, or whether both were acting as secondary (or accidental) hosts.

Nearctaphis zabapsis (Richards)    Colour of apterae in life has not been observed, probably dark; BL c.1.5 mm. On roots of Trifolium sp. in Colorado, USA.

Neoamphorophora  Mason

Aphidinae: Macrosiphini

Two species with clavate siphunculi related to Wahlgreniella but with shorter antennae with a shorter terminal process, and apterae have no rhinaria on ANT III.

Neoamphorophora kalmiae Mason    Apterae are pale green with pale appendages and variably developed, ill-defined, dark brown to black dorsal pigmentation; BL 1.6-2.1 mm. Immatures are pale and slightly waxy. Alatae have paired dark sclerites on abdominal tergites 3 and 4, and sometimes on other tergites. On Kalmia angustifolia in north-eastern USA and eastern Canada. Monoecious holocyclic, with oviparae and alate males in September (MacGillivray 1967).

Neoamphorophora ledi (Wahlgren)    Apterae in life are yellowish green to dark green or brownish pink, with darker dorsal abdominal markings, their siphunculi dark brown with paler bases; BL 1.8-2.9 mm. Alatae have a variably developed dark dorsal abdominal patch and 35-40 secondary rhinaria on ANT III only.  On Rhododendron (=Ledum) palustre in northern Europe, and now reported from Rhododendron spp. in Japan by Barjadze & Özdemir (2017) who provided a full redescription of apterae and alatae. Sasaki et al. (2021) reported several more localities in Japan. Monoecious holocyclic with alate males (Heie 1995).

Neoantalus  Remaudière


Two species differing from Pterasthenia in the absence of wax glands, and alatae having secondary rhinaria on ANT III-V (Remaudière & Autrique 1985, Quednau 2010).

Neoantalus aeschynomenidis (van Harten & Ilharco)    Apterae are green, BL 1.25-1.7 mm. In small numbers on growing tips of Aeschynomene dimidiata in Angola. Very close to N. humulariae and it could be the early seasonal form of N. humulariae (Remaudière & Autrique 1985, Quednau 2010).

Neoantalus humulariae (van Harten & Ilharco)    Apterae are bright yellow to dark green; BL 1.5-1.9 mm. On undersides of leaves and sometimes on flowers of Humularia welwitschii in Angola. Alatae have also been trapped  in Burundi (Remaudière & Autrique 1985). Partially monoecious holocyclic, with oviparae and alate males collected along with apterae and alate viviparae in early July (original description).

Neobetulaphis Basu

Calaphidinae: Calaphidini

An oriental genus on Betulaceae, nominally with five species, but the differences between Neobetulaphis alba in Japan and N. pusilla in India are those to be expected between small and large specimens respectively of the same species.  A.K. Ghosh & Quednau (1990) reviewed and keyed the three Indian species.

Neobetulaphis alba Higuchi   Apterae are pale yellow in life, BL c. 1.5 mm.  Alatae are undescribed.  On Betula platyphylla var. japonica in Japan (Higuchi 1972), and also recorded from China (Qiao et al. 2005b).  The life cycle is unknown.  Perhaps a synonym of N. pusilla.

Neobetulaphis chaetosiphon Quednau & Chakrabarti   Colour in life is not recorded, apterae are presumably rather dark, brownish; BL 2.1-2.3 mm.  Alatae have no dorsal black abdominal patch.  On Alnus nepalensis and Betula spp. (alnoides, utilis) in India (Meghalaya, Sikkim, West Bengal).  Apterae have also been recorded from Quercus sp. (A.K. Ghosh & Quednau 1990). The life cycle is unstudied, but viviparae have been found through the winter.  As noted by Chakrabarti (1988), the distinction between this species and N. immaculata when size and seasonal variation are taken into account needs to be verified.

Neobetulaphis immaculata Ghosh   Apterae are pale yellow; BL 1.2-1.5 mm.  Alatae are undescribed.  Originally described from Rosaceae in Meghalaya, India (A.K. Ghosh 1976), but Chakrabarti (1988) recorded it from a more likely host, Betula alnoides, in Uttar Pradesh. The life cycle is unknown.

Neobetulaphis pusilla Basu   Apterae are yellowish green with dark green cross-bands on thorax and abdomen; BL 1.7-2.3 mm.  Alatae have a black patch on abdominal tergites 4-6 and dark siphunculi.  On undersides of leaves of Betula spp. (alnoides, utilis), and possibly Alnus sp., in India (Himachal Pradesh, Sikkim, Uttar Pradesh, West Bengal), and also recorded from China (Qiao et al. 2005b). A record from Turkey (Görür et al. 2011a) requires additional confirmation. Chakrabarti (1988) described the apterous male and ovipara from Uttar Pradesh in October, but viviparous morphs occur through the winter months in Sikkim and West Bengal.  The east Asian N. alba may be a synonym, and N. hebeiensis (Zhang, Zhang & Zhong), described from oviparae collected on Betula platyphylla in China (as a subspecies of Callipterinella callipterus; W. Zhang et al. 1995b) is not clearly distinct.

Neochromaphis Takahashi

Calaphidinae: Pterocallidini

Two east Asian species feeding on twigs and branches of Betulaceae, and visited by ants.  They are stout-bodied aphids with pigmented wings, long dorsal abdominal hairs and a very short antennal terminal process.  Quednau (2003) provided a key and illustrations.  (A third western Palaearctic species on Ostrya, ostryae Börner, was placed in this genus by Eastop & Hille Ris Lambers (1976), but is here regarded as a Pterocallis).

Neochromaphis carpinicola (Takahashi)  All viviparae are alate, stout-bodied, dark brown with forewings almost wholly pigmented (see Moritsu 1983); BL 1.7-2.0 mm.  On undersides of twigs and branches of Carpinus spp. in Japan.  Alate males and apterous oviparae occur in October-November (Takahashi 1923, Quednau 2003).

Neochromaphis coryli Takahashi   All viviparae are alate, stout-bodied, brownish, with maculate forewings; BL 1.6-2.2 mm.  On twigs and branches of Corylus spp. in China, Japan and Korea (the latter as carpinicola; Paik 1965).  Quednau (2003) illustrated an ovipara collected on Corylus avellana in mid-September.  2n=18 (Chen & Zhang 1985b).

Neocranaphis Ghosh & Quednau

Calaphidinae: Panaphidini

Two Asian bamboo-feeding species originally placed in Shivaphis, but distinguished by the setiform empodial hairs and other characters (A.K. Ghosh & Quednau 1990). Phyllaphoides is also closely related. Quednau (2003) provided a key and illustrations of both species.

Neocranaphis arundinariae (Takahashi)   All viviparae are alate, elongate-bodied, white, dusky on thorax, without dorsal abdominal markings, secreting cottony wax on anterior part of body, hind legs and (slightly) on antennae; BL c. 2.1 mm. On undersides of leaves of Arundinaria (Pleioblastus) sp. in Taiwan (Liao, 1976, as Shivaphis arundinariae), and also recorded from mainland China (Qiao et al. 2005b). Sexual morphs and life cycle are unknown.

Neocranaphis bambusicola (David, Rajasingh & Narayanan)   All viviparae are alate, elongate-bodied, white, with powdery wax covering body, legs and antennae; BL 2.7-3.0 mm. On undersides of leaves of Bambusa sp. in West Bengal, India. An ovipara and an apterous oviparoid (intermediate) female were collected in November (David et al. 1970, as Cranaphis).

Neodermaphis Sorin

Hormaphidinae: Nipponaphidini

Two species in Japan closely related to Dermaphis but the apterae are smaller and flatter, with differences in the cuticular sculpturing and the ommatidia, and greatly reduced antennae (Sorin 2006).  The apterous viviparae are known only from single collections and the alatae are undescribed.

Neodermaphis lineata Sorin   Apterae are elongate oval, black; BL c. 0.7 mm. Collected from twigs of Quercus glauca in Japan (Honshu) in January (Sorin 2006).

Neodermaphis unebiensis Sorin   Apterae are black with white wax powder on lateral perpendicular areas; BL c. 0.9 mm. Collected on branches of Quercus glauca in Japan (Honshu) in May (Sorin 2006).

Neohormaphis Noordam

Hormaphidinae: Nipponaphidini

Three species in east Asia with alatae having unusual wing venation, the media, Cu1a and Cu1b all arising from a common stem. A fourth species tentatively assigned to this genus has been collected on Quercus (= Cyclobalanopsis) championii in Hong Kong (BMNH collection, leg. J.H. Martin). Immature alatae of the Hong Kong aphid secrete curious glassy filaments resembling miniature fibre-optic bundles, from large circular dorsal wax glands (J.H. Martin, personal communication). The two named Chinese species were reviewed by Qiao et al. (2018).

Neohormaphis calva Noordam     Galls on upper sides of leaves of Distylium stellare in Java are pale greenish, semiglobose, downy, about 1.5 cm in diameter, usually arising singly from the mid-rib or a smaller vein.  Alatae (BL 1.4-1.8 mm, with black eyes and dark grey abdomen) leave the gall in September through a small hole on the underside of the leaf (Noordam 1991). Migration occurs to Quercus sp.; BL of apterae on Quercus is 0.8-1.0 mm, appearance in life unknown, but probably secreting some form of wax. (Immature alatae on Quercus may secrete wax filaments as they have similar wax glands to the undescribed species from Hong Kong mentioned above.)

Neohormaphis quercisucta Qiao, Guo & Zhang   Apterae are black, with short wax threads on marginal area and with 3-4 thick wax threads dorsally; BL 0.7-1.0 mm. Alatae and life cycle are unknown. On an unidentified Quercus sp. in China (Yunnan). The description of this species bears some resemblance to that of the secondary host form of Neohormaphis calva, although with clear differences including the reticulated tergum and shorter HT II (Qiao et al. 2005a, as Lithoaphis – but note that there must be errors in the measurements given in their table). Redescriptions of immatures and adult apterae were provided by Qiao et al. (2018).[The transfer from Lithoaphis was made by Chen et al. (2014).]

Neohormaphis wuyiensis Qiao & Jiang    Apterae on undersides of leaves of unidentified species of Lithocarpus and Quercus in China are smoky blue, secreting thick radiating plates and bars of wax (Jiang et al. 2008). Wax glands of immature and adult apterae were illustrated by Chen & Qiao (2012c). Alatae are undescribed and the life cycle is unknown.

Neomariaella Swedo &Osiadacz

Aphidinae: Macrosiphini

One species possibly related to Brachycolus or Brachycorynella but with a short rounded cauda as well as very short conical siphunculi, and apterae having 5-segmented antennae.

Neomariaella lambersi (Szelegiewicz)     Apterae are pale green to yellow, dusted with grey wax powder; BL 1.1-1.8 mm. On young shoots of Myricaria spp., described from Poland (Szelegiewicz 1961b, as Mariaella lambersi), and since recorded from Slovakia, alpine France, Turkey, Iran, Kazakhstan and Mongolia (Szwedo & Osiadacz 2010). Monoecious holocyclic, with oviparae and apterous (alatiform) males in Poland in October (BMNH collection, leg. J. Nast). Remaudière et al. (2006, as Mariaella lambersi) provided a redescription, including the males and oviparae.

Neomyzus van der Goot

Aphidinae: Macrosiphini

About eight Asian species with distinctive black dorsal markings, previously placed as a subgenus of Aulacorthum but differing in several morphological features (shape of head, hairs on R IV+V, immatures with spinulose hind tibiae) that place them closer to the Myzus group of genera.

Neomyzus circumflexus (Buckton)  Mottled Arum Aphid   Plate 19c  Apterae are shining nearly white or pale yellow to bright green, with distinctive sclerotic dorsal markings,  consisting of transverse bands or paired patches on the thorax and a large roughly U-shaped, patch on the abdomen (see; BL 1.2‑2.6 mm. Extremely polyphagous, feeding on numerous species of both monocots and dicots, and even ferns and gymnosperms. In temperate climates N. circumflexus is found especially in glasshouses and on house plants (e.g. Cineraria,  Cyclamen, Fuschia, Zantedeschia). Piron (2017) recorded many new host records from the Netherlands (as Aulacorthum circumflexum). Virtually world‑wide, presumably due to transportation by man. Its origin is unknown, but east Asia seems most likely. Apparently it is entirely anholocyclic; no sexual morphs have been recorded. See also Blackman & Eastop 2000, p.244, as Aulacorthum (Neomyzus).  2n=8 ( except one record of 2n=10; Kar & Khuda-Bukhsh, 1986) .

Neomyzus codonopsis (Miyazaki)   Apterae are greenish white with distinctive irregular black markings across the thorax and abdomen (see Takahashi 1965a, as Aulacorthum taiwanum); BL 1.6-2.3 mm. Occurring singly on leaves of Codonopsis lanceolata in Japan. Alatae are undescribed. Formerly this species was placed as a subspecies of taiwanus.  2n=36.

Neomyzus dendrobii (A.N. Basu)  (Fig.25d)  Apterae are creamy yellow, ornamented with a pattern of brown to purplish brown segmental markings; BL 1.6-2.4 mm. On young stems of  Dendrobium sp. in West Bengal, India. Anholocyclic.

Neomyzus dicentrae (A.N. Basu)    Apterae are pale yellowish with brown dorsal markings; BL 1.6-2.5 mm. Alatae have a large dark dorsal abdominal central patch. Forming colonies on undersides of leaves of Dicentra thalictrifolia (= Dactylicapnos scandens) in north-east India (West Bengal).

Neomyzus parthenocissi (Takahashi)    Apterae are green with a brown-black dorsal abdominal patch; BL 1.0-1.3 mm. Described from Parthenocissus tricuspidata (Vitaceae), and also collected on Schizophragma hydrangeoides (Hydrangeaceae; BMNH collection, leg. R.L. Blackman). Only known from Japan. 2n=12.

Neomyzus primulus (A.K. Ghosh, Banerjee & Raychaudhuri)    Apterae are white with black dorsal markings; BL 1.6-1.8 mm. On Primula sp. in Sikkim, India.

Neomyzus strobilantheae (L.K. Ghosh)    Colour of apterae in life is unrecorded; BL 2.1-2.4 mm. On young shoots of Strobilanthes atropurpureus in Himachal Pradesh, India. The generic placement of this species is uncertain.

Neomyzus taiwanus (Takahashi)    Apterae are white with black dorsal markings; BL c. 2.3 mm. On Codonopsis javanica (ssp. japonica) in Taiwan (Tao 1963). Alatae are unknown.

Neonipponaphis Takahashi

Hormaphidinae: Nipponaphidini

A genus for two species with prosoma distinctly separate from abdominal tergites 2-7 and bearing numerous fine hairs.

Neonipponaphis pustulosus Chen & Qiao   Apterae are aleyrodiform, reddish brown or blackish brown in life, brown in mounted preparations; BL 1.4-1.6 mm. Living on twigs of Castanopsis eyrei, attended by ants (Chen & Qiao 2012b). In China (Fujian province). A redescription of the aptera was provided by Qiao et al. (2018). Alatae and life cycle are unknown.

Neonipponaphis shiiae Takahashi  (fig. 36B) Apterae are aleyrodiform, blackish brown to almost black with a reddish tinge in life, but pale in cleared preparations; BL 1.3-1.7 mm.  Throughout the year on branches and shoots of Castanopsis cuspidata and C. sieboldii in Japan (Takahashi 1962; Sugimoto 2019), often under ant shelters (Aoki et al. 2021). Alatae appear in May, and are black with dusky wings. The life cycle is unknown.

Neopemphigus Mamontova & Kolomoets

Eriosomatinae: Pemphigini

One palaearctic species feeding on poplar roots, closely related to Pachypappa and possibly belonging in that genus.

Neopemphigus turajevi Mamontova & Kolomoets   In dense wax-dusted colonies causing gall-like swellings on fine roots of Populus bolleana (= P. alba var. pyramidalis) in Ukraine (Mamontova & Kolomoets 1981).  Alate sexuparae appear in September.  Apparently it lives without host alternation on Populus, but the life cycle needs to be clarified.  Severe damage by this aphid has been reported (Kolomoets 1984).

Neophyllaphis Takahashi


About 18 species on Podocarpaceae and Araucariaceae in the southern hemisphere and mountains of the tropics, and extending northward into China and Japan.  Subgenus Chileaphis occurs only in South America. An unusual feature of Neophyllaphis is that several species have winged oviparae.  Where sexual morphs are known, they generally occur in mixed colonies with parthenogenetic morphs, often over several generations in spring, summer or autumn.  Hille Ris Lambers (1967b) and Russell (1982) provided reviews, and Quednau (2010) keyed the species and provided illustrations of all available morphs. The Chinese species were reviewed and keyed by Qiao et al. (2001a), and the Javanese species by Noordam (2004).  The structure and function of adhesive vesicles on the posterior abdominal sternites was studied by White & Carver (1971).

Neophyllaphis araucariae Takahashi   Apterae are broadly oval, bright yellow to yellowish orange; BL 1.1-1.5 mm.  On young growth of Araucaria spp. in Mauritius (original description), Java, New Guinea, Australia, Hawaii, Costa Rica, Bahamas, Mexico and USA (Florida, California).  Perhaps native to Norfolk Island or Australia (Carver 1959), or south-east Asia.  Probably mainly anholocyclic, but the alate ovipara is known from Hawaii (Carver 1971).  2n=18 (Hales & Lardner 1988).

Neophyllaphis brimblecombei Carver   Apterae are cream-coloured under a coating of greyish wax; BL 1.4-2.1 mm.  On Podocarpus elatus in Queensland, Australia, and on P. chinensis and P. macrophyllus in Hong Kong (Martin & Lau 2011).  Qiao et al. (2001a) reported this species from Eucalyptus robusta in China.  Hales (1971) gave an account of the biology (although her aphids on P. spinulosus were N. lanata).  Alate oviparae and males occur in spring and summer (Carver 1971); eggs and oviposition were described by Carver & Hales (1974).  2n=26 (Hales & Lardner 1988).

Neophyllaphis burostris Qiao & Zhang  Appearance in life is unrecorded: BL of aptera c. 1.6 mm. Alatae are undescribed. On Podocarpus macrophyllus in Guangxi, China (Qiao et al. 2001a).

Neophyllaphis (Chileaphis) cuschensis Nieto Nafría & Delfino   Apterae are brown with thorax and abdomen covered in ash-coloured wax powder; BL 1.5-2.3 mm. In compact groups on young branches of Podocarpus glomeratus in Peru (Mier Durante et al. 2008). Alatae have a series of unpaired dark spinal abdominal sclerites. Alate oviparae were noted, but are undescribed.

Neophyllaphis fransseni Hille Ris Lambers   Appearance in life is unrecorded; BL of aptera 1.0-1.6 mm.  On Podocarpus spp. in Java.  Alate oviparae are produced in January-March (Russell 1982; and BMNH collection, leg. D. Hille Ris Lambers).

Neophyllaphis gingerensis Carver   Apterae are purplish blue, with a coat of white wax; BL 1.4-1.8 mm.  On Podocarpus alpina, living singly or in small groups on leaves, in the mountains of Australian Capital Territory.  Apterous oviparae and alate males occur in January-February (Carver 1959).  This is the only species of the genus known to have apterous oviparae, perhaps as an adaptation to the mountain habitat (Eastop 1966).  Milner & Holdom (1986) recorded a fungal pathogen.  2n=14 (Hales & Lardner 1988).

Neophyllaphis grobleri Eastop   Apterae are orange-coloured under a thick coat of wax; BL 1.7-1.8 mm.  On Podocarpus spp. in Africa (Angola, Cameroon, Ethiopia, Kenya, South Africa, Tanzania), feeding on both young and full-grown trees.   Heavy attack can stunt growth of seedlings (Eastop 1955).  Alate oviparae (for description see Russell 1982) and males (Dürr 1971) occur in spring.  Ponsen (1990) studied the digestive system.  2n=18*.

Neophyllaphis (Chileaphis) iuiuyensis Mier Durante & Ortego   Apterae are pale green with longitudinal bands of varying intensity, pale yellowish legs and antennae, and thorax and abdomen very lightly covered in ash-coloured wax powder; BL 1.7-2.5 mm. In compact groups on young branches of Podocarpus parlatorei in Argentina (San Salvador de Jujuy). Alatae have a series of unpaired dark quadrate spinal abdominal sclerites. Alate oviparae and males were collected at the same time as viviparae in November (Mier Durante et al. 2008).

Neophyllaphis lanata Hales & Lardner   Apterae are deep reddish brown to reddish purple, covered in a woolly white wax exudate; BL c. 1.5 mm.  On stems and leaves of young shoots of Podocarpus spinulosus (a shrub) in Australia.  Hales & Lardner (1988) described enzyme, chromosome and other differences from N. brimblecombei, with which it was previously confused.  Alate oviparae and males start to occur in early spring (September).  2n=24 (Hales & Lardner 1988).

Neophyllaphis (Chileaphis) michelbacheri (Essig)  Apterae are broadly oval, whitish grey in life due to covering of wax powder; BL c. 1.8 mm.  Alatae have dark dorsal abdominal cross-bands.  In Chile.  Described from Pilgerodendron uviferum, but also recorded (Carrillo 1980) from Podocarpus nubigena, which is more likely to be the normal host. Alate oviparae and males occur in January (Russell 1982).  Oviparae are unusual in having numerous scent plaques on fore and middle tibiae as well as on the hind tibiae (Essig 1953b).

Neophyllaphis podocarpi Takahashi   Plate 8 c,d   Apterae are broadly oval, reddish purple thickly dusted with bluish white wax, often in transverse intersegmental bands; BL 1.5-2.0 mm.  On leaves, young twigs and fruit stems of Podocarpus spp. and Nageia nagi in China, Japan, Taiwan, Vietnam, Java, Malaya, and introduced to North America,  Hawaii, Italy (Limonta 1990), Spain (Pérez Hidalgo et al. 2015) and the Azores (Aguiar et al. 2013).  It often occurs in dense colonies, causing stunting and curling of new growth (Johnson & Lyon 1988, p.294).  Alate oviparae and males occur in Japan in both spring and autumn (Shibata 1955); elsewhere, sexual morphs appear rarely.  Russell (1982) summarises all available information.  2n=26 from Japan (Blackman 1986), or 2n=24 from China (Chen & Zhang 1985); perhaps indicative of more than one species under this name.

Neophyllaphis (Chileaphis) podocarpini Carrillo   Apterae are green and red in life; BL c. 2.2 mm.  On Podocarpus salignus in Chile; this species could not live on P. nubigena (Carrillo 1980).  Alate oviparae and males occur along with parthenogenetic morphs in December.

Neophyllaphis (Chileaphis) propinqua Quednau   Colour of aptera in life is unknown; BL 2.1-2.5 mm. On Podocarpus lambertii in Brazil (Quednau 2010). Alate oviparae and males were collected in October-December. 

Neophyllaphis (Chileaphis) pueblohondensis Quednau   Colour of aptera in life is unknown; BL 2.0-2.4 mm. On Podocarpus sp. in Venezuela (Quednau 2010). Other morphs are unknown.

Neophyllaphis rappardi Hille Ris Lambers   Apterae are dark brown-purple, covered with wax; BL c. 1.7 mm.  On Agathis labillardieri in Biak, Papua (Indonesia).  Found in small colonies on undersides of very young needles of older trees in deep forest shade (Hille Ris Lambers 1967b).  Sexual morphs are unknown.

Neophyllaphis totarae Cottier   Apterae are dark purplish brown, dusted with whitish wax powder; BL c. 2.8 mm.  On Podocarpus spp. in New Zealand, usually feeding singly on leaves or in small colonies on young twigs.  Alate oviparae and males occur in spring and summer (Cottier 1953, and BMNH collection). The native parasitoid reported by Valentine (1975) has now been described (Mackauer & Finlayson 2012). 2n=10*.

Neophyllaphis varicolor Miller & Halbert   Apterae are purple to dark wine-red, wax-dusted; BL 1.2-1.6 mm. Immatures may be yellow, orange, red or purple. Deescribed from Florida, USA, in colonies on new growth of Podocarpus spp. and Afrocarpus sp. (Miller & Halbert 2014); now also recorded from Costa Rica, and from a Podocarpus sp. in Vietnam, where it is presumably indigenous (Zúñiga-Centeno et al. 2019). Sexual morphs have not been recorded.

Neophyllaphis viridis Ilharco   Apterae are dark green, covered with thick wax powder; BL 1.5-1.8 mm.  On Podocarpus (= Afrocarpus) falcatus in South Africa, found in a mixed colony with N. grobleri (Ilharco 1973; Dürr & Martin 1976). The alate vivipara was described by Russell (1982).  Sexual morphs are unknown.

Neoprociphilus Patch

Eriosomatinae: |Pemphigini

One nearctic species with host alternation between Acer  and Smilax.

Neoprociphilus aceris (Monell)  (fig.18A)  Apterae in life are covered in white wax; BL 3.3-5.3 mm.  On undersides of twigs and branches of Acer saccharum in spring, sometimes extending onto petioles and leaves.  Throughout range of A. saccharum in eastern North America.  Heteroecious holocyclic; alatae produced in May-June migrate to found colonies on stems and leaves of Smilax spp. (Smith & Graham 1967).  Apterae on Smilax are pale brown to purplish, producing varying amounts of wax wool, which forms a tuft at end of abdomen; BL 3.7-4.5 mm. Sexuparae return to A. saccharum from late August to October, although overwintering can also occur as hibernating close-knit clusters of first instar nymphs in leaf litter, even in regions with severe winters outside the range of A. saccharum.  In spring these first instars may climb plants and be dispersed by wind, with wax filaments acting as sails (Lee & Robinson 1976).  2n=14.

Neopterocomma Hille Ris Lambers

Aphidinae: Macrosiphini

A little-known genus of three species, two from basal and underground parts of Salix in Europe, the third from Populus in China.  No alate morph is known, and in the one case where sexuales have been collected the males appear to be paedogenetic.

Neopterocomma asiphum Hille Ris Lambers   Apterae are reddish (Heinze 1961), probably wax-covered; BL 2.5-3.1 mm.  On lower part of trunk of various Salix spp., attended by ants.  Widely distributed in central and eastern Europe.  Other morphs and life cycle are unknown.

Neopterocomma populivorum Zhang   Appearance in life is not recorded; BL of aptera c. 2.8 mm.  On Populus sp. (feeding site not recorded) on Changbei Mountain, Jilin, China (Zhang & Zhong 1990).  Other morphs and life cycle are unknown.

Neopterocomma verhoeveni Hille Ris Lambers   (fig. 106B)  Apterae are covered with fine wax powder, dirty reddish or purplish grey; BL 2.0-2.6 mm.  In small colonies on underground parts of older branches of Salix repens, attended by ants.  Only known from the Netherlands.  Monoecious holocyclic; sexuales were found in October.  Males are yellowish white, dwarfish, and apparently only moult once to attain maturity (Hille Ris Lambers 1956a), although this is so unusual as to require further confirmation.

Neoreticulaphis Sorin

Hormaphidinae: Nipponaphidini

One species described from Quercus in Japan, related to Reticulaphis or Neothoracaphis but with circular pore-like prosomal ornamentation and hind legs concealed beneath body.

Neoreticulaphis nipponica Sorin   Aptera are blackish brown to black, almost circular with dorsum somewhat domed; BL c. 0.8 mm. On Quercus gilva in Japan, feeding on uppersides of leaves along mid-ribs. Apparently anholocyclic with only one generation per year (Sorin 1999b). Alatae are unknown.

Neorhopalomyzus Tao

Aphidinae: Macrosiphini

Two east Asian Myzus-like species with a sexual phase on Lonicera, migrating to unknown secondary hosts.

Neorhopalomyzus lonicericola (Takahashi)    Apterae (fundatrices) are dirty yellow, with black-tipped siphunculi; BL 2.7-3.5 mm. On Lonicera spp., feeding on undersides of leaves, which turn yellow and spotted with purple, but are not curled (Hori 1938b, as Amphorophora lonicericola). All progeny of fundatrices are alatae, with 8-14 secondary rhinaria on ANT III only, migrating to an unknown secondary host. Return migrant females (with secondary rhinaria distributed III 34-58, IV 19-34, V 0-12), oviparae and alate males (with secondary rhinaria distributed III 54-79, IV 23-35, V 1-7) are found on Lonicera in late September-October. In China, Japan, Korea (Lee et al., 2002c) and east Siberia.

Neorhopalomyzus lonicerisuctus Zhang, Zhang & Zhong    Apterae (fundatrices) are dark brown, dusted with white wax powder; BL c.2.6 mm. On Lonicera spp. in China.

Neosaltusaphis  Hille Ris Lambers


A single species related to Subsaltusaphis, but with the posterior corners of abdominal tergite 7 and the anterior corners of the pronotum developed into processi, and the dorsal hairs minute and spatulate. The genus was reviewed and illustrated by Quednau (2010).

Neosaltusaphis bodenheimeri Hille Ris Lambers (Fig.20a) Apterae are straw-coloured, with antennae black on distal halves; BL c. 2.3 mm. Alatae are unknown. Described from  Austria (Styria), and now also recorded from Kazakhstan (Kadyrbekov 2016d). The host is “presumably a Carex species” according to the original description, although swept from a wet meadow which included other Cyperaceae.

Neosappaphis Hille Ris Lambers

Aphidinae: Macrosiphini

Two species resembling Anuraphis but with smooth flangeless siphunculi (flangeless in the type species) and 4- or 5-segmented antennae.

Neosappaphis franzi Hille Ris Lambers    Appearance of apterae in life is unknown; BL 1.6-1.95 mm. Apterae have secondary rhinaria distributed III 2-3, IV 11-24 (also present in immatures), alatae are unknown. Sieved from a meadow in which Armeria (presumably maritima) was the dominant plant, on Islas Cies, north-west Spain. Alatae are unknown.

Neosappaphis paradoxa (Mamontova-Solukha)    Apterae are pale greenish; BL c.1.9 mm. On roots of Sisymbrium polymorphum in Ukraine; also in Kyrgyzstan, and in Kazakhstan, where it is also recorded from S. loeselii (Kadyrbekov 2017a).

Neosensoriaphis Quednau


A genus for one Nothofagus-feeder related to Neuquenaphis, but the aptera has finger-like processes only on abdominal tergites 7 and 8, and a strong pattern of polygonal reticulation of the dorsal cuticle (Quednau 1990, 2010).

Neosensoriaphis parva Quednau   Appearance in life is unknown, probably dark under a covering or dusting of wax; BL of aptera 1.0-1.4 mm.  On Nothofagus obliqua (incl. var. macrocarpa) in Chile.  Biology is unknown; an apparent fundatrix, and a single alata, were collected in November (BMNH collection, leg D. Hille Ris Lambers).

Neosymydobius Baker

Calaphidinae: Myzocallidini

About 14 species superficially resembling Symydobius but associated with nearctic oaks.  Probably the resemblance to Symydobius is a result of convergence due to occupation of similar feeding sites and ant attendance, as Neosymydobius is closer to Myzocallis or Tuberculatus, whereas Symydobius is nearer to Calaphis. They usually feed on bark of terminal twigs, extending onto leaf petioles and along basal parts of main leaf veins.  Quednau (1994) reviewed the genus, described five new species and provided a key, and Quednau (1999) illustrated all the species.

Neosymydobius acutipilosus Quednau   Colour of apterae in life is unknown, probably brownish with banded antennae and black hind legs; BL 1.5-2.0 mm. On Quercus undulata in New Mexico, USA (Quednau 1994).

Neosymydobius agrifoliae (Essig)  Apterae are plump-bodied, dark reddish brown with a paler yellowish Y-shaped mark on dorsum (Essig 1917), banded antennae and dark hind legs; BL 1.6-2.3 mm.  Alatae have dark-bordered wing veins.  Monoecious holocyclic on Quercus spp. (agrifolia, chrysolepis, douglasii) in California.  Apterous males and oviparae occur in October (BMNH collection, leg. R.C. Dickson).

Neosymydobius ajuscanus Remaudière   Apterae are somewhat flattened dorsoventrally, yellowish white with black-banded antennae; BL 1.1-1.8 mm.  Collected on undersides of leaves of Quercus mexicana, and from Q. grisea, in Mexico.  Alatae are present in September-October.  Biology and sexuales are unknown.

Neosymydobius albasiphus (Davis)  Apterae and alatae are shining black or brown with a whitish spinal stripe (Hottes 1926a) and/or white waxy areas around pale siphunculi (Richards 1968), banded antennae and black hind legs; BL 1.4-1.9 mm.  On Quercus spp. (alba, macrocarpa, michauxii, palustris, stellata) in eastern North America.  Oviparae and alate males occur in September-October (Davis 1914 and BMNH collection, leg. R.L. Blackman).  N. canadensis Richards 1965, described from Q. macrocarpa in Ontario, differs only in absence of the white waxy areas around the siphunculi, and possibly in larval chaetotaxy; it was synonymised with albasiphus by Remaudière (1982).

Neosymydobius butzei Remaudière   Apterae are dark shiny chocolate-brown, with black hind legs; BL 1.6-2.1 mm.  On Quercus mexicana in Mexico; Remaudière (1982) also described a variant form (“var. papillata“), which was found on Q. centralis (= Q. laeta) and Q. peduncularis as well as Q. mexicana.  Biology and sexuales are unknown, and it is believed to be anholocyclic.

Neosymydobius chrysolepis Swain  Apterae and alatae are brown, with conspicuously pale siphunculi; BL 1.7-2.1 mm.  On Quercus spp. (chrysolepis, engelmanni, gambelii) in western USA (California, Utah).  Oviparae and alate males occur in October (BMNH collection, leg. G.F. Knowlton).

Neosymydobius luteus Tissot   Apterae are yellowish (only one aptera is known, and they are apparently rare in comparison with alatae).  Alatae have brown head and thorax and pale green-yellow abdomen; BL 1.2-1.4 mm.  On Quercus nigra and Q. cinerea in Florida (Tissot 1932) and Louisiana.  Biology and sexuales are unknown.

Neosymydobius memorialis Hottes & Frison   Apterae have a yellowish head and abdomen and light dusky brown thorax; BL 1.2-1.5 mm.  Alatae have  chestnut-brown head and thorax and lemon yellow abdomen.  On Quercus stellata in Illinois, USA, and also recorded from Q. lyrata in Louisiana (BMNH collection, leg. H.B. Boudreaux), and from Q. texana (Quednau 1999).  Oviparae were collected in October in Illinois (Hottes & Frison 1931), and in mid-November in Texas (Quednau 1999).

Neosymydobius mimicus Hottes   Apterae are shining dusky to dark greenish brown; BL c. 1.6 mm.  Alatae have a green abdomen with darker brownish-green cross-bars.  On Quercus alba in Minnesota, USA (Hottes 1926a), and also recorded from Q. macrocarpa in Manitoba (Robinson 1971).  Biology and sexuales are unknown.

Neosymydobius neomexicanus Quednau   Colour of apterae in life is unknown, probably brownish with antennae almost entirely black and hind tibiae dark except for distal third; BL 1.6-1.9 mm. Alatae are undescribed. On Quercus arizonica in New Mexico, USA (Quednau, 1994). Biology and sexuales are unknown.

Neosymydobius paucisetosus Quednau   Colour in life is unknown; BL of aptera 1.7-2.5 mm. On various Quercus spp. (agrifoliae, chrysolepis, douglasii, dumosa, lobata, wislizeni) in California, USA. Oviparae and an apterous male were collected on Q. chrysolepis in September-October (Quednau 1994).

Neosymydobius peregrinus Quednau   Colour of aptera in life is unknown, probably brownish with banded antennae and dark hind tibiae; BL 1.7-2.0 mm. Alatae are undescribed. On Quercus turbinella and Q. arizonica in New Mexico, USA (Quednau 1994). Biology and sexuales are unknown.

Neosymydobius quercihabitus Miller   Apterae and alatae are dark brown, with conspicuously banded antennae: BL 1.5-2.2 mm.  On Quercus stellata in eastern USA (Maryland, Massachussetts).  Quednau (1971) provided a redescription.  Biology and sexuales are unknown.

Neosymydobius rumorosensis Quednau   Apterae and alatae are chocolate brown, shiny, with banded antennae and a pale middle section to hind tibia; BL of aptera 1.9-2.4 mm (Quednau 1994). On Quercus douglasii and Q. dumosa in California, USA, and on an unidentified Quercus sp. in Mexico (La Rumorosa). Biology and sexuales are unknown.

Neothelaxes  Chakrabarti & Quednau


Three species, two of them on Parthenocissus in north India, related to Thelaxes but with dorsal body hairs arranged in single rows, and differences in the form of R IV+V and in embryonic chaetotaxy. The biology of the Indian species is unstudied, and their apterous viviparae are undescribed (Chakrabarti & Quednau 1996). The generic pacement of the third species, recently described from apterae in galls on Pilea in China (Jiang et al. 2021a), is uncertain.

Neothelaxes parthenocissi Chakrabarti & Quednau    Appearance in life is unknown; BL of alata 1.0-1.5 mm. On Parthenocissus semicordata in Uttar Pradesh, India. Monoecious holocyclic; oviparae were collected in October.

Neothelaxes pileata Qiao   Apterae are dirty pale green or yellowish green, covered in life in white wax powder; BL 1.2-1.3 mm. Alatae are unknown. Found in irregularly spherical galls on leaves of Pilea martinii in China (southern Tibetan plateau).    

Neothelaxes viticola Chakrabarti & Quednau    Appearance in life is unknown; BL of alata 1.6-2.2 mm. On Parthenocissus semicordata in Uttar Pradesh, India.

Neothoracaphis Takahashi

Hormaphidinae: Nipponaphidini

About ten species mainly known from Quercus in east Asia.  Apterae are very small, black, sclerotic, and lack siphuncular pores, and their legs have much reduced tarsi, with the fore- and mid-legs concealed but the hind legs usually projecting a little behind the body.  Host alternation and gall formation on Distylium is only established for one species (yanonis).  Several species seem to be anholocyclic on evergreen Quercus, with the apterous vivipara the only known morph.  Most species were originally described in Thoracaphis.  Takahashi (1958d) and A.K. Ghosh & Raychaudhuri (1973a) gave accounts of the genus and keyed species under Microthoracaphis, which is here included within Neothoracaphis.  Chen et al. (2011c) reviewed the genus, Tao (1966) keyed the species in Taiwan, Qiao et al. (2018) reviewed and keyed the Chinese species, and A.K. Ghosh (1988) reviewed the species in India. Aoki et al. (2019b) examined mitochondrial DNA sequences of four species in Japan.

Neothoracaphis elongata (Takahashi)  Apterae are elongate oval, black, with pale hind legs projecting behind body; BL 0.45-0.65 mm. They live throughout the year on foliage of evergreen Quercus spp. in Japan (Takahashi 1958d) and China (Yunnan; Chen et al. 2011c). Specimens collected from Quercus (= Cyclobalanopsis) edithiae and Q. neglecta (= Q. myrsinifolia) in Hong Kong, which in life have five pairs of distinctly curved wax rods (Martin & Lau 2011), may also be either this species or N. saramoensis. (N.B. not to be confused with Parathoracaphis elongata (Takahashi), which is a quite different species.)  

Neothoracaphis garwhalensis Chakrabarti & Raha   Apterae are oval, black, with pale hind legs projecting a little behind body; BL 0.7-0.8 mm.  In ant-attended colonies on leaves of Quercus spp. (incana, serrata) in Uttar Pradesh, India (Chakrabarti & Raha 1985).  The life cycle is unknown.

Neothoracaphis glaucae (Takahashi)  Apterae are dimorphic (Aoki et al. 2019b). The larger form (“L-type”) is oval, indented at frontal margin, black, with much white wax mid-dorsally, and with yellow hind legs protruding behind body; BL 0.65-0.85 mm.  The smaller form of aptera (“S-type”) is more elongate oval, shining brown-black with a prominent spinal ridge; BL 0.4-0.6 mm. Both phenotypes live on the leaves of Quercus glauca in Japan. Apterae of the smaller type are common throughout the year, while those of the larger type occur in small numbers in summer and early autumn. Alatae (BL 1.0-1.2 mm) develop in November to January, from nymphs that are larger than the adult apterae and covered in numerous long semi-transparent wax filaments. Possibly these alatae are sexuparae, but the life-cycle is still unresolved (Aoki et al. 2019b).

Neothoracaphis quercicola (Takahashi)  Adult apterae are black, broadly oval, without wax; BL c. 0.9 mm (Takahashi 1924a; as Astegopteryx).  Immatures are yellow and secrete much wax.  Alatae have black head and thorax and dusky yellowish brown abdomen.  Living parthenogenetically all year on undersides of leaves of Quercus variabilis in Taiwan (records from Q. serrata may be in error), and now recorded also forming large colonies on upper and lower surfaces of leaves of  Q. acutissima in China (Hubei, Yunnan, Guangxi; Chen et al. 2011c).  Alatae are produced in November-March (Takahashi 1931a).

Neothoracaphis querciphaga (Takahashi)  Apterae are black, narrowly oval, with protruding hind legs; BL 0.4-0.7 mm.  Living all year on foliage of Quercus myrsinaefolia in Japan (Takahashi 1958d).

Neothoracaphis saramaoensis (Takahashi)  Apterae are black, elongate oval, indented on frontal margin, with white wax dorsally, including a pair of long whitish wax “horns” arising from middle of anterior part of prosoma, and also “a wide, very thin whitish secretion behind the hind end” (Takahashi 1935c).  Hind legs projecting posteriorly, pale brown.  BL 0.5-0.7 mm.  On undersides of leaves of Quercus variabilis in Taiwan. It has now also been reported to occur on Lithocarpus spp. in China (Yunnan; Chen et al. 2011c). The small S-type morph of N. glaucae (q.v.) on Q. glauca in Japan is almost indistinguishable and was originally thought to be this species (Takahashi 1958d).

Neothoracaphis semicarpifolia Chen, Yang & Qiao  Apterae are oval, flattened, strongly sclerotised, with white wax mid-dorsally and fine bluish-white wax threads around the entire margin; BL 0.6-0.8 mm. On undersides of leaves of Quercus semecarpifolia at an altitude of 3330m in Tibet (Chen et al. 2011c). The life cycle is unknown. 

Neothoracaphis sutepensis (Takahashi)  Apterae are broadly oval, black with much white wax mid-dorsally, and black hind legs protruding behind body; BL 0.5-0.7 mm.  On undersides of leaves of a tree of the Fagaceae (probably Quercus sp.) in Thailand (Takahashi 1941, as Thoracaphis). Apterae believed to be of this species were redescribed from unidentified Quercus spp. in Taiwan by Tao (1969, as Microthoracaphis)) and in India (Manipur, Meghalaya) by A.K. Ghosh (1988). The life cycle is unknown.

Neothoracaphis tarakoensis (Takahashi)  Apterae are broadly oval with frontal margin indented, black, with much white wax mid-dorsally, and black hind legs protruding behind body; BL 0.9-1.0 mm.  On undersides of leaves of Quercus spinosa in Taiwan (Takahashi 1937b).  The life cycle is unknown.  The distinction between this species and N. yanonis, apart from size, is unclear.

Neothoracaphis yanonis (Matsumura)  Host-alternating between Distylium racemosum and deciduous Quercus spp. (dentata, serrata) in Japan and China.  Small, prominent galls develop in April-May, protruding from both sides of Distylium leaves (fig. 130B, and Moritsu 1983, as Nipponaphis yanonis). All second generation are alatae, with black head, thorax, antennae and legs, yellowish brown abdomen and dark-bordered wing veins; BL 1.2-1.6 mm.  Apterae on undersides of Quercus leaves are black with white wax mid-dorsally, without marginal wax, with black hind legs projecting behind body (fig. 91K); BL 0.5-0.6 mm.  Alate sexuparae return to Distylium in October-November and produce abundant sexuales on undersides of leaves (Takahashi 1958d, Ngakan & Yukawa 2020). The male genitalia were described and illustrated by Wieczorek et al. (2012). Shen (1986) studied the life cycle and chemical control measures in China, and Ito (1984) recorded invasion of galls by the kleptoparasitic moth Nola innocua in Japan. N. hangzhouensis Zhang (in Zhang & Zhong 1982b), is a synonym (Chen et al. 2011c).

Neotoxoptera Theobald

Aphidinae: Macrosiphini

A genus for six Myzus-like aphids with clavate siphunculi and dark-bordered wing-veins, probably of oriental origin. Three species are now widespread and at least the adventive populations seem to be entirely anholocyclic; Peréz Hidalgo et al. (2011b) provided a key to separate these species. Three species in east Asia host-alternate with Caprifoliaceae as primary hosts, as in the somewhat similar genus Rhopalomyzus. Accounts are available for Australia (Carver 1980), Japan (Miyazaki 1971, Sorin 1971) and Korea (Lee et al. 2002c).

Neotoxoptera abeliae Takahashi    Apterae (fundatrices) are green, BL c. 2 mm. Alatae have black head, thorax, antennae and siphunculi, and broadly banded wing veins. On Caprifoliaceae (Abelia, Lonicera) in spring, apparently migrating in the second generation to an unknown secondary host. In Japan, Korea and east Siberia. Fundatrices provisionally assigned to this species were found on Abelia triflora at 2,600m in Pakistan (Naumann-Etienne & Remaudière 1995). Pashtshenko (1988a) recorded oviparae from Weigela praecox, but these may have been N. weigeliae (q.v.). The type specimens were collected from Abelia spathulata; the host was erroneously given as Deutzia gracilis in the original description.

Neotoxoptera formosana (Takahashi)    Apterae are shining magenta-red to dark reddish brown or almost black (see influential points/gallery); BL 1.6-2.3 mm. Alatae are very dark red to black with the wing-veins heavily black-bordered, the borders being of rather constant width along the lengths of the veins. On leaves of Allium spp., or on bulbs in store. In east and south-east Asia, Australia, New Zealand, Hawaii, Panama, Guadaloupe (Etienne & Champoiseau 2011), South America (Argentina, Brazil, Chile, Columbia, Peru, Venezuela), North America, St Helena, Canary Islands (Peréz Hidalgo et al. 2011b), and throughout much of Europe; Norway, Finland, UK (Halstead 2000), Germany, Italy (Barbagallo & Ciampolini 2000), France and Netherlands (Piron 2010a). Apparently completely anholocyclic everywhere. Hori & Komatsu (1997) studied repellency of rosemary oil and its components to this aphid, and Hori (2007) analysed the attractive compounds present in Allium spp. 2n=12.

Neotoxoptera oliveri (Essig)  Plate 19d  (Fig.59b)   Apterae are dark-red to almost black; BL 1.1-2.0 mm. Alatae have wing veins heavily black-bordered, the borders widening out at base and apex of each vein. The host range of N. oliveri is remarkably similar to that of Myzus ascalonicus; most commonly on certain Alliaceae (Allium), Caryophyllaceae (Stellaria) and Violaceae (Viola), but also recorded from plants in several other families. Almost cosmopolitan; recorded from southern Europe (France, Portugal, Madeira, Serbia), Africa, Pakistan, Korea, Java, Australia, New Zealand, western USA, Central America (Mexico, Panama, Bermuda, Costa Rica) and South America (Argentina, Brazil, Chile, Peru). Apparently anholocyclic everywhere; Naumann-Etienne & Remaudière (1995) recorded a single male from Pakistan. It was not possible to distinguish N. sungkangensis Hsu from N. oliveri on the basis of the published description.

Neotoxoptera violae (Pergande)  (Fig.59a)  Apterae are purplish or dark wine-red; BL 1.0-1.7 mm. Alatae are dark wine-red with broadly banded wing-veins. On undersides of leaves of Viola spp. In North and South America, Hawaii, Australia, New Zealand, China, Taiwan and Korea, and southern Europe (Spain, Melía 1991; Italy, Barbagallo & Cocuzza 1998; Canary Islands (Peréz & Oromí 2004); and most recently France, Germain & Deogratias 2008). African records are all (as yet) referable to N. oliveri. No sexual morphs are known. 2n=12 (this record was erroneously attributed to N. oliveri in Blackman & Eastop 2000).

Neotoxoptera weigeliae Lee and Seo  (Fig.61d)   Apterae (fundatrices?) are greenish; BL 1.0-1.3 mm. Alatae are dark brown with large black dorsal abdominal patch and forewings with dark pterostigma (cf. Micromyzus diervillae) and diffusely bordered veins. On Weigela spp. in Korea, Japan, and possibly east Siberia (Pashchenko 1988a, as oviparae of  N. abeliae). Heteroecious holocyclic, probably migrating in the the second generation to an unknown secondary host.

Neotoxoptera yasumatsui Sorin    Apterae are reddish to deep brown; BL c.1.4-1.5 mm. Alatae are blackish brown with a large central dorsal abdominal patch and broadly banded wing-veins. On Kalimeris lautureana (as Aster lautureanus) in Japan, forming small colonies that distort apical parts of subterranean stems and young leaves at or just below ground level (original description). Heteroecious holocyclic; migration to the primary host Weigela praecox is recorded in east Siberia (Pashchenko 1988a). Apterae collected on Artemisia vulgaris var. indica in Japan were described as a subspecies, N. yasumatsui artemisiae Sorin (2001).


Neuquenaphis Blanchard


A South American genus of about fifteen species associated with Nothofagus, closely related to Sensoriaphis in Australia and New Zealand. Apterae usually have rows of conspicuous pigmented finger-like hair-bearing spinal, marginal, and sometimes also pleural, processes. Alatae generally have fewer, less conspicuous processes.  Quednau & Remaudière (1994b) revised the genus, provided keys to apterae and alatae, and sorted out much of the nomenclatural confusion. Quednau (2010) reviewed the world fauna and provided illustrations of all available morphs. Nieto Nafría et al. (2019b) provided keys to the known apterae and alatae. Apparently all the described species are holocyclic, but apart from that their biology is virtually unknown.  Several species seem to feed on more than one species of Nothofagus. Blackman et al. (2003) studied karyotype variation and Gaete-Eastman et al. (2004) studied genetic diversity in relation to diet breadth.

Neuquenaphis aurata Mier Durante & von Dohlen   Colour of apterae in life is yellow-gold to yellowish straw, with dusky dorsal processes; BL of apterae 2.3-3.2 mm. Alatae are creamy yellow with dark brown head and thorax (Nieto Nafría et al. 2019b). On Nothofagus betuloides in southern Argentina (Santa Cruz, Tierra del Fuego).  Specimens on slides are very similar to N. blackmani.

Neuquenaphis blackmani Nieto Nafría & Brown   Colour of apterae in life pale green; BL of apterae 2.1-3.0 mm. Alatae are pale green with brown head and thorax (Nieto Nafría et al. 2019b). On leaves of Nothofagus dombeyi and N. nitida (both trees are evergreens) in Chile (Los Rios, Los Lagos).  2n=12 (Blackman et al. 2003, as Neuquenaphis sp. A)

Neuquenaphis bulbicauda Hille Ris Lambers   Colour of apterae dark grey-brown, immatures green; BL of apterae 2.2-3.1 mm. On leaves of Nothofagus dombeyi in Chile (Hille Ris Lambers 1968). Alatae are unknown. [The alata and sexual morphs described by Hille Ris Lambers under bulbicauda are other species; the alata is N. valdiviana, and the sexual morphs are those of N. neobulbicauda.] 2n=14 (Blackman et al. 2003).

Neuquenaphis (Spicaphis) chilensis Essig  (=Spicaphis michelbacheri Essig 1953b p. 69)   Colour in life is unknown, but probably rather dark with a covering of powdery wax; BL of apterae and alatae 2.2-3.1 mm.  Alatae have dark-bordered wing veins.  On leaves of Nothofagus dombeyi in Chile.  2n=10 (Blackman et al. 2003).

Neuquenaphis edwardsi Laing   Colour in life is unknown, probably rather dark with a covering of powdery wax; BL of apterae and alatae 1.8-2.6 mm.  On various Nothofagus spp. (antarctica, glauca, obliqua) in Argentina and Chile, including Tierra del Fuego.  Apterous oviparae and alate males occur in April-May (BMNH collection, leg. G. Monsalve). The male genitalia were described and illustrated by Wieczorek et al. (2011). 2n=12  (Blackman et al. 2003).

Neuquenaphis (Spicaphis) essigi Hille Ris Lambers   Appearance in life is unknown, probably rather dark, perhaps with greyish wax.  Apterae are very broadly oval;  BL of apterae and alatae 2.0-2.4 mm.  On Nothofagus obliqua (incl. var. macrocarpa) in Chile (Hille Ris Lambers 1968).  Sexuales are unknown. 2n=12 (Blackman et al. 2003).

Neuquenaphis michelbacheri Essig (1953 p. 66)   Apterae are unknown, alatae have dark spots at ends of wing veins; BL of alata 1.7-2.3 mm. On Nothofagus dombeyi in Chile.

Neuquenaphis neobulbicauda Quednau & Remaudière  Appearance in life is unknown; BL of aptera 2.1-2.3 mm. Alatae are undescribed. On Nothofagus dombeyi in Chile. Apterous oviparae  (BL 1.5-1.9 mm) and and an alate male were collected in February (Hille Ris Lambers 1968, as bulbicauda, redescribed by Quednau & Remaudière (1994b). 

Neuquenaphis palliceps Hille Ris Lambers  Appearance of apterae in life is unknown; larger and probably paler than most other described species. Alatae pinkish with three short bars of wax on anterior abdomen; BL of apterae and alatae 2.2-2.9 mm.  On Nothofagus spp., especially N. dombeyi, in southern Argentina and southern Chile.  Apterae collected in November-December were possibly fundatrices (Hille Ris Lambers 1968), and an alate male was collected in April (BMNH collection, leg. G. Monsalve). 2n=6 (Blackman et al. 2003).

Neuquenaphis ramirezi Nieto Nafría & Ortego   Apterae are dark green to greenish brown; BL 1.7-2.3 mm. Other morphs are unknown. On Nothofagus pumilio, and (less  frequently) N. antarctica in southern Chile ((Nieto Nafría et al. 2019b). 2n=16 (Blackman et al. 2003, as Neuquenaphis sp. B). 

Neuquenaphis schlingeri Hille Ris Lambers   Appearance of apterae in life is unknown, probably rather pale. Alatae pale brown with white wax markings and darkish wing veins; BL of apterae and alatae 1.4-1.8 mm.  On Nothofagus spp. (alessandrii, glauca, obliqua) in Chile (Hille Ris Lambers 1968 and BMNH collection).  Alate males and apterous oviparae occur in April-May (BMNH collection, leg. G. Monsalve). 2n=12 (Blackman et al. 2003).

Neuquenaphis sensoriata Hille Ris Lambers  Plate 8 a,b   Apterae in life dark brownish green, with pale dorsal processes and a dusting of wax. Alatae have basal part of ANT III conspicuously pale compared with dark rest of antenna, dark but unbordered wing veins, white wax markings on head and thorax, and two rows of white patches on dorsal abdomen including spinal processes; BL of apterae and alatae 1.7-2.1 mm.  Immatures are shiny red-brown (slide label, BMNH collection).  On Nothofagus spp. (especially glauca, obliqua) in Chile.  Fundatrices occur in November (BMNH collection, leg. D. Hille Ris Lambers), sexuales are not recorded. 2n=16 (Blackman et al. 2003).

Neuquenaphis similis Hille Ris Lambers   Apterae are elongate oval, of unknown colour but probably rather pale; BL 2.6-2.8 mm.  Alatae are unknown.  In Chile; described from Nothofagus obliqua (Hille Ris Lambers 1968), but all subsequent collections, including possible fundatrices in November, were from N. pumilio (BMNH collection, leg. D. Hille Ris Lambers). 2n=14 (Blackman et al. 2003).

Neuquenaphis staryi Quednau & Remaudière  Appearance in life is unknown; BL of alata 1.4-1.9 mm (BL of aptera unrecorded). On Nothofagous alessandri in Chile (Quednau & Remaudière 1994b). Oviparae occur (along with apterous and alate viviparae) in late May; Ramírez et al. (2008) studied leaf colour preferences of autumn populations.  2n=14 (Blackman et al. 2003).

Neuquenaphis valdiviana Carrillo  Apterae according to the original description are lemon yellow with rows of dark finger-like processes; BL c.1.7 mm.  On Gunnera chilensis (= tinctoria) in Valdivia, Chile (Carrillo 1980). Attempts to transfer this aphid to various Nothofagus spp. were unsuccessful (Carrillo 1980), and subsequent collections have been made from Gunnera, but it has since also been recorded from N.  dombeyi (Quednau & Remaudière 1994b). 2n=6 (Blackman et al. 2003).

Nevadaphis Drews

Aphidinae: Aphidini

One little-known species in North America related to Aphis, with greatly reduced siphunculi and cauda.

Nevadaphis sampsoni Drews    Apterae are dirty yellow to reddish brown, often mottled; BL c. 1.9 mm. In colonies on roots of Seriphidium tridentata, sometimes ant-attended, in USA (Nevada, California). Other morphs are unknown.

Nevskya Ossiannilsson

Calaphidinae: Saltusaphidini

Three species closely related to Iziphya, but with accessory rhinaria on ANT VI placed close to the primary rhinarium, no modification of the basal dorsal hairs on the tibiae, and dorsal hairs very numerous, short and mushroom-shaped. Nevskya tuberculata (Zhang & Zhang) differs from either of the species below in having a pair of tubercles posteriolaterally on the pronotum, and a longer antennal terminal process.  Quednau (2010) reviewed the genus, keyed the species and illustrated all available morphs. [Nevskya Ossiannilsson 1953 has been reinstated as the valid name for the genus in place of Nevskyella Ossiannilsson 1954; see Quednau 2010 p.70 (footnote).]

Nevskya fungifera Ossiannilsson  (= Subsaltusaphis sinensis Zhang, Zhang & Zhong 1995, = Nevskyella similifungifera Qiao & Zhang 2004; synonymies by Quednau 2010)  Apterae are yellow or brownish yellow, sometimes with dark paired dorsal patches on metathorax and abdomen, or uniformly brownish; BL c.1.6-1.7 mm. Alatae have bordered wing veins with dark spots at their apices. On Carex spp. and Cyperus spp., living on all above-ground parts of the plant. Described from Sweden, and also found in Kazakhstan (Kadyrbekov 2009b, as Nevskyella fungifera) and China (as Subsaltusaphis sinensis and Nevskyella similifungifera). Monoecious holocyclic, with oviparae and apterous males in Sweden in September, and fundatrices in May (original description).

Nevskya meridionalis (Hille Ris Lambers & van den Bosch)    Apterae are creamy to yellowish, with paired dark dorsal patches; BL 1.5-1.7 mm.  On undersides of leaves of  Carex halleriana in France, on C. praecox in Slovakia (Holman 2009), and alatae have been found on Carex sp. in Italy.  Monoecious holocyclic, with oviparae and apterous males in October (original description).

Nevskya tuberculata (Zhang & Zhang)   Described from a single alata trapped in China, but Quednau (2010) recorded and illustrated the apterous morph, collected from a Carex sp. in South Korea.  Biology is unknown.

Nietonafriella Ortego

Aphidinae: Macrosiphini

One South American species resembling Uroleucon (Lambersius) but with an enlarged clypeus.

Nietonafriella euclypteata Ortego    Apterae are greenish yellow to dark green, sometimes with a darker green, orange or reddish brown band between and around bases of siphunculi; BL 1.6-2.2 mm. Alatae have 11-19 secondary rhinaria on ANT III. On apices of young stems of Eupatorium patens in western Argentina. Monoecious holocyclic, with apterous males (Ortego 1998b).

Nigritergaphis Zhang, Lou & Qiao

Aphidinae: Macrosiphini

One species described from Isodon in China with long stiff dorsal hairs, extensive dark dorsal sclerotisation and short siphunculi and cauda.

Nigritergaphis crassisetosa Zhang, Lou & Qiao   Apterae are shining black, BL 1.4-1.6 mm, immatures are green. On Isodon eriocalyx, curling, twisting and blistering the leaves, in Yunnan province, China (B. Zhang et al. 2013). Alatae have dark dorsal abdominal cross-bands and secondary rhinaria distributed ANT III 18-34, IV 7-12 and V 1-3. Apterae collected in October had 5-segmented antennae, and those collected in December had 6-segmented antennae. No sexual morphs were observed.

Nippodysaphis Hille Ris Lambers

Aphidinae: Macrosiphini

One species in Japan with a short cauda, resembling Dysaphis (Pomaphis), but lacking marginal or spinal tubercles.

Nippodysaphis deutziae (Hille Ris Lambers)  (Fig.32d)  Appearance in life is unknown; BL (fundatrix) c.2.35 mm. On Deutzia crenata in Japan. Spring populations heavily curl leaves (Miyazaki 1971), and migrate in the second generation to an unknown secondary host. Emigrant alatae have a dark dorsal abdominal patch, and secondary rhinaria distributed III 3-10, IV 0-5. Presumed gynoparae from Japan and Korea in the BMNH collection have barely coalescing transverse dorsal abdominal cross-bands and secondary rhinaria distributed III 19-26, IV 8-11, V 2-5.

Nippolachnus Matsumura

Lachninae: Tuberolachnini

An oriental genus of 4-5 species characterized by the absence of an ocular tubercle, the triommatidia being below the compound eyes.  They live on woody Rosaceae, mainly Eriobotrya and Pyrus.  A.K. Ghosh (1982b) reviewed the species in India, and Kanturski et al. (2018d) reviewed the genus, used DNA evidence to take one species out of synonomy, made SEM studies of the sensilla, and provided a key to species.

Nippolachnus bengalensis Basu & Hille Ris Lambers   Apterae are long-bodied, pale green; BL 2.6-3.3 mm.  Alatae have a variable sclerotic pattern on the dorsal abdomen.  On undersides of leaves of Eriobotrya dubia in West Bengal, India (Basu & Hille Ris Lambers 1968); also recorded from Pyrus pashia and Photinia arguta in Meghalaya, and from cultivated pears in Bhutan (Agarwala 1983).  Specimens from Pyrus granulosa (= Sorbus corymbifera) in Malaya (Takahashi 1950; as N. piri) also appear to be this species.  The life cycle and sexual morphs are unknown.

Nippolachnus himalayensis (van der Goot)  Apterae are broadly oval, dirty greyish to brick red with whitish intersegmental bands of wax, and mainly black legs: BL 3.5-5.5 mm.  In large colonies on undersides of leaves of Eriobotrya petiolata in West Bengal, India (as eriobotryae; Basu & Hille Ris Lambers 1968).  The life cycle and sexual morphs unknown.

Nippolachnus micromeli Shinji  Apterae in life are very pale, almost colourless; BL 2.2-3.0 mm. On udersides of leaves of Rhaphiolepis and Sorbus in East Asia (Japan, Korea, China). This species has for a long time treated as a synonym of N. piri, but there is now DNA and morphological evidence that it should be regarded as a separate species (Kanturski et al. 2018). Sexual reproduction and overwintering as eggs occur in Japan on Rhaphiolepis; Otake (1995, as N. piri) studied aspects of its overwintering and spring colony development on this host.

Nippolachnus piri Matsumura  Plate 15 a,b Apterae are long-bodied, pale green with some bluish white dorsal wax, with pale brown legs and antennae; BL 2.3-3.7 mm.  On the undersides of leaves, feeding along main veins, of Eriobotrya japonica, and also recorded from Pyrus spp. There are also Japanese records, possibly of vagrants of either this species or N. micromeli, from Castanea crenata and Betula platyphylla (Higuchi & Miyazaki 1969). Aphids on Raphiolepis and Sorbus identified in the literature as N. piri should probably now all be referred to N. micromeli (Kanturski et al. 2018d).  In India, China, Taiwan, Vietnam, Laos and Japan.  Oviparae and alate males occur on E. japonica in Japan in October-November (Takahashi 1923).  In India (West Bengal, Sikkim) populations are apparently anholocyclic and host specific to Pyrus communis, on which large damaging populations may build up in summer months (M.R. Ghosh & Raychaudhuri 1981).  Neonippolachnus betulae Shinji, described from Betula in Japan and not since recognised, is probably this species, and Nipponaphis xitianmushanus, on E. japonica in China, may or may not be distinct (Zhang & Zhong 1982b).

Nipponaphis Pergande

Hormaphidinae: Nipponaphidini

About 15 east Asian species, some with host alternation between galls on Distylium and Fagaceae, others apparently anholocyclic on Fagaceae, Lauraceae and Moraceae. The galls of the species studied in detail (N. distyliicola and N. monzeni) take more than a year to mature, and the complete life cycle can take 2-5 years. Apterae on secondary hosts are aleyrodiform, with the strongly sclerotised prosoma bearing pustulate ornamentation.  Alatae have smoky wings.  Qiao et al. (2018) reviewed and keyed two of the Chinese species, Takahashi (1962) reviewed the species on Distylium in Japan, Sorin (1987) illustrated differences in embryonic chaetotaxy, and Aoki et al. (2015) tabulated data on chaetotaxy of first instar nymphs. Many of the Japanese species are illustrated by Moritsu (1983).  A.K. Ghosh & Raychaudhuri (1973b) reviewed and keyed the genus; the three species they placed in subgenus Pseudonipponaphis are here transferred to Schizoneuraphis. Aoki et al. (2015) reviewed the species forming green galls on Distylium in Japan, applying molecular and morphological methods.

Nipponaphis brevipilosa Noordam   Apterae are pale brown with antennae, legs and margins of abdominal plate yellowish; BL 1.3-1.6 mm. In dense colonies on young and older twigs of Castanopsis argentea in Java (Noordam 1991). Alatae were collected in May. The life cycle is unknown.

Nipponaphis distychii Pergande  (= Nipponaphis litseae Takahashi)   Large galls are produced on the stems of Distylium racemosum in Japan and Korea. They are of two types; (1) globular or fig-shaped, yellowish green with green longitudinal vein-like stripes (Moritsu 1983, Paik 1972) or (2) spindle-shaped and pale green (fig. 130H and Sorin 1987b, as N. litseae). Aphids forming the latter type of gall were previously thought to be a different species, N. litseae, but molecular analysis has now shown that the two names should be treated as synonyms (Aoki et al. 2015). Alatae (secondary rhinaria distributed ANT III 39-48, IV 20-25, V 10-15) emerge in autumn; according to the original description (Pergande 1906, quoting Kuwana) they have black head and pterothorax, purplish yellow prothorax and dark purplish brown abdomen; BL 1.9-2.2 mm.  The wings are only slightly dusky – more so at their bases.  They migrate to Neolitsea sericea, where the blackish brown, almost circular apterae feed on the branches (Takahashi 1959b, as N. litseae). Records of N. distyliicola and N. coreana from Neolitsea in Korea may be referable to this species.  Monzen’s (1954) account under the name distychii refers to N. monzeni (Takahashi 1962a).

Nipponaphis distyliicola Monzen   Plate 3 e,f    Closed green fig-shaped or globular galls on the twigs of Distylium racemosum in Japan. Occasionally the galls arise from deformed leaves (Kurosu & Aoki 2021). Alatae (BL 1.5-2.5 mm, antennal rhinaria distributed III 21-39, IV 10-19, V 8-14) emerge in autumn and fly to Quercus spp. (acuta, glauca, crispula, miyagii, myrsinifolia).  Apterae on branches and shoots of Quercus are almost circular, black; BL 1.2-2.2 mm.  Alate sexuparae (BL 2.5-2.7 mm) return to Distylium in April-May, but anholocyclic populations also persist on Quercus (Takahashi 1962a; Sorin 1958, as gigantea).  The eggs produced in spring on Distylium soon hatch and the fundatrices form galls on lateral buds, but these remain small until the following year, so that the complete life cycle takes two years (Kurosu & Aoki 1998b). Ito & Hattori (1983) reported predation by Nola innocua, and Kurosu et al. (1995) described first instars (not sterile soldiers) of N. distyliicola attacking lepidopteran and dipteran larvae introduced into galls.  The male genitalia were described and illustrated by Wieczorek et al. (2012). N. coreana on Quercus myrsinaefolia in Korea (Paik 1965) appears to be a synonym, to judge from paratypes in the BMNH collection.

Nipponaphis ficicola Hille Ris Lambers & Takahashi   Apterae are very broadly oval, wrinkled, dark purple to blackish in life; BL 1.0-1.3 mm.  On the bark of about 1 cm diameter branches and twigs of Ficus benjamina and Ficus sp. in Java (Hille Ris Lambers & Takahashi 1959).  Apterae were collected in June-November.  The life cycle is unknown.

Nipponaphis javanica Noordam   Apterae are brownish, with some wax powder; BL. c. 1.8 mm. In dense colonies on young shoots, and on under and upper sides close to mid-ribs of youngest leaves, of Castanopsis acuminatissima in Java (Noordam 1991).  Alatae were collected in August and January. The life cycle is unknown.

Nipponaphis karatanei (Sasaki)  This species, described from Magnolia fuscata (= M. figo var. figo) in Japan (Sasaki 1936). was placed in Nipponaphis by Monzen (1954), but is unlikely to belong to this genus.  The association with Magnolia is also doubtful, because the galls described appear to be those of a psyllid that lives on Persea, and the description of the aphid has many inconsistent features making its taxonomic position uncertain.

Nipponaphis loochooensis Sorin   (= N. machili? of Sorin 1987)   Galls on twigs of Distylium racemosum in Okinawa, Japan are glossy, yellowish green, fig-shaped, 3.3-5.9 cm long and c. 3 cm wide (Sorin 1996).  Alatae emerge in late January and migrate to Machilus spp. (thunbergii, japonica) larvipositing on the undersides of the leaves. Apterae on the secondary host are blackish brown, box-like, almost circular; BL c. 1.8 mm, and feed on young shoots and twigs. On M. japonica, which may be the preferred host, they also colonise the undersides of leaves, but on M. thunbergii they are only found on the twigs (Aoki et al. 2019a). They differ from other Machilus-feeding Nipponaphis (machili, machilicola) in having fewer and shorter dorsal hairs and lacking hairs on perpendicular margins of the prosoma. Alate sexuparae return to Distylium in April. Sorin (1996) provided descriptions of all morphs except the fundatrix.

Nipponaphis machili (Takahashi)  Apterae are blackish brown, almost circular; BL c. 1 mm. Along the mid-ribs on the undersides of leaves of Machilus sp. in Taiwan (Takahashi 1933b), and also reported to occur on Phoebe sp. in Sichuan, China (Tao 1999).

Nipponaphis machilicola (Shinji)  (= N. amamiana Takahashi; Aoki et al. 2019a)   Green, globular galls are produced on twigs of Distylium racemosum in Okinawa, Japan (Aoki et al. 2015). Alate emigrants from these galls have BL 1.7-2.0 mm and secondary rhinaria distributed ANT III 34-44, IV 16-22, V 13-19, and fly to form colonies on twigs and branches of Machilus spp. (thunbergii, japonica). Apterae on the secondary host are almost circular, dorsally flattened, dark bluish brown with pale yellow markings (see Moritsu 1983 or Aoki et al. 2019a); BL 1.5-2.2 mm. Populations on Machilus are also recorded from Honshu (Kanagawa Prefecture), where they may persist on the secondary host throughout the year. Alate sexuparae produced in March in Okinawa and in late May in Kanagawa were described by Aoki et al. (2015). [But for the feeding site, body size and greater variation in shape of dorsal pustules, these apterae strongly resemble those of N. machili, and its taxonomic status requires confirmation.  N. amamiana Takahashi (1962a) appears to be a synonym.]

Nipponaphis manoji Ghosh & Raychaudhuri   Apterae are almost circular, dorsally flattened, dark brown, not strongly sclerotised; BL 1.2-1.4 mm (A.K. Ghosh & Raychaudhuri 1973b).  In large, ant-attended colonies on leaves and young stems of Litsea spp. (?corymbosa, monopetala) in West Bengal, India.  Aphids on Machilus yunnanensis in China (Yunnan) have also been identified as this species (Huang et al. 2012). The life cycle is unknown; alatae (sexuparae?) were collected in February.  A Nipponaphis species described (but not named) from apterae on Litsea polyantha (= monopetala) in Nagaland (Raha & Raychaudhuri 1981) is very similar, except for the  apparent lack of posteriomedial hairs on abdominal tergite 7.

Nipponaphis minensis Zhang    The host of this species, described from China (Fujian), is not given in the original description, but is listed as a Quercus sp. by Qiao et al. (2018), who provide a redescription.

Nipponaphis monzeni Takahashi   Galls on  Distylium racemosum in Japan are large, hard, variously shaped, pyriform, conical or almost globular, pale brown, maximum diameter 2-6 cm (figs 130E-G; Sorin 1958, Moritsu 1983, Kurosu & Aoki 2009). Self-sacrificing repair of damage to the gall by first instar larvae was described by Kurosu et al. (2003), and the longer-term consequences of gall repair were studied by Kutsukake et al. (2009). Alatae (BL c. 3.0 mm) emerge from a round hole in the side of the gall in November-December and migrate to evergreen Quercus spp. (glauca, myrsinaefolia, phillyraeoides), Castanopsis sieboldii, and also Lithocarpus edulis. [A.K. Ghosh (1988) recorded this species from Machilus (= Persea) in northern India, but evidently had a Sinonipponaphis sp.] The blackish brown oval apterae (fig. 91A) (BL 1.5-1.7 mm) live on twigs and branches of the secondary hosts through the winter. Sexuparae return to Distylium in April. Eggs hatch in May (Takahashi 1962a), and galls are initiated in June, but they remain small for at least 21 months and may not grow to maturity until the spring of the third, fourth or fifth year (Kurosu & Aoki 2009). No colonies remain on secondary hosts during the summer months.

Nipponaphis multisetosa Noordam   Apterae are brownish; BL 1.3-1.7 mm. On young and older twigs of Castanopsis javanica in Java (Noordam 1991). Other morphs and life cycle are unknown.

Nipponaphis semiglabra Noordam   Apterae are brown with a transparent coat of wax, the prosoma having a paler brown spinal band, two paler thin transverse lines along the posterior margin of the pronotum and abdominal tergite 1, and seven pairs of small black spots; BL 1.2-1.5 mm. In small numbers on older twigs of Lithocarpus bennettii in Java (Noordam 1991). Other morphs and life cycle are unknown.

Nudisiphon Chakrabarti & Bhattacharya

Aphidinae: Macrosiphini

A genus for one Indian species with flangeless siphunculi. Apterae have a peculiar cauda with a curved, horn-like hairless apex, whereas in alatae the cauda is elongate triangular with a hairy apex.

Nudisiphon folisacculata (Kumar & Burkhardt)    Apterae are presumably pale, wax-covered; BL 2.2-2.4 mm. Alatae have numerous secondary rhinaria distributed over half the circumferences of  ANT III-V. On Spiraea sp. in India (Himachal Pradesh), causing leaves to fold upwards and parallel to the midrib, forming “fluffy sac-like pseudogalls” (original description, as Xenosiphonaphis). An alate male was collected in August. Also recorded from the same region on Indigofera (?) sp. (Chakrabarti & Bhattacharya 1982, as Nudisiphon chitinicauda).

Nurudea Matsumura

Eriosomatinae: Fordini

Three species forming flower-like galls on Rhus in east Asia, and migrating in autumn to unknown secondary hosts, likely to be mosses.  Tsai & Tang (1946) provided a review of the Chinese species, and Tao (1970) keyed the species in Taiwan (although his synonymy is not followed here). Two of the species considered under this name in Blackman & Eastop (1994) have been transferred back to the genus in which they were originally described (Floraphis meitanensis and F. choui).

Nurudea ibofushi Matsumura  (= sinica Tsai & Tang)  Galls are pale yellowish brown with a pinkish tinge, irregular in shape, maximum diameter 4.5-5.5 cm; on leaves of Rhus chinensis (= semialata) in China and Japan.  Alatae (BL c.1 mm) with brown head and thorax and pale yellow-green or yellow-brown abdomen emerge in autumn (Tsai & Tang 1946; as N. sinica). Molecular studies (Ren et al. 2013, 2017, 2019) have grouped this species with Schlechtendalia chinensis and suggested possibly synonymy, but the morphological differences indicate that this requires further investigation.

Nurudea shiraii (Matsumura)  Galls are inflorescence-like, densely pubescent, pinkish (see fig. 4a in C. Zhang et al. 2008), on leaflets of Rhus chinensis in China, Taiwan and Japan.  Alatae (BL 1.0-1.4 mm) with black head, blackish brown pterothorax and yellowish brown abdomen, emerge in autumn and fly to an unknown secondary host (Takahashi 1921). Ren et al. (2013) found genetic divergence between Chinese and Japanese populations assigned to this species.

Nurudea yanoniella (Matsumura)  (= Fushia rosea Matsumura)  Galls are rosy red (see fig. 4c in C. Zhang et al. 2008, as Nurudeopsis rosea), on leaflets of Rhus chinensis in China, Japan and Korea.  Alatae (BL 1.0-1.4 mm) emerge in autumn and fly to an unknown secondary host. Ren et al. (2013) found genetic divergence between Chinese and Japanese populations assigned to this species.

Nurudea zhengii Ren &Qiaio  Galls are rosy red and flower-like, formed on the leaves of the apical part of the stem of Rhus hypoleuca in China (Ren et al. 2018). Alatae (BL 1.2-1.5 mm) appear in late September and fly to an unknown secondary host.