The Aphids

Four African species with a spinulose head, no antennal tubercles, usually 5-segmented antennae, Rhopalosiphum-like siphunculi and a short rounded cauda; an unusual combination which means that their systematic position is uncertain. The two species of which apterae are known feed on grass roots, and the morphological convergence with Rhopalosiphum is probably due to this rather than to any taxonomic affinity. Ilharco (1972b) provided keys to the alate morph.

Hallaphis ilharcoi van Harten Apterae are dark olive-green (immatures are brownish yellow); BL 1.3-1.8 mm. Alatae have 3-7 secondary rhinaria on ANT III. On roots of grasses (Tricholaena rosea?) a few cm below the surface, attended by ants (Pheidole sp.). Only known from Angola (Ilharco 1972b).

Hallaphis rhodesiensis (Hall) Apterae are green; BL 1.7-2.1 mm. On roots of unidentified Poaceae in Africa south of Sahara (see Millar 1994).

Hamamelistes Shimer

Hormaphidinae: Hormaphidini

A genus of five species with similar host plant associations to Hormaphis, and with a similar disjunct distribution in North America and east Asia, but with a gall (or “pseudogall”) dwelling stage on the secondary host Betula as well as on the primary host Hamamelis. The life cycle takes two years. Alatae have two oblique veins in the hind wing. Aoki et al. (2001) revised the Japanese species and clarified their life cycle and taxonomy with the aid of DNA sequence data. Phylogeny and evolution of the genus based on nuclear and mitochondrial DNA sequences were studied by von Dohlen et al. (2002) and Chen et al. (2011b). First instars have been shown to have a role in defending the inhabitants of the galls on Hamamelis (Shibao & Fukatsu 2003, Shibao et al. 2010). Qiao et al. (2018) gave an account of the Chinese species.

Hamamelistes betulinus Horvath Plate 3c, d Apterae in blister-like pseudogalls on birch are greenish or dark brown to black, BL 1.4-2.0 mm, secreting white wax (see influentialpoints.com/Gallery). They have short, 3- or 4-segmented antennae, and lack siphuncular pores in European populations, although siphunculi are present in some generations in Japan (Aoki et al. 2001). Alatae (BL 1.3-2.0 mm) have 5-segmented antennae and pigmented siphuncular pores. H. betulinus is anholocyclic on Betula spp., mainly on B. pendula, in Europe and northern Asia, east to Mongolia. It feeds on the undersides of the leaves causing pale yellowish blisters to develop on the upper surfaces (fig. 127), and overwinters as aleyrodiform/ coccidiform first instar larvae on the twigs. For a more detailed account of the anholocyclic European population see Heie (1980). In Japan, the secondary host is most commonly B. platyphylla and there is host alternation, sexuparae being produced in May-June and flying to Hamamelis japonica, where sexual morphs develop on the leaves and eggs are laid on twigs and trunks (Aoki et al. 2001). These hatch the following year and the developing fundatrices induce coral-like galls to develop from flower buds. Populations remaining on Betula produce secondary migrants that recolonise other birch trees and continue parthenogenetic reproduction. 2n=12 (for anholocyclic European population).

Hamamelistes kagamii (Monzen) Galls formed from flower buds or lateral buds of Hamamelis japonica in Japan are large, stalked, pouch-like, with 7-13 horn-like protruberances, yellowish and covered with brownish hairs (Monzen 1954). All the second generation in the galls develop as alatae and migrate to Betula grossa, where colonies of brown apterae on the undersides of the leaves cause blistering and yellowing. Sexuparae are produced in May-June, and there is presumably a two-year life cycle, with no evidence of anholocycly (Aoki et al. 2001). Shibao et al. (2010) studied defensive behaviour of the gall generations on Hamamelis.

Hamamelistes miyabei (Matsumura) This aphid produces rather large yellowish green spiny galls from the lateral leaf buds of Hamamelis japonica in Japan, similar to those formed from flower buds by H. spinosus in North America. The galls develop in June-July, and all aphids of the second generation in the galls are alatae, leaving the galls in late July-September. DNA sequencing has shown that the cockscomb-like galls on the upper surfaces of the leaves of Betula maximowicziana, containing reddish yellow apterae previously described as H. cristafoliae, are the secondary host populations of this species (Aoki et al. 2001). Alatae produced in the birch galls in June of the following year are sexuparae that migrate back to Hamamelis, but in August the alatae produced are secondary migrants which presumably recolonise B. maximowicziana and continue parthenogenetic reproduction, as in H. betulinus. First instar nymphs have a defensive role against predators, both in the cockscomb galls on birch (Akimoto et al. 1996, as H. cristafoliae), and in the spiny galls on Hamamelis (Shibao & Fukatsu 2003). [Bhattacharya et al. (1983) found a similar aphid causing folded-leaf pseudogalls on Betula alnoides in northern India, but they described an apterous ovipara from these galls, perhaps indicating a distinct species.]

