The Aphids


(in alphabetical order)


















Idiopterus Davis

Aphidinae: Macrosiphini

A genus for one distinctive fern-feeding species.

Idiopterus nephrelepidis Davis  Plate 19g  (Fig.43i)   Apterae are black with pale legs, pale antennae ringed with black, siphunculi black basally and pale distally, and a black cauda (see; BL 1.2-1.6 mm. Alatae have broadly black-bordered forewing veins and a large white spot on the pterostigma. On many genera and species of ferns (Acrostichum, Adiantum, Asplenium, Anemia, Blechnum, Blotiella, Ceropteris, Dicranopteris, Dryopteris, Gymnocarpium, Nephrolepis, Pityrogramma, Polypodium, Polystichum, Pteridium, Pteris). Records from other plants are probably all vagrants. Perhaps neotropical in origin (Holman 1974), now almost cosmopolitan, but confined to glasshouses and caves in northern temperate regions. Apparently entirely anholocyclic. See Steffan (1962) and Heie (1994) for further information. 2n=13.

Illinoia Wilson

Aphidinae: Macrosiphini

About 45 North American species and one from the Caucasus.  Illinoia are pale spindle-shaped aphids with long appendages, most easily recognised by the long weakly to moderately clavate siphunculi, similar to those of Amphorophora  but ornamented on the distal, constricted region with a few rows of polygonal reticulation. There is no host alternation. Evolution of host relationships within the genus is somewhat obscure; many species in the two principal subgeneric groups are associated with Ericaceae, but others feed on taxonomically diverse plants, including some on trees. The species from cultivated Rhododendron mostly bear 5 hairs on the first tarsal segments.  The subgenus Masonaphis contains about 8 species, 6 of them on Ericaceae in North America.  Illinoia s.str. contains about 30 species, 9 of them (including the Russian one) associated with Ericaceae, with 6 of the species known from eastern North America.  Seven species of Illinoia s.str. and one Masonaphis live on Compositae.  The 3‑4 species of subgenus Oestlundia are associated with Rubus and the 2 subspecies of Amphorinophora live on Lonicera.  MacGillivray (1958) revised Illinoia (= Ericobium) using the name Masonaphis for the full genus.  Blackman (2010) reviewed and keyed the 5 species introduced to Europe from North America.

Illinoia alni (Mason)  Apterae are spindle-shaped, pale green with a dark green median stripe and dark spots at bases of siphunculi; BL 2.5-3.1 mm.  Immatures are pale green.  In small colonies on undersides of leaves of Alnus rugosa in north-eastern USA and eastern Canada (MacGillivray 1958).  Oviparae were collected in late August .  2n=10.

Illinoia andromedae (MacGillivray)    Apterae are shiny bright green with darker appendages, BL 1.9-2.3 mm. Specimens beaten from mixed vegetation of  Andromeda glaucophylla and Vaccinium myrtilloides fed on leaves of A. glaucophylla, which is thus the only confirmed host plant.  Only known from apterae collected in Quebec, Canada.

Illinoia azaleae (Mason)  (Fig.46a)   Apterae are rather shiny deep green; BL 1.9-2.7 mm. Immatures are pale greyish green. Alatae have quite dark wing-veins. On young stems and leaves of Rhododendron spp. (especially azaleas); also sometimes on other Ericaceae (Andromeda, Leucothoe, Pieris, Vaccinium) and occasionally on some other plants (Comptonia, Viola, Tulipa). In eastern North America, and introduced to Europe, South Africa, Hawaii, Australia, New Zealand and Argentina (Ortego et al. 2004). Probably mainly anholocyclic; oviparae sometimes occur, but males are undescribed. In colder climates it is an indoor or glasshouse pest of potted azaleas and can cause defoliation. A form described by Tissot & Pepper (1944) from Kalmia latifolia in Pennsylvania was regarded by MacGillivray (1958) as a subspecies, Illinoia azaleae ssp. kalmiaflora. It differs in the ratio of R IV+V to HT II from I. azaleae on Rhododendron (1.6-1.7 in kalmiaflora, 1.1-1.45 in azaleae), and is probably a good species. On the other hand, Amphorophora rhododendronia Mason, regarded as a subspecies of azaleae by MacGillivray, should probably be treated as a synonym. 2n=10.

Illinoia borealis (Mason)  Colour in life is unknown, probably dark green or brownish green with dark siphunculi; BL 1.7-2.5 mm.  On new growth of Ericaceae (Vaccinium, Gaultheria), and also recorded from Myrica aspleniifolia (= Comptonia peregrina) (MacGillivray 1958). In north-eastern USA and Canada (Quebec). Oviparae and apterous males are recorded from Quebec early in August ( Specimens in the BMNH collection from Clintonia identified as borealis seem to be pallida (q.v.).

Illinoia brevitarsis (Gillette & Palmer)    Apterae are pale yellowish green to yellow; BL 2.3-3.2 mm. On leaves of Clematis ligusticifolia in western USA. Monoecious holocyclic, alate males and oviparae occurring in late September-October (Palmer 1952). [In the BMNH collection there is an undescribed Macrosiphum sp. from the same host in Oregon (leg. D. Hille Ris Lambers, no. T85), distinguishable by the characters given in the key.]

Illinoia canadensis (MacGillivray)  Apterae are rather shiny, yellowish green mid-dorsally, darker green laterally with dark spots at the bases of the dark-tipped siphunculi; BL c. 2.7 mm.  On Myrica sp. in New Brunswick, Canada (MacGillivray 1958).  Life cycle is unknown.

Illinoia ceanothi (Bartholomew)    Apterae are green, BL 2.0-2.5 mm. On leaves of Ceanothus spp. in western USA (California, Oregon, Idaho, Washington; Alatae have 5-9 secondary rhinaria on basal part of ANT III (BMNH collection, leg. R.C. Dickson and H.L.G. Stroyan). Monoecious holocyclic, with oviparae and apterous males in late August to October (BMNH collection, leg. R.C. Dickson).

Illinoia corylina (Davidson)    Apterae are green with dark-tipped siphunculi: BL 2.3-2.5 mm. Apterae have 2-4 secondary rhinaria on ANT III, alatae have 23-28. Described from Corylus rostrata (= cornuta var. californica), and perhaps monoecious holocyclic on this host, as there are photographs by A. Jensen of a fundatrix and an aptera on C. cornuta at (and see also However Davidson (cited in Mason 1925) stated that it was more common on “ninebark” (Physocarpus capitatus). In western USA (California, Oregon, Washington). [In the BMNH collection (leg. D. Voegtlin) there are 2 apterae and 2 immatures that run to I. corylina in MacGillivray’s (1958) key, but collected from Aquilegia formosa. A. Jensen has collected an unidentified Illinoia from Aquilegia in Washington and British Columbia (, so perhaps there is another similar but undescribed species of Illinoia on this host.]

Illinoia (Amphorinophora) crystleae (Smith & Knowlton)    Apterae are pale whitish green, with pale appendages, dark only on tarsi and R IV+V; BL 3.7-5.0 mm. On leaves of Lonicera involucrata in western North America. Oviparae and alate males occur in August-September (MacGillivray 1958, Palmer 1952). A form described by Essig on L. ledebourii in California (as Amphorophora bartholomewi) is regarded as a subspecies, I. crystleae ssp. bartholomewi. 2n=16*.

Illinoia (Oestlundia) davidsoni (Mason)  Plate 25d   Apterae are pale green, slightly waxy, with brown-black siphunculi; BL 3.3-4.3 mm. On leaf petioles and undersides of leaves of Rubus parviflorus in western North America.  Amphorophora arnicae Glendenning is regarded as a synonym, with host alternation between Rubus and Arnica spp. suspected (MacGillivray 1958), but this has still not been confirmed experimentally. The collection data for the BMNH specimens suggests that there may be two similar species on Arnica, one migrating to Rubus and the other not.  2n=12.

Illinoia dzhibladzeae Shaposhnikov    Apterae are yellowish brown, shining brown dorsally, siphunculi paler with black apices; BL 1.9-2.5 mm. On undersides of leaves and on young shoots, flowers and fruits of Rhododendron flavum and Rhododendron sp. in Ukraine, Georgia and Turkey (BMNH collection, leg. N. Tuatay).

Illinoia finni (MacGillivray)    Apterae are undescribed, but some large greenish white apt. collected on Vaccinium sp. in Idaho (BMNH collection, leg. J.H. Martin) could be this species; BL of these apterae is 3.3-3.4 mm. BL of alate paratypes is 2.2-2.5 mm (2 specimens), collected on “huckleberry” (Gaylussacia or Vaccinium; we have included it under both these genera) in Ohio, USA. This species is similar to I. pepperi (MacGillivray 1958).

Illinoia goldmaryae (Knowlton)  (Fig.49a)   Apterae are bright green (Essig 1942, as Amphorophora patchiae); BL 1.8-2.9 mm. Forming dense colonies on young growth of species of Aster, Conyza, Erigeron and Solidago (MacGillivray 1958), and there are also records from other genera of composite plants.  Widespread in USA and eastern Canada, also in Central America (Panama; Quiros et al. 2009) and South America (Peru, Argentina; Ortego & Nieto Nafría 2017). Also introduced to England, where it was common in the London area on cultivated Michaelmas daisies and golden rods in 1960-62, and has since been recorded from suction traps (Blackman 2010).  It has also been reported from the Azores (Borges et al. 2010), and a trapped alata is recorded from Tunisia (Boukhris-Bouhachem et al. 2007). Monoecious holocyclic in USA, with fundatrices on Solidago in April and males in September (; the population introduced to England was apparently anholocyclic. 

Illinoia gracilicornis (MacGillivray)    Apterae are whitish to mid-green; BL 1.3-2.4 mm. On Holodiscus spp. in western USA and Canada (British Columbia). Monoecious holocyclic, with oviparae and apterous males in September. Jensen (2021) gives full descriptions of asll morphs. [The host recorded in the original description (only of oviparae) was Cercocarpus montanus, but Jensen (2021) has shown that this was almost certainly a misidentification of  Holodiscus.] 

Illinoia grindeliae (Williams)    Apterae are greyish green, siphunculi pale with dusky apices; BL 1.6-2.5 mm. Alatae may have 1-2 secondary rhinaria on ANT IV and V, as well as 17-21 on III. On Grindelia squarrosa, and possibly specific to this host, in western and central USA and Canada. MacGillivray (1958) differentiated some samples from Colorado and Utah as a subspecies (palmerae), giving the host as Grindelia sp., but paratypes that she donated to the BMNH collection are labelled as collected from G. squarrosa, and specimens subsequently collected do not provide evidence of any correlation between morphology and geographical distribution or host plant.

Illinoia lambersi (MacGillivray)  (Fig.46b)   Apterae and alatae exist as three distinct colour morphs, green, pink and yellow, which can often occur in mixed populations (see; BL 2.0-3.7 mm. Alatae have 21-30 secondary rhinaria on ANT III.  On young leaves, shoots and flowers of Rhododendron spp., including deciduous varieties. The principal host is Rhododendron, but there are also records from Daboecia cantabrica (e.g. Piron 2017), and colonies sometimes occur on Ilex aquilifolium (records from British Columbia, Canada, Forbes & Chan 1989, and England, BMNH collection, leg. V.F. Eastop).  In western North America, and introduced into Europe in about 1970 (Stroyan 1972b, Hille Ris Lambers 1973a), South America (Chile, Fuentes-Contreras et al. 1997), and Japan (Morimoto & Kiritani 1995).  Holocyclic on Rhododendron in California, USA, but European populations are anholocyclic (Hille Ris Lambers 1973a).  2n=10.

Illinoia liriodendri (Monell)  Apterae are spindle-shaped, pale green lightly dusted with wax, with antennae and siphunculi black except at bases, and legs pale green except for black tibial apices and tarsi (see; BL 1.7-2.5 mm.  A red colour form also occurs.  Monoecious holocyclic on Liriodendron tulipifera in North America, feeding on undersides of leaves. Introduced to Japan in 1991 (Sugimoto 1999), more recently to France (1998), Italy (2001), UK and Germany (2004), and Korea (Kim et al. 2011). It is also now recorded from Luxembourg, Slovenia, Poland, Hungary (Bozsik 2012), Serbia, Greece and Egypt. Oviparae and alate males occur in October.  Davis (1909) gave a full description, Dreistadt (1987) studied honeydew production, Zuparko & Dahlsten (1993) surveyed its parasitoids in California, and Dreistadt & Dahlsten (1988) discussed the timing of control measures.  2n=10.

Illinoia macgillivrayae (Hille Ris Lambers)    Apterae are pale yellowish green; BL 2.6-3.2 mm. Alatae have c.7-9 secondary rhinaria on ANT III (1 specimen, BMNH collection). In small colonies on young leaves of Raphiolepis indica, and subsequently found on other Rosaceae (Chaenomeles japonica, Pyrus kawakamii, Holodiscus discolor and Spiraea sp.), as well as two samples from Corylus cornuta (BMNH collection, leg. C.-k. Chan, and In western USA (California, Oregon) and Canada (British Columbia). It is very similar to I. spiraeae, but larger and with a relatively longer R IV+V.

Illinoia masoni (Knowlton)    Apterae are bluish green, with pale appendages; BL 1.8-2.8 mm. On leaves of Helianthus spp. in western USA (Palmer 1952). In laboratory tests, Rogers & Thompson (1978) found various degrees of resistance to this species in North American perennial species of Helianthus.

Illinoia (Oestlundia) maxima (Mason)    Apterae are white with black siphunculi (unpublished observations, R.L. Blackman); BL 2.9-3.9 mm. Alatae have a dark pterostigma and a dark spot at tip of forewing. On leaves of Rubus parviflorus in western North America (British Columbia to California). Monoecious holocyclic with an abbreviated life cycle, oviparae and alate males being produced in the 3rd or 4th generation from late May (California) to July (Frazer & Forbes 1968). Population dynamics of this aphid (Gilbert 1980) and its interactions with a parasitoid (Gilbert & Gutierrez 1973) have been studied intensively. 2n=12.

Illinoia (Masonaphis) menziesiae Robinson    Apterae are grey-green to salmon pink (see; BL 2.0-2.5 mm. Alatae are undescribed. On undersides of leaves of Menziesia ferruginea, where it causes a yellowing and downward curling of apical portions of leaves. In western USA (Idaho, Montana, Washington, A. Jensen, and Canada (Alberta, original description, and British Columbia, BMNH collection, leg. C.-k. Chan.) Monoecious holocyclic, with oviparae and apterous males collected in early September-October. (The siphunculi are unusual for this genus in having little or no distal swelling. The specimens described by Robinson are probably fundatrices.) 2n=10*.

Illinoia morrisoni (Swain)    Apterae are spindle-shaped, long-legged, rather dark apple green, with antennae dark beyond the base of ANT III, and legs and siphunculi tipped with black (see influential points/Gallery); BL 1.5-2.3 mm.  On terminal leaves and shoots of numerous conifers, usually Cupressaceae (Actinostrobus, Callitris, Calocedrus, Chamaecyparis, Cupressocyparis, Cupressus, Juniperus, Thuja, Widdringtonia); also on Araucariaceae (Araucaria) and Taxodiaceae (Metasequoia, Sequoia, Taxodium). There is only one record from Pinaceae (Cedrus deodora; BMNH collection, leg. H.G. Walker).  In western North America, Panama (Quirós et al. 2009), Costa Rica and Venezuela (Villalobos Muller et al. 2010), and introduced into Europe (UK, France, Portugal; Rabasse et al. 2005b, Wieczorek et al. 2019). Alatae have also been trapped in northern Italy (Coceano & Petrović-Obradović 2006). The life cycle is unknown; no sexual morphs are described and, in California at least, it seems to be anholocyclic.  2n=10.

Illinoia pallida (Mason)    Colour of apterae in life is unrecorded; BL 1.7-2.6 mm. Alatae are undescribed. On Clintonia borealis in north-eastern USA and Canada (Maine, New Brunswick). The aptera was redescribed by MacGillivray (1958).

Illinoia (Masonaphis) paqueti (MacGillivray)  (Fig.46c)   Apterae are whitish, yellowish or reddish, wax-dusted, with black-tipped siphunculi; BL 3.1-4.0 mm. On felty undersides of old senescing leaves of Rhododendron (=Ledum) groenlandicum in Canada (Quebec, Labrador) and Alaska (BMNH collection). Monoecious holocyclic, with oviparae and alate males in August (MacGillivray 1958).

Illinoia patriciae (Robinson)  Apterae are spindle-shaped, yellowish green to green; BL 1.7-2.3 mm.  On new needles of Tsuga heterophylla in British Columbia, Canada (Robinson 1969) and Oregon, USA (  Differences from the one other conifer-feeding Illinoia species, I. morrisoni (which has not been recorded from Tsuga) are given by Robinson.  The life cycle is not known.

Illinoia pepperi (MacGillivray)  (Fig.56c,d)  Apterae are green or red, with rather dark antennae, legs and siphunculi; BL 1.7-2.8 mm. On Vaccinium spp. in north-eastern North America. Monoecious holocyclic with oviparae and alate males in October (MacGillivray 1958). Elsner & Kriegel (1989) studied distribution and seasonal phenology in Michigan. 2n=28.

Illinoia phacelia (Essig)    Apterae are green with dark-tipped appendages; BL c. 2.5-2.7 mm. On undersides of leaves of Phacelia sp. in western USA (California, Oregon). The alata was redescribed by MacGillivray (1958).

Illinoia (Masonaphis) pinawae (Robinson)  (Fig.46e)  Apterae are pale yellowish green; BL 1.9-2.5 mm. Alatae are undescribed. On Rhododendron (=Ledum) groenlandicum in Canada (Manitoba, Ontario).

Illinoia (Masonaphis) rhododendri (Wilson)  (Fig.46b,d)  Apterae are green or pink, sometimes with dorsal cuticle sclerotic and rather smoky; BL 2.4-2.8 mm. On Rhododendron sp(p) in north-western USA. The life cycle is uncertain but may be abnormal, with fundatrices in late July, and possible early production of sexuales (MacGillivray 1958).

Illinoia (Masonaphis) rhokalaza (Tissot & Pepper)    Apterae are light yellowish brown, sometimes with greenish tinge, with dark brown to antennae, legs and siphunculi; BL 2.2-2.9 mm. On Rhododendron spp., Leucothoe fontanesiana and Kalmia latifolia in eastern USA (MacGillivray 1958).

Illinoia richardsi (MacGillivray)    Apterae are white with pale appendages, with only apices of antennal segments, apices of tibiae and tarsi darker to blackish; BL 2.6-3.2 mm.  Alatae are also very pale but have brownish head, thorax, antennae and legs, and rather dark wing veins. On undersides of leaves and in inflorescences of Anaphalis margaritacea and Gnaphalium bicolor (= Pseudognaphalium bioletti); often found in the inflorescences with Uroleucon russellae. Widely distributed in USA and Canada. Hille Ris Lambers (1966) described a form in California with thicker and less swollen siphunculi as a subspecies, I. richardsi ssp. pacifica.  Sexual morphs and life cycle unknown.  2n=10.

Illinoia (Oestlundia) rubicola (Oestlund)    Apterae are greenish yellow, often with a broad dark green spinal stripe (see influential points/Gallery); BL 3.1-4.6 mm. Alatae have a dark pterostigma and a large black spot at the tip of each forewing. On stems of Rubus idaeus var. strigosus, and occasionally on other closely-related Rubus spp., widely distributed in North America. Monoecious holocyclic, with oviparae and alate males in October. 2n=12.

Illinoia simpsoni (MacGillivray)    Colour of apterae in life is unknown; BL 1.8-2.1 mm. Alatae are undescribed. On Linnaea borealis in Maine, USA.

Illinoia spiraeae (MacGillivray)  Colour of apterae in life is unknown; BL 2.7-2.9 mm. Alatae have not been described. On Spiraea spp. in north-western USA (Oregon, Idaho) and British Columbia, Canada, and there are also samples in the BMNH collection from certain other woody Rosaceae, including Photinia davidiana, and also from plant genera in other families (Deutzia, Rhus, Salix, Yucca) indicating that this species has polyphagous tendencies. Very similar to I. macgillivrayae; the difficulties of distinguishing between these two species are discussed by Jensen (2021)  2n=10.

Illinoia spiraecola (Patch)    Apterae are various hues of red, green and yellow (see influential points/Gallery); BL 2.5-3.2 mm. On Spiraea spp., widely distributed in North America. Specimens identified as this species have also been collected from undersides of leaves of a tall Thalictrum sp. It was redescribed by MacGillivray (1958).

Illinoia subviride (MacDougall)    Apterae are delicate green, similar in colour to hairs of the host plant. On Aster alpinus in Washington, USA. 2n=10.

Illinoia thalictri (MacGillivray)    Apterae are very pale sea-green to yellowish green, lightly wax-dusted; BL 2.0-2.3 mm. On leaves and petioles of an unidentified Thalictrum sp. in Colorado, USA, and recorded from T. fendleri in New Mexico (Miller et al. 2016); a photograph of an Illinoia sp. on Thalictrum ( is probably this species. Monoecious holocyclic with oviparae and alate males in September (MacGillivray 1958).

Illinoia wahnaga (Hottes)    Apterae are pale whitish green to milky white, with black eyes; BL 2.2-2.6 mm. Living singly on undersides of leaves of  plants in genera formerly placed in Convallariaceae (Convallaria, Maianthemum) in North America (Colorado, British Columbia, Manitoba). Oviparae and apterous males occur in September-October in Manitoba (Robinson 1965). 2n=10.

Illinoia wilhelminae (Hille Ris Lambers)  Apterae are spindle-shaped, colour in life is not recorded; BL c. 3.1 mm.  The siphunculi show little or no trace of swelling.  Described from a single aptera collected in Quebec, Canada, on Alnus rugosa, but the form of the last rostral segment suggests that this was not its normal host plant (Hille Ris Lambers 1962).  It does not seem to have been recorded since.

Impatientinum Mordvilko

Aphidinae: Macrosiphini

A genus which possibly has host-alternation between Smilax and Impatiens in Asia, although this needs further confirmation; the two species that have spread to Europe produce their sexual morphs on Impatiens. Five species living on Cuphea in America (Mexico, Panama and Brazil) have also been placed in this genus. Reviews are available for Europe (Heie, 1994), Japan (Miyazaki, 1971), Korea (Lee & Havelka, 2001a) and America (Nieto Nafría et al. 2012).

Impatientinum americanum  Remaudière    Apterae have a shining black dorsum, antennae dark except for basal part of ANT III, and dark siphunculi; BL 1.5-2.6 mm. On Cuphea aequipetala and Cuphea sp. in Mexico. Alatae were described by Nieto Nafría et al. (2012). The record from Panama (Quiros et al. 2009) should be referred to I. chiriquense.

Impatientinum asiaticum Nevsky  Plate 25f (Fig 33a,c)   Apterae are bright green, pink or red, with an extensive shiny brown-black dorsal shield, black siphunculi and yellow to whitish cauda (see influential points/Gallery); BL 2.1-3.1 mm. On undersides of leaves along main veins, and later in dense colonies on flowerstalks, of Impatiens spp., especially I. parviflora. Probably originally in Central Asia (Tien Shan), now widely distributed in Europe (Holman 1971, Heie 1994) and Central Asia (Kadyrbekov 2017a). A subspecies, I. asiaticum ssp. dalhousiense, was described from Smilax parvifolia (= elegans) in north-west India (Verma 1969, as I. impatiensae dalhousiensis).  I. asiaticum s.str. is monoecious holocyclic on Impatiens in Europe (with alate males), but the north Indian subspecies may be heteroecious holocyclic between Smilax and Impatiens. 2n=16.

Impatientinum balsamines (Kaltenbach)    Apterae are green with an extensive shiny black dorsal shield, black siphunculi and a pale cauda; BL 2.0-2.7 mm. Singly or in small colonies on undersides of leaves of Impatiens spp. but especially associated with I. noli-tangere. Probably east Asian in origin, now widely distributed in Europe and Asia. Monoecious holocyclic with alate males. 2n=16.

Impatientinum chiriquense Nieto Nafría & Mier Durante   Apterae probably have a shiny black dorsum in life, and dark antennae and siphunculi; BL 1.7-2.0 mm. Apterae and alatae were collected in May on Cuphea infundibulum (= appendiculata) in Panama (Nieto Nafría et al. 2012). Other morphs are undescribed.

Impatientinum holmani Peña-Martínez & Nieto Nafría   Apterae probably have a shiny black or dark brown dorsum in life with dark siphunculi, but antennae mainly pale; BL 1.7-2.3 mm. Monoecious holocyclic on Cuphea sp. in Mexico, with alate males collected in October (Nieto Nafría et al. 2012).

Impatientinum impatiens (Shinji)  (Fig.33b,d)   Apterae are yellowish green to bluish green or orange-red with an extensive shiny black dorsal shield, black siphucunli and a pale cauda; BL 2.4-3.5 mm. On Impatiens spp. (especially I. textori) in Asia (north India, east Siberia, Japan, Korea, Taiwan, Laos). Fundatrices were described from Smilax china in Japan (Miyazaki 1971), and it has also been collected from this host in Laos (Yoshitomi et al. 2014). There is presumably a host alternation between Smilax (primary host) and Impatiens, but this needs to be confirmed experimentally. Macrosiphum smilaceti Takahashi is probably a synonym. 2n=16.

Impatientinum oaxacense Nieto Nafría & Pérez Hiidalgo   Apterae probably have a shining black dorsum in life, with dark siphunculi, the antennae and legs mainly pale; BL c.2.3 mm. Only known from the holotype aptera, collected in August on Cuphea sp. in Mexico (Nieto Nafría et al. 2012).

Impatientinum paranaense de Carvalho, Cardoso & Lazzari    Apterae are green with abdomen olive green, antennae, legs and siphunculi dark brown, and cauda green; BL 1.5-2.0 mm. Alatae have 15-24 secondary rhinaria distributed over the length of ANT III. On flower stems of Cuphea calophylla in the state of Paraná, Brazil (de Carvalho et al. 2004). The life cycle is unknown.

Impatientinum (Neoimpatientinum) smilaceti Agarwala, Mondal & Raychaudhuri    Colour of apterae in life is not recorded, presumably green or reddish with broad dark marginal longitudinal bands, black siphunculi and a pale cauda; BL 2.8-3.0 mm. On Smilax macrophylla in Sikkim, India. [This is a secondary homonym of Macrosiphum smilaceti Takahashi, if that species is I. impatiens.]

Inbaria Barjadze, Halbert, Matile & Ben-Shlomo

Eriosomatinae: Fordini

A genus erected for one gall-forming Middle-Eastern species originally described in Geoica, distinguished by the very large single rhinaria on each of ANT III-VI in the emigrant alatae (Barjadze et al. 2018c).

Inbaria swirskii (Remaudière & Inbar)   Galls on Pistacia atlantica are of irregular shape, bulbous, c. 25 mm in diameter, arising from leaf stem between bases of leaflets. Apterous fundatrigeniae and emigrant alatae were collected from the galls in October (Remaudière et al. 2004). Secondary hosts and other life cycle stages are unknown. Only known from Jordan.

Indiaphis A.N. Basu

Aphidinae: Macrosiphini

Four Indian species on Ericaceae, closely related to Ericolophium, and possibly not warranting a separate genus. A.K. Ghosh (1991) provided a key to species. Alatae have dark dorsal abdominal cross bands or a quadrate patch.

Indiaphis crassicornis A.N. Basu   Plate 23g  (Fig.46h)   Apterae are pale glassy white, dusted with wax; BL 1.7-2.0 mm. Alatae have secondary rhinaria distributed III 21-40, IV 0-8.  On Rhododendron spp. in India (Punjab, West Bengal). Alate males were collected in Punjab in November (BMNH collection, leg. Commonwealth Institute of Biological Control).

Indiaphis indica (L.K. Ghosh, Verma & Raychaudhuri)    Colour of apterae in life is unrecorded, probably pale with dark-tipped siphunculi; BL 1.4-2.4 mm. Alatae have secondary rhinaria distributed III 44-56, IV 1-7, V 0-2. On an unidentified Rhododendron sp. in India (Himachal Pradesh, Uttar Pradesh).

Indiaphis rostrata A.K. Ghosh & Raychaudhuri    Apterae are pale yellow; BL 1.8-2.0 mm. On undersides of leaves of an unidentified Rhododendron sp. in India (Sikkim). Only known from apterae; these were collected in March and may be fundatrices, which would explain the 5-segmented antennae with a short terminal process. (However they are probably not fundatrices of I. crassicornis, because of the long R IV+V.)

Indiaphis setosa (Hille Ris Lambers & A.N. Basu)    Apterae are green to dark green, with brown legs and dark brown siphunculi; BL 1.6-2.2 mm. Alatae have secondary rhinaria distributed III 8-12, IV 0. On Agapetes (= Pentapterygium) serpens, often in rather large colonies on tender parts of shoots and sometimes on undersides of leaves. In West Bengal, India.


Indiochaitophorus Verma

Calaphidinae: Panaphidini

One elm-feeding species related to Chromocallis, but with shorter antennal hairs, unbordered forewing veins and a heavily sclerotised apterous morph (see Quednau 2003)

Indiochaitophorus furcatus Verma   Apterae are rather elongate oval, pale brown with thick black dorsal longitudinal stripes; BL 1.3-2.0 mm. Alatae have paired dark spinal and marginal sclerites on each of abdominal tergites 1-6. Recorded from undersides of leaves and twigs of Ulmus wallichiana in India (Kashmir, Himachal Pradesh), and on U. villosa in Pakistan (BMNH collection, leg. M.A. Ghani). Heavy infestation of riverside trees was reported in Kashmir in June (Verma 1970a). Records from Algeria (Laamari et al. 2013) and Turkey (Şenol et al. 2017) are doubtful and require additional confirmation. The life cycle is unknown.

Indoidiopterus Chakrabarti, A.K. Ghosh & Raychaudhuri

Aphidinae: Macrosiphini

One east Asian species on Geranium. The species was transferred to Neotoxoptera by Naumann-Etienne & Remaudière (1995), but the unusual nature of the spiracular plates and the dorsal hairs, taken together with the wing venation of the alatae, seems to warrant the reinstatement of the genus.

Indoidiopterus geranii (Chowdhuri, R.C. Basu, Chakrabarti & Raychaudhuri)  (Fig.30e,f)   Colour of apterae in life is unrecorded; BL 1.6-1.8 mm. The alata was described by Chakrabarti et al. (1972).  On Geranium spp. in north-west India and Pakistan. Alate males were trapped in Pakistan (Naumann-Etienne & Remaudière 1995, as Neotoxoptera geranii). 2n=12.

Indomasonaphis Verma

Aphidinae: Macrosiphini

Three species in north India and Pakistan, at least two of which have host alternation between Rhododendron and Compositae/Asteraceae. They have characters (e.g. spinulose hind tarsi, alatae with numerous secondary rhinaria) shared with other members of a group of Asian genera associated with Rhododendron (Chaetomyzus, Ericolophium, Indiaphis).

Indomasonaphis anaphalidis (A.N. Basu)  (Fig.34a, Fig.46g)    Apterae are elongate oval, rather shiny pale green to lemon yellow, BL 3.4-4.8 mm. Alatae have variably developed black dorsal abdominal patch. Males are alate. Originally described from Anaphalis triplinervis in West Bengal, feeding on tender shoots and leaves without causing noticeable damage, and since recorded from various other Compositae/Asteraceae (Inula, Ageratum, Gerbera, Senecio, Artemisia), as well as some unlikely hosts (Eurya japonica, Morus alba, Oxalis corniculata). Heteroecious holocyclic with a sexual phase on Rhododendron spp. in northern India (Verma 1971, as Indomasonaphis indica; Chakrabarti et al. (1983) as Indumasonaphis tuberculatus) and Pakistan (Naumann-Etienne & Remaudière 1995).

Indomasonaphis inulae (A.K.Ghosh & Raychaudhuri)    Apterae are elongate-bodied, with colour in life unrecorded, probably pale; BL 2.8-3.4 mm.  On Rhododendron sp. and Inula cappa in northern India. Host alternation between these two plants (with Rhododendron as the primary host) is believed to occur (Chakrabarti et al. 1983, Chakrabarti & Banerjee 1993a). 2n=32 (Kurl & Chauhan 1988).

Indomasonaphis rumicis (Chakrabarti & Raychaudhuri)    Apterae are green to brown; BL c.3.8-3.9 mm. Alatae have dark dorsal abdominal cross-bands. On undersides of leaves of an unidentified Rumex sp., and (as 2 alate males) on Oxyria digyna, in northern India (Uttar Pradesh). Also collected (as immature alatae) on Gerbera (assuming that “Garbera” was a typographic error), which is a more probable host for a species in this genus. Alate males were found in October (original description).

Indomegoura Hille Ris Lambers

Aphidinae: Macrosiphini

Four species in east Asia with dark, somewhat swollen siphunculi reticulated at apices. Two species have Staphyleaceae as primary hosts. Of the other two species one was described from an  unidentified plant, and the other is only known from alatae collected on a shrub that was subsequently identified as Corchorus sp. and unlikely to be the true host.  G. Zhang & Qiao (1998b) reviewed the genus in China.

Indomegoura indica (van der Goot)   Plate 20i   Apterae are orange yellow, covered in white wax, with stout, dark siphunculi; BL  3.1-4.2 mm. On Staphylea bumalda and Euscaphis japonica in Japan, and recorded as common in the Tokyo area, feeding on the undersides of the branches, which may be severely affected (Takahashi 1923, Miyazaki 1971). Populations in summer occur on Hemerocallis spp., but there is some uncertainty about the life cycle as Takahashi (1923), in describing the biology, recorded apterous sexuparae. and both apterous and alate males on the primary host, indicating that only part of the population may migrate.  In Japan, Korea, China, Taiwan and India. 2n=10.

Indomegoura nigrotibiae (Tao)    Colour of apterae in life is unknown; BL 2.2-2.5 mm. On Staphylea bumalda (Miyazaki 1971). In Taiwan, Japan and Korea (Lee et al. 2002). Alatae are undescribed, and the life cycle is unknown.

Indomyzus Ghosh, Ghosh & Raychaudhuri

Aphidinae: Macrosiphini

Myzus-like aphids in east Asia with long dorsal hairs like Micromyzodium but having apterae with secondary rhinaria on ANT III and also sometimes on IV. Alatae are undescribed. The type species I. sensoriatus is recorded from both India (A.K. Ghosh et al. 1971f) and China (Su et al. 2011) but in both cases the host plant was unidentified.

Indomyzus glaricae Sathe & Jadhav  NOMEN NUDUM  Apterae are dark brown, shiny; BL c. 1.5 mm. Differences from the type species of the genus include fewer secondary rhinaria, siphunculi with a much smaller flange, and more hairs on the cauda. On Gliricidia maculata in India (Sathe & Jadhav 2008). Other morphs and life cycle are unknown. Unfortunately the name and description are invalid as no type specimens were designated.

Indonipponaphis Ghosh & Raychaudhuri

Hormaphidinae: Nipponaphidini

Two little-known species described from Quercus, related to Nipponaphis but without spinal hairs on abdominal tergite 7, and with paired spinal and marginal tubercles on prosoma and abdominal plate.

Indonipponaphis fulvicola Sorin   Galls on Distylium racemosum in Japan are almost globular, up to 27 mm, in cusps of young leaves, and appear to be very rare (Fukatsu et al. (2000). Alatae migrate in late October-November to Quercus acuta and Q. gilva. Apterae on Quercus are brown, almost circular, dorso-ventrally flattened, strongly sclerotised; BL c. 2.1 mm (Sorin 1979b).  Sorin (1997) gave an account of the life cycle and described all the morphs on Distylium including sexuparae and sexuales. Fukatsu et al. (2000) described attacking behaviour by unmodified but long-lived 1st instars of the gall generation. Only known from Japan; records from India in Holman (2009) appear to be in error.

Indonipponaphis tuberculata Ghosh & Raychaudhuri   Apterae are brownish, oval, strongly sclerotised; BL 1.9-2.1 mm.  On stem of Quercus dealbata (= Q. floribunda?) in Meghalaya, India (A.K. Ghosh & Raychaudhuri 1973).  Biology and other morphs are unknown.

Iowana Hottes

Aphidinae: Aphidini

One nearctic species with very short and sparse hairs on body and appendages, a short rounded cauda, short flangeless siphunculi, well-developed marginal tubercles and a little-differentiated anal plate. Until recently it was known from only two apterae collected in Iowa, USA in 1925, but Favret et al. (2004) rediscovered it in Illinois and described all morphs. A second species described in this genus from China is also included here, although more likely to belong to another genus, possibly Aphis (Protaphis).

Iowana frisoni Hottes    Apterae are pale green with  light reddish brown head and prothorax and brown siphunculi; BL 1.8-3.1 mm. Alatae have the media usually once-branched and 12-22 secondary rhinaria on ANT III. In ant shelters at bases of flowering stems and in leaf axils of Silphium laciniatum and S. terebinthinaceum (no colonies were found on two other Silphium spp.). The populations studied in Illinois by Favret et al. (2004) were most commonly attended by Lasius neoniger. Monoecious holocyclic, with oviparae and apterous males produced in October.

Iowana zhangi L. Zhang    Colour of apterae in life is unrecorded; BL c.1.6 mm. On Ophiuros exaltatus in Heilongjiang province, China (L. Zhang 2000). Other morphs and life cycle are unknown.

Ipuka van Harten & Ilharco

Aphidinae: Macrosiphini

Two species on Compositae/Asteraceae in Africa and east Asia , possibly most closely related to Aulacorthum, but antennae of apterae as well as alatae bear numerous secondary rhinaria on III-V, and immatures have spinulose hind tibiae (van Harten & Ilharco 1976). The host of a third, African, species is unknown.

Ipuka dispersa (van der Goot)   Plate 22e   Apterae are brownish grey with ill-defined dark transverse bars on thorax and abdomen, darker brown patches around bases of siphunculi, dark red eyes, antennae dark except at base of III, legs yellowish except for black knees and tarsi, siphunculi black and cauda pale yellowish (see aphids of Karnataka website); BL 1.6-2.2 mm. On Emilia spp., often living concealed in leaf sheaths. In Africa, Sri Lanka, China (Su et al. 2011), south-east Asia and Australia. Apparently anholocyclic.

Ipuka melantherae van Harten & Ilharco     Apterae are greenish yellow to yellowish orange, with an orange spot around siphunculi, which are blackish, and a pale yellowish cauda; BL 1.8-2.2 mm. On undersides of leaves of Melanthera sp. in Angola.

Iranaphias Remaudière & Davatchi

Aphidinae: Macrosiphini

One species only known from Iran, apparently related to Hydaphias but with first tarsal chaetotaxy 3,3,3, much shorter siphunculi and cauda, and apterae with numerous, long  pointed dorsal hairs (Remaudière & Davatchi 1959).

Iranaphias dehbani (Remaudière & Davatchi)    Apterae are very broadly oval, dark-coloured with slight pruinosity; BL 1.0-1.4 mm. Alatae have secondary rhinaria distributed III 16-24, IV 0-3.  On Galium sp. in Iran (Izadbar), at 2,600 m altitude. Life cycle is unknown.

Israelaphis Essig


A small, distinctive and probably rather ancient genus of grass-feeding aphids found only in the Mediterranean region and western Iberia. The life cycle is very unusual, and related to the Mediterranean climate.  The parthenogenetic phase exploits short-lived annual grasses with a winter growth period following autumnal rains, sexuales being produced and diapausing eggs laid in spring.  Alatae are apparently very rare and only known for one species, so dispersal is very limited. Barbagallo & Patti (1999) reviewed the genus and discussed its possible affinities. Nieto Nafría & Mier Durante (1998) reviewed the Iberian species.

Israelaphis alistana Mier Durante    Apterae are dark green, with rather darker appendages than I. carmini; BL 2.1-2.5 mm. On grasses (Anthoxanthum, Vulpia) in Spain (Zamora). Monoecious holocyclic with oviparae and apterous males in May-June (Mier Durante 1978, as I. tavaresi ssp. alistana). Oviparae are dark green (cf. carmini). Slide-mounted specimens are not reliably separable from I. carmini, and in view of the low dispersal ability of these aphids (no alatae are known), it seems likely that the original subspecies designation of alistana was correct, and that it is a local isolated variant population of carmini. 2n=18.

Israelaphis carmini  Essig    Apterae are greenish yellow, with slight wax covering, and pale appendages; BL 1.9-3.0 mm. On grasses (Avena, Bromus, Hordeum, Koeleria) in hilly regions of Israel, Sicily and Portugal. Monoecious holocyclic, with oviparae and apterous males in March-May (Ilharco 1965, as I. tavaresi; Barbagallo & Patti 1999). Oviparae are brown (cf. alistana). 2n=18.

Israelaphis ilharcoi Barbagallo & Patti    Apterae are grass-green or yellowish green, with slight whitish wax secretion, head brownish, appendages dark-tipped; BL 2.3-3.0 mm.  Alatae are unknown. On grasses, especially Bromus subg. Anisantha, in Sicily and Portugal. Monoecious holocyclic, with eggs hatching in October, apterous viviparae developing during the winter months and giving rise to oviparae and apterous males in spring (Barbagallo & Patti 1999).

Israelaphis lambersi Ilharco  Plate 5a   Apterae are greenish yellow to yellow; BL 2.0-2.9 mm. On upper surfaces of leaves of grasses (Avena, Bromus, Phalaris, Phleum) in Portugal and southern Spain. Alatae are very rare and have abnormal wing venation (Ilharco 1966b). Monoecious holocyclic; apterous males are produced in early spring (original description). 2n=16.

Iziphya Nevsky

Calaphidinae: Saltusaphidini

About 12 species of squat-bodied, distinctively marked sedge-feeding aphids, leaping from host when disturbed. Alatae have banded wing veins. Six species are nearctic, five palaearctic and one holarctic. The taxonomy of Iziphya is difficult due to seasonal polymorphism and environmental variation in important diagnostic characters (body pigmentation, antennal length, shape of dorsal hairs. Quednau (2010) reviewed the genus, keyed the apterae and alatae of all species and illustrated all available morphs. He sank several names in synonymy and also concluded that some species were less host specific than originally believed.

Iziphya albipes  Richards    Apterae have dorsum mostly black with a pale area on anterior abdomen, a yellowish median area on head, and yellowish tibiae and tarsi; BL c.1.5-1.8 mm. In eastern North America; recorded from Carex pennsylvanica in Minnesota, USA (Quednau 2010), and on Carex sp. growing on rocky hillsides in Quebec, Canada (original description).

Iziphya americana (Baker)  (Fig.19d)  Apterae are yellowish or greyish, usually with a pair of large dark dorsal abdominal patches anterior to siphunculi, not fused medially; BL c.2.1-2.4 mm.  On Carex spp. (lepidocarpa, tribuloides) in bogs and wet meadows of northern USA and Canada.  Oviparae have been swept from C. tribuloides in October in Oregon (BMNH collection, leg. D. Voegtlin).

Iziphya bufo (Walker)   Plate 4d   Apterae are yellowish or greenish yellow with variable blackish grey markings, consisting usually of a pair of large dark areas on the metathorax and a dark arc joining siphunculi, but sometimes with dorsum almost completely black; BL 1.4-2.1 mm. Alatae have a dark central dorsal abdominal patch and broadly banded wing veins. On various Carex spp., especially C. arenaria, and also on Cyperus rotundus (BMNH collection, leg. R. van den Bosch), with odd records from Juncus (possibly misidentified Juncobia leegei) and grasses such as Cynodon dactylon (as I. maculata); often found in drier situations such as sandhills, drained forest soil and steppe regions, but also sometimes in bogs. Oviparae and apterous males occur in September-October. I. bufo is widespread in Europe, and in south-west and central Asia. This is a very variable species, particularly with regard to the shape of the abdominal and tibial hairs, and its morphological discrimination from I. memorialis is not at all clear.  We here follow Quednau (2010) in regarding I. maculata Nevsky, I. austriaca Börner and I. oettingenii Quednau as synonyms of I. bufo.

Iziphya eurymelaina Quednau     Colour in life not observed, but dorsum with extensive dark pigmentation; BL 1.2-1.7 mm. Vacuum-trapped, possibly from Carex, in Illinois, USA (Quednau 2010).

Iziphya flabella (Sanborn)    Apterae are yellowish green with variable dark dorsal markings usually consisting of paired dark areas on head, metathorax and abdomen, leaving a clear spinal stripe, but early and late season individuals may be much more extensively pigmented (see Quednau 2010); BL 1.5-2.0 mm. On leaves of  various Carex spp., especially on rocky hillsides or in dry forest soil. Widely distributed in North America. Oviparae and apterous males occur in September to early October. Quednau (2010) reported one incidental record from Germany, on C. pilulifera.

Iziphya ingegardae Hille Ris Lambers  (Fig.19c)   Apterae have the dorsum strongly arched and are yellowish with dark dorsal markings, including an irregular dark band forming an arc between the siphunculi, fused at least partially in midline; BL 1.8-2.2 mm. On Carex canescens and C. leporina in bogs in northern Europe (Sweden, Germany, Poland). Oviparae and apterous males occur in Sweden in early September (Ossiannilsson 1959).

Iziphya mackaueri Quednau    Apterae pale ochreous dorsally except for intersegmental muscle plates and dark spots at hair bases (Quednau 2010); BL c.2.3 mm. On Carex sp. in a wet bog in Ontario, Canada.

Iziphya memorialis Börner    Apterae are probably yellowish with dark dorsal markings; BL 1.6-1.9 mm. On Carex spp. (praecox, stellulata, possibly montana) in both wet and dry habitats in Europe (Czech Republic, Austria, Germany, Poland and Sweden).  Sexuales appeared in Sweden in early September (Ossiannilsson 1959).

Iziphya mordvilkoi Quednau & Shaposhnikov    Apterae are yellowish green with dark dorsal markings; BL c. 2.1 mm.  On Carex sp. in east Siberia.

Iziphya spenceri Richards    Colour of aptera in life was not observed, probably yellowish with dark dorsal markings; BL c. 1.7-1.8 mm. Described from an unknown host in British Columbia, Canada, and also recorded from Carex sp(p). in western USA from Washington to New Mexico (, but it seems to have a holarctic distribution, occurring on Carex sp. in dry steppe country in Mongolia (Quednau 1969; as mongolica), in Kazakhstan (Kadyrbekov 2011b), and also found on C. liparicarpos in Hungary (Szelegiewicz 1981a). Quednau (2010) illustrated an ovipara collected in Oregon in late September.

Iziphya variabilis Quednau    Colour of apterae in life was not observed, probably yellowish, with variable dark markings; BL c.1.7-1.8 mm. On Carex sp. in wet sites in mountainous steppe country in Mongolia and east Siberia (east Baikal region). Oviparae occur in October (Quednau 2010).

Iziphya vittata Richards    Apterae are rather elongate-bodied, yellowish or greyish with variable black dorsal markings, often consisting mainly of spots at hair bases, with antennae black except for basal part of segment III; BL 1.8-2.3 mm. On leaves of Carex spp. in wet places, open soil near lake shores, sometimes on road sides, widely distributed in North America. Oviparae occur in late July-August in Quebec (Hille Ris Lambers 1960, as I. punctata; Quednau 2010).