Hamamelistes similibetulae (Qiao & Zhang) Apterae are brown, flattened, broadly oval; BL 1.0-1.2 mm. They feed singly on the undersides of mostly older leaves of Betula albosinensis, causing conical rounded protrusions on the upper sides of the leaves (Qiao & Zhang 2004c, as Hormaphis; the revised generic placement was confirmed using molecular methods by Chen et al. 2011b). In Tibet. Other morphs and life cycle are unknown.

Hamamelistes spinosus Shimer Apterous fundatrices in the spiny galls on Hamamelis virginiana (fig. 128A) are about 2 mm long, almost globular, dark purplish brown, covered with wax meal. Apterae feeding on the undersides of birch leaves are smaller, dark brownish red to purple with abdominal tufts of white wax. There is also a black, rugose, coccid-like morph that overwinters on birch twigs. The alatae may be distinguished from those of Hormaphis hamamelidis migrating between the same hosts by their 5-segmented antennae and two oblique veins in the hind wing. This species occurs throughout North America. The life cycle is complex, and extends over two years, as determined by Pergande (1901). Large numbers of alatae emerge from the galls produced by fundatrices in spring on H. virginiana, and fly to Betula spp. (papyrifera, lutea, nigra), where an overwintering coccidiform morph lives on the twigs until the following spring. The progeny of this morph move to the undersides of young leaves, producing reddish brown to crimson-coloured swellings of the leaf lamina between the veins. The apterae in these pseudogalls are dark brownish red to purple with abdominal tufts of white wax (see influentialpoints.com/Gallery). The return migrants to witch hazel produce sexuales which lay overwintering eggs. 2n= c.50*.

Hannabura Matsumura

Calaphidinae: Calaphidini

A genus for two species, one east Asian and the other North American, related to Calaphis but with fewer hairs on the first tarsal segment.

Hannabura alnicola Matsumura Apterae are pale yellowish, with mainly whitish appendages (Matsumura 1917); BL 1.5-1.6 mm. On Alnus spp. (hirsuta var. sibirica, japonica, matusumurae) in Japan. Oviparae were collected in July-August in Hokkaido (Higuchi 1972); biology is otherwise unknown.

Hannabura alnosa (Pepper) Apterae are creamy-white with a reddish-brown/orange line along each side of the body, and the very long antennae are dark (see influentialpoints.com/Gallery); BL 1.3-1.6 mm. They position themselves close to the larger veins on the undersides of leaves of the host (Alnus rugosa), and are very inconspicuous. Alatae are very pale green with dark antennae and legs. Sexuales occur in October; males are alate (Pepper 1950, as Calaphis alnosa). In north-eastern North America.

Hayhurstia del Guercio

Aphidinae: Macrosiphini

One species on Amaranthaceae with very small, slightly clavate siphunculi with a small flange, distinctly shorter than the finger-like cauda.

Hayhurstia atriplicis (L.) Plate 14b Apterae are green, covered with white wax powder (see influentialpoints.com/Gallery); BL 1.5-2.9 mm. On Amaranthaceae, usually Atriplex and Chenopodium spp, living inside pod-like yellowish pseudogalls formed by rolling leaves upwards. Widespread in Europe and Asia, and also in north and central Africa, and in North and central America. Mamontova-Solukha (1963) described a population on Chenopodium sp. in Ukraine as a subspecies, H. atriplicis ssp. chenopodii, and Kadyrbekov (2017a) reported that this form also occurs on two Chenopodium spp. in Kazakhstan. Monoecious holocyclic in coller climates, with oviparae in October (pictured at aphidtrek.org); males apterous or alate (Heie 1992, Blackman 2010). 2n=14.

Helosiphon Leclant

Aphidinae: Macrosiphini

One European species, possibly related to Anuraphis, with unusual features of the siphunculi and anal plate.

Helosiphon eryngii Leclant Plate 9g Apterae are yellow to grey-green orange with dark appendages, sometimes with a reddish area anterior to the siphunculi; BL 1.4-2.1 mm. Living on Eryngium campestre, at bases of petioles and at stem bases, attended by ants. In the south of France and Spain, and aslo reported from Greece (Ttsitsipis et al. 2007). Monoecious holocyclic, with apterous males (original description).

Hemipodaphis David, Narayanan & Rajasingh

Eriosomatinae: Eriosomatini

Two species, one known only from galls on Zelkova in Japan, and the other known only from a monocotyledenous secondary host in India. The taxonomic position of the genus was reviewed by Akimoto (1983).

Hemipodaphis persimilis Akimoto Spring generations form leaf-roll galls on Zelkova serrata in Japan; see http://www.agr.hokudai.ac.jp/systent/akimoto/gall/Hemipodaphis.jpg. Emigrant alatae are very small (BL 0.6-0.7 mm), brown, emerging in July-August to migrate to an unknown secondary host (Akimoto 1983). Populations persist in the galls until October (Akimoto 1992). Aggressive behaviour by first instar larvae was described by Aoki (1978, as Colopha sp. nr. caucasica). [This species was transferred to Zelkovaphis by Barbagallo (2002), but reinstated in Hemipodaphis by Sano & Akimoto (2011).] 2n=36 (Blackman 1986).

Hillerislambersia A.N. Basu

Aphidinae: Macrosiphini

One species in north India with very long fine-pointed dorsal body hairs, thick siphunculi with distal reticulation, and a short cauda.

Hillerislambersia darjeelingi A.N. Basu Plate 28a Apterae are pale green, with a longitudinal dark green stripe on each side of thorax and anterior abdomen; BL 2.3-2.9 mm. Described from large colonies on undersides of leaves of a Lonicera sp. in West Bengal, causing inward rolling and hardening of leaves. It has since recorded from Holboellia latifolia (possibly the true host?) and Rhododendron sp (Raychaudhuri, 1980). Life cycle is unknown; apparently anholocyclic in type locality.

Himalayaphis A.K. Ghosh & Verma

Aphidinae: Macrosiphini

One very large species in northern India with markedly clavate siphunculi, long antennae and long dorsal hairs with knobbed apices and tuberculate bases. The nearest relatives would appear to be Rhododendron-feeding genera such as Chaetomyzus and Neoacyrthosiphon.

Himalayaphis anemones A.K.Ghosh and Verma Apterae are greenish, BL 4.2-4.7 mm, collected from undersides of leaves of Anemone sp. in Himachal Pradesh, India. Alatae are still undescribed and the life cycle is unknown.

Holmania Szelegiewicz

Aphidinae: Macrosiphini

One species in Mongolia with long thick capitate dorsal hairs and 5 hairs on first tarsal segments like Chaetosiphon, and a similar host association with Potentilla, but with siphunculi having both hairs and polygonal reticulation. R IV+V is relatively long and hairy.

Holmania chaetosiphon Szelegiewicz Plate 16d Apterae are probably whitish in life with dark joints to antennal segments, dark antennal terminal process, dark femoral and tibial apices, tarsi and R IV+V; BL 1.7-2.0 mm (based on 4 apterae in BMNH collection, leg. H. Szelegiewicz). Described from one vagrant alata on Stipa, but Potentilla sibirica (= pennsylvanica) was subsequently recorded as the host (Holman & Szelegiewicz 1972), and the aptera was later described, from specimens collected on Potentilla longifolia (Szelegiewicz & Holman 1982).

Hoplocallis Pintera

Calaphidinae: Myzocallidini

Four palaearctic species on Quercus related to Myzocallis, but with antennal terminal process always shorter than ANT VI base, with small pronotal hairs in clusters, and with distinctive dorsal pigmentation and sclerotisation. All viviparae are alate. Immatures are much broader and flatter than adult alatae and have four longitudinal rows of brown sclerites bearing clusters of hairs. Quednau (1999) reviewed, keyed and illustrated the species.

Hoplocallis microsetosus Quednau & Chakrabarti Colour in life is not recorded; BL of alata c. 2.4 mm. On Quercus incana in Uttar Pradesh, India (Quednau & Chakrabarti 1976), on Q. incana and Q. leucotrichophora in Pakistan (Naumann-Etienne & Remaudière 1955, Quednau 1999)) and also on Q. baloot (= Q. ilex) in Afghanistan (BMNH collection, leg. DHRL). Sexuales are unknown.

Hoplocallis microsiphon Quednau & Barbagallo Colour in life is not recorded; BL of alata 1.6-2.1 mm. On various Quercus spp., but perhaps usually on Q. calliprinos, in the Mediterranean region (Israel, Lebanon, Italy, Turkey, Greece), and Czech Republic (Quednau & Barbagallo 1991). Oviparae are reported from Italy (Patti & Barbagallo 1997), and an alate male was collected in Greece in December.

Hoplocallis picta (Ferrari) Alatae are rather elongate, pale yellow-green with banded antennae and a series of dark spinal markings (see influential points/gallery); BL 1.8-2.1 mm. On undersides of leaves of Quercus spp. throughout the Mediterranean area, in south-west Asia, India (A.K. Ghosh & Quednau 1990), and introduced into South America (Argentina, Chile). In Europe it seems to be extending its distribution northward; there are trap records from England, and it has recently been found on Q. ilex in South Wales (Baker 2009b). In Israel males and oviparae appear in December (Swirski 1953, as Myzocallis bodenheimeri). 2n=14.

Hoplocallis ruperti Pintera Alatae rather elongate, yellow to whitish with dark lateral as well as spinal markings; BL c. 2.0 mm. Immatures are yellow with contrasting dark brown quadrate dorsal patches. On leaves of Quercus spp. (cerris, robur) in central, southern and eastern Europe; Austria, Italy (Sicily), Bulgaria, Czech Republic, Hungary, Turkey. Oviparae and males are produced in October in the Czech Republic (Pintera 1952).

Hoplochaetaphis Aizenberg

Calaphidinae: Myzocallidini

One small oak-feeding species in southern Europe and south-west Asia. Colonies include apterous viviparae with very long dorsal hairs; they appear to be adapted to living in leaves rolled by caterpillars.

Hoplochaetaphis zachvatkini (Aizenberg & Moravskaja) (= H. parvula) Apterae are broadly pear-shaped, variably coloured pinkish red, rich red, orange or dull brown, with a blackish brown dorsal sclerotic pattern that is particularly extensive in spring, sometimes dusted with a thin mealy wax so that they resemble caterpillar faeces (Aizenberg & Moravkaja 1959); BL 0.8-1.3 mm. Alatae have dark head and thorax and paired dark patches or cross-bands on abdominal tergites 3-7 or 4-7, broadest and often merged into a patch on abdominal tergites 4-5, and dark triangles at ends of forewing veins, but especially a large dark patch at the distal end of Cu_1b. On Quercus spp. often in leaves rolled by caterpillars. In Mediterranean region and south-west Asia. Sexual morphs were recorded in October in Turkey (on Quercus sp., Tuatay & Remaudiere 1965), Iran (on Q. persica; BMNH collection, leg. S. Hodjat) and Italy (on Q. pubescens, as H. parvula, Patti 1983). Vereshchagin & Aukshtikal’nene (1981) made ecological observations.

Hoplochaitophorus Granovsky

Calaphidinae: Myzocallidini

Four North American oak-feeding species apparently related to Neosymydobius. Colonies consist mainly of apterae, which typically have Chaitophorus-like pigmented spiny dorsal hairs.

Hoplochaitophorus dicksoni Quednau Colour of apterae in life is unknown; BL not given. Alatae unknown. On Quercus pungens × arizonica In Arizona, USA (Quednau 1999).

Hoplochaitophorus heterotrichus Quednau Apterae are whitish green with banded antennae, mostly black hind legs, a chestnut brown spot covering the middle of the thorax, and a variably developed pattern of dorsal abdominal markings (Quednau 1971); BL c. 2.2 mm. Alatae have dark-bordered forewing veins. On basal part of lower leaf surface of Quercus prinus in north-eastern USA (Pennsylvania, New York). Oviparae were collected in New York in early October (Quednau 1999).

Hoplochaitophorus quercicola (Monell) Apterae are greenish with dark dorsal markings (see influentialpoints.com/Gallery); BL 1.8-2.1 mm. Alatae have dusky forewings with dark-bordered veins. First described from specimens colonising undersides of leaves of Quercus prinus in Illinois, USA (Monell 1879), but Granovsky (1933) recorded it living in dense colonies along veins on upper surface of foliage, and according to Thomas (1879; as Callipterus quercifolii) it occurs on both sides of leaves along the veins. Recorded from various Quercus spp. (alba, macrocarpa, prinus, rubra, etc.) in North America (except north-west). Apterous males and oviparae (the latter with a long, drawn-out ovipositor) occur in October (in Illinois).

Hoplochaitophorus spiniferus Quednau Colour in life not recorded, hind tibiae of apterae dark brown; BL of aptera c. 2.3 mm. On Quercus bicolor in New York, USA (Quednau 1972). Biology and other morphs are unknown.

Hormaphis Osten-Sacken

Hormaphidinae: Hormaphidini

A genus for three species, two nearctic and one palaearctic, migrating from Hamamelis to Betula, or monoecious on Hamamelis. Conical pouch galls protruding from the upper sides of the leaves are produced on Hamamelis, but the apterous forms on Betula are aleyrodiform with a fringe of white wax, living freely on the leaves. The antennae of alatae are either 3-segmented (North American species) or 5-segmented (the palaearctic species) and there is a single oblique vein in the hind wing. The life cycles and nomenclature of the North American species were elucidated by von Dohlen & Stoetzel (1991), with descriptions of all morphs. Phylogeny and evolution of the genus based on nuclear and mitochondrial DNA sequences were studied by von Dohlen et al. (2002).

Hormaphis betulae (Mordvilko) Plate 3b Small, flattened, subcircular yellowish green or yellowish brown aphids with a fringe of radiating wax filaments, dispersed on undersides of birch leaves, not causing any leaf deformation. Legs and antennae are greatly reduced. Alatae produced on birch have 5-segmented antennae but differ from those of the other European birch-feeding hormaphidine, Hamamelistes betulinus, in having no siphunculi and usually only one oblique hind-wing vein. In Europe and Asia. H. betulae is anholocyclic on Betula spp. in northern and central Europe, and probably in Siberia and central Asia. It is believed that overwintering may occur as a larval stage in the soil or in moss, and that the aleyrodiform generations may have only three larval instars (see Heie 1980b). A holocycle exists in Japan, the aphid described as H. gallifoliae (Monzen 1954), forming conical pouch galls on the upper sides of leaves of Hamamelis japonica, being the primary host form of H. betulae (Aoki & Kurosu 1991). Alatae leave the galls in May-June to deposit larvae on leaves of Betula platyphylla var. japonica. The complete mitochondrial genome of this species has been published (Y.-Q. Li et al. 2015). 2n=18 (Blackman 1986).

Hormaphis cornu (Shimer) Apterous fundatrices form conical galls on the upper sides of the leaves of Hamamelis virginiana (see influentialpoints.com/Gallery). They are dark purplish brown; BL 1.3-2.0 mm. The alatae (BL 1.3 -1.9 mm) that leave the galls in June have only three antennal segments, opaque wings with one oblique vein in the hind wing, and no siphunculi. They migrate to Betula nigra, where they produce very small aleyrodiform apterae, dark brown to black with a fringe of radiating wax filaments, that feed on the undersides of the leaves (see influentialpoints.com/Gallery). There is also a record from B. papyrifera. After three generations on birch leaves sexuparae are produced which fly back to Hamamelis in September-October (von Dohlen & Stoetzel 1991). In eastern North America, mostly at low elevations and latitudes south of 41°N. 2n=?18

Hormaphis hamamelidis (Fitch) Apterous fundatrices in conical galls on Hamamelis virginiana (fig. 128b) are dark purple with white wax on the abdomen; BL 0.9-1.3 mm. The galls are slightly smaller than those of H. cornu and have little or no basal constriction. There is an abbreviated life cycle on witch hazel without host alternation (von Dohlen & Gill 1989). Alatae leaving the galls in late July to September are sexuparae (BL 1.1-1.7 mm), and give rise to sexual morphs on the undersides of the leaves. This species occurs in north-eastern North America at higher elevations and higher latitudes than H. cornu, although there is some overlap (von Dohlen & Stoetzel 1991). 2n=?18.

Hyadaphis Kirkaldy

Aphidinae: Macrosiphini

About 17 palaearctic species associated with Caprifoliaceae and/or Umbelliferae/Apiaceae, with 14 of the 22 recorded host genera in the tribe Apieae. The best‑known species alternate between Lonicera and the aerial parts of Umbelliferae/Apiaceae. Morphologically, Hyadaphis are similar to Lipaphis, and both these genera were placed in Rhopalosiphum in some earlier accounts. Heie (1992) reviewed the species found in Fennoscandia and Denmark, and Blackman (2010) those from UK and Ireland. Remaudière & Halbert (2000) provided keys to all morphs of the three species introduced to America, and Kadyrbekov (2005e) keyed the species in Central Asia, transferring two species (coerulescens and ferganica) from Rhopalomyzus to this genus. Nieto Nafría et al. (2016a) provided a key to apterae of all species except H. veratri. Molecular studies would be useful to better define life cycles and host relationships, and to confirm and clarify the differences from related genera.

Hyadaphis agabiformis (Nevsky) Apterae are yellowish orange with a fine pruinose secretion; BL c. 1.3-1.4 mm. (Specimens in ethanol turn black.) On Bupleurum czimganicum (= B. falcatum ssp. cernuum) in southern Kazakhstan.

Hyadaphis anethi Nieto Nafría, Pérez Hidalgo & Brown Colour of apterae in life is unknown; BL 1.4-1.7 mm. Alatae have secondary rhinaria distributed ANT III 32-43, IV 8-17, V 3-8. On an Anethum species, probably A. graveolens, at high altitude (1,900-2,000 m) in Pakistan. The life cycle is unknown, but two alate males were collected in late August (Nieto Nafría et al. 2016a).

Hyadaphis bicincta Börner Apterae are grey-green, probably pruinose; BL 1.2-2.0 mm. In upwardly-rolled leaves of Lonicera nigra in Europe (France, Austria, Czech Republic).

Hyadaphis bupleuri Börner Apterae are green or brownish green, with darker appendages, BL 1.6-1.9 mm. Specimens remain pale in ethanol (see agabiformis and mongolica). In often dense colonies on upper sides of leaves, young shoots and inflorescences of Bupleurum spp. In Europe (Austria, France, Germany, Hungary, Poland, Czech Republic, Ukraine), and Iran (Rezwani 1987). Monoecious holocyclic with oviparae and apterous males in September (BMNH collection, leg. C. Börner).

Hyadaphis bupleuriphila Barjadze & Özdemir Apterae are pale green with pale brown appendages; BL 1.5-2.0 mm. Alatae have secondary rhinaria distributed ANT III 21-47, IV 2-12, V 0-3. On Bupleurum sp., and also on another unidentified species of Apiaceae (on which dense colonies were found on the inflorescences), in Turkey (Barjadze & Özdemir 2018). [This is the species referred to as “Hyadaphis sp. (Turkey, BMNH colln)” in Blackman & Eastop (2006).]

Hyadaphis coerulescens (Narzikulov) Apterae are yellowish green; BL 1.7-2.6 mm. On Lonicera spp. in Central Asia. Oviparae and alate males occur in September-October (Mukhamediev & Akhmedov 1982, as Rhopalomyzus). Possibly migrating to Bunium sp. or Schrenkia ugamica (Kadyrbekov 2017a).

Hyadaphis coriandri (Das) Plate 13a (Fig.41b) Apterae are dirty greenish in colour with dark green dorsal mottling and rust‑red patches at bases of dark brown siphunculi, variably dusted with white mealy wax (see aphids of Karnataka website); BL 1.3‑2.1 mm. On numerous species of Umbelliferae/Apiaceae (including Anethum, Carum, Coriandrum, Cuminum, Daucus, Ferula, Foeniculum, Pimpinella), living mainly in the umbels. Coriandrum is particularly susceptible to attack (Blackman & Eastop 2000). Occasionally found colonizing plants outside the Umbelliferae/Apiaceae (Mentha, Amaranthus, Glycine). Probably of Asian origin, now in Portugal, Spain, the Mediterranean region, the Middle East, Central Asia, India, Pakistan, Africa, USA (Florida, Hawaii) and South America (Peru, Argentina). Probably anholocyclic over much of its range, but a holocycle with a sexual phase on Lonicera nummulariifolia occurs in Iran, and on three other Lonicera spp. in Kazakhstan (Kadyrbekov 2013f, 2017a). The progeny of the fundatrix all migrate to Umbelliferae/Apiaceae in the second generation (Remaudière & Halbert 2000). 2n = 12*, 13 and 14.

Hyadaphis ferganica (Mukhamediev & Akhmedov) Apterae are dark green; BL 2.3-2.6 mm. On Lonicera nummularifolia in Central Asia (Uzbekistan, Kyrgyzstan). Oviparae and alate males occur in October (Mukhamediev & Akhmedov 1982, as Rhopalomyzus).

Hyadaphis foeniculi (Passerini) Plate 13b (Fig.38g) Apt. greyish green or light green with dark appendages (see also influentialpoints.com/Gallery); BL 1.3-2.6 mm on primary hosts, 1.4-2.0 mm on secondary hosts. Alatae have secondary rhinaria distributed ANT III 36-71, IV 6-28, V 0-2 (-5). On Lonicera spp., especially L. xylosteum, and sometimes Symphoricarpos, curling leaves upwards in spring, migrating to various Umbelliferae/Apiaceae, where they colonise stems, leaves and inflorescences. Common secondary hosts are Angelica, Apium, Conium, Foeniculum, Pastinaca, Peucedanum and Pimpinella, but apparently not Daucus. Widespread in Europe, especially in the north, eastward to Turkey, Iran and Iraq, in North America (New York, New Brunswick, California), and reported to be a major pest of fennel in north-east Brazil (Ramalho et al. 2012). Confused in the literature with H. passerinii, and populations with intermediate conditions can occur, indicating that hybridisation may occur where holocyclic populations are sympatric. There is another undescribed species close to H. foeniculi in Lebanon (BMNH collection, leg. D. Hille Ris Lambers); apterae from L. etrusca have the cauda paler and more pointed than usual. 2n=14 for samples of H. foeniculi from Conium and Foeniculum. One sample from Lonicera had 2n=13 (a foeniculi × passerinii hybrid?), and one sample from Foeniculum had a mixture of 2n=12 and 2n=14 individuals (Blackman & Eastop 2006).

Hyadaphis galaganiae Nevsky Apterae are yellow-green to yellowish tinged with grey, with dark appendages; BL 1.6-2.0 mm. On Muretia (=Galagania) fragrantissima, Ferula spp. and Chaerophyllum bulbosum ssp. prescotti in Kazakhstan and Kyrgyzstan (Kadyrbekov 2005d). The siphunculi are short, and the description does not distinguish it from H. coriandri. H. ferulae Kadyrbekov 1992 is a synonym (Kadyrbekov 2005d).

Hyadaphis haplophylli Kadyrbekov Apterae are yellow or yellowish green with red eyes; BL 1.4-1.8 mm. On inflorescences of Haplophyllum dshungaricum (an anomalous host for a Hyadaphis), attended by ants. There is a subsequent record from Bunium setaceum (Kadyrbekov 2017a; listed in error as B. setosum), which is more likely to be the normal host. In south-east Kazakhstan (Kadyrbekov 2005d).

Hyadaphis levantina Nieto Nafría, Pérez Hidalgo & Brown Colour of apterae in life is unknown; BL 1.4-2.5 mm. Alatae have secondary rhinaria distributed ANT III 44-66, IV 9-23, V 0-2. On Lonicera nummulariifolia at high altitude (1,500-2, 200m) in Israel and Lebanon (Nieto Nafría et al. 2016a). The life cycle is unknown.

Hyadaphis mongolica Szelegiewicz Apterae are pale green to yellowish green, with dark appendages; BL 1.5-1.8 mm. Specimens become black in ethanol. On Bupleurum scorzoneurifolium, colonising especially the flower stems, in Mongolia and North Korea (Lee et al. 2002c).

Hyadaphis parva Nieto Nafría, Pérez Hidalgo & Brown Colour of apterae in life is unknown; BL c.1.0-1.4 mm. The host was apparently not an umbelliferous plant, and was identified with some uncertainty as Andrachne cordifolia (= Leptopus cordifolius). In mountains of Pakistan (Azad Jammu and Kashmir). Other morphs and life cycle are unknown (Nieto Nafría et al. 2016a).

Hyadaphis passerinii (del Guercio) (Fig.38f) Apterae are greyish green or light green with dark appendages (see influentialpoints.com/Gallery); BL 1.2-2.4 mm on primary hosts, 1.0-1.9 mm on secondary hosts. Alatae have secondary rhinaria distributed ANT III 23-45, IV 3-18, V 0 (-2). On Lonicera spp., especially caprifolium and periclymenum, curling leaves upwards in spring, migrating to various Umbelliferae/Apiaceae, particularly Daucus, and also on Conium and Pastinaca, colonising stems, leaves and inflorescences. Anholocyclic on Umbelliferae/Apiaceae in warmer climates. In Europe, especially the south, Mediterranean region, Middle East, Central Asia, Pakistan, India, and introduced to southern Africa, Australia, New Zealand, North and South America. Confused in literature with H foeniculi (q.v.). 2n = 12 for 12 samples from Lonicera and one sample from Daucus (cf. foeniculi).

Hyadaphis polonica Szelegiewicz Apterae are green to yellowish green, coated with wax powder, with dark green appendages; BL 1.6-2.0 mm. On upper part of root of an umbelliferous plant, probably Carum carvi, in Poland, east Kazakhstan (Kadyrbekov 2005d), and recorded also in Finland, from Lonicera, its primary host (Heie 1992).

Hyadaphis tataricae (Aizenberg) Apterae are yellow-green to brownish yellow, coated with greyish wax powder (see aphidtrek.org); BL 1.1-2.5 mm. On Lonicera spp. in Europe, Iran (Rezwani 2010), Central Asia and North America. Monoecious holocyclic, with oviparae and alate males in late August-November (Voegtlin 1984b). Remaudière & Halbert (1999) provide discriminants from H. coriandri and H. foeniculi/passerinii for all morphs. 2n=14.

Hyalomyzus Richards

Aphidinae: Macrosiphini

A Myzus-like genus with nine North American and three little-known Indian species which possibly do not belong here. Hyalomyzus are related to Ovatus but have swollen siphunculi, and apterae have a strongly wrinkled tergum. Three American species apparently host-alternate between woody Rosaceae (usually Crataegus) and secondary host plants in several families; however as pointed out by Voegtlin (1984a), there is in no case any published evidence of host transfer tests to verify the life cycle. The other species are known only from herbaceous plants. Bhattacharya (1994) reviewed the Indian species, and L.K. Ghosh (1987) reviewed the species in Himachal Pradesh, India. Stoetzel et al. (1999) reviewed the genus and provided a cladistic analysis and key to the North American species.

Hyalomyzus collinsoniae (Pepper) (Fig.48c,d) Apterae are greenish to purple or wine-coloured; BL 1.3-2.0 mm. Alatae have dark-bordered wing veins and numerous secondary rhinaria distributed III 42-65, IV 18-42, V 1-9. On Collinsonia canadensis and Salvia azurea (Lamiaceae); also collected on a Crataegus sp., which is presumably the primary host. In eastern USA (New York, Pennsylvania, Mississippi, North Carolina). It is closely related to H. eriobotryae; Stoetzel et al. (1999) discussed the differences, largely restricted to alatae.

Hyalomyzus eriobotryae (Tissot) Plate 18b Apterae are greenish purple to brown, with light fluffy wax at tip of cauda; BL 1.0-2.0 mm. Alatae are Myzus-like but have secondary rhinaria distributed III 40-60, IV 22-40, V 5-19, and no dark dorsal abdominal patch. On young growth of Crataegus spp. in spring in eastern North America (Ontario to Florida), and also recorded from North Dakota and Washington. Originally described from alate ?gynoparae and males collected in December in Florida on Eriobotrya japonica , and at the same time recorded from apple and hawthorn (Tissot 1935). Crataegus spp. are presumably the normal native primary hosts. Apparently heteroecious holocyclic, with the presumed secondary host populations recorded from Labiatae/Lamiaceae; Lycopus virginicus and Collinsonia canadensis in North America, and Hyptis capitata in Cuba (Holman 1974). However host alternation has not been conclusively demonstrated (Stoetzel et al. 1999). Spring populations of Hyalomyzus on Crataegus could be other species so far known mainly or entirely from secondary hosts, such as H. collinsoniae (Stoetzel et al. 1999).

Hyalomyzus fragaricola L.K. Ghosh Colour of apterae in life is unrecorded; BL c. 1.6-1.7 mm. Alatae are unknown. On Fragaria sp. in north-east India (Himachal Pradesh).

Hyalomyzus himachali Bhattacharya Colour of apterae in life is unrecorded; BL 1.9-2.1 mm. Alatae are unknown. On Rubus sp. in north-east India (Himachal Pradesh.

Hyalomyzus jussiaeae Smith Apterae are dirty greenish brown, with reddish area around the siphunculi; BL 1.3-1.9 mm. Alatae have secondary rhinaria distributed III 26-39, IV 9-24, V 3-9. On Ludwigia (= Jussiaea) spp. in Central America (Cuba, Puerto Rico, Honduras) and eastern USA. Also collected on Drosera capillaris and Isnardia (= Ludwigia?) intermedia (as H. tissoti; Nielsson & Habeck 1971), and it has been recorded in large numbers on another semi-aquatic plant, Polygonum punctatum, in Florida (Heppner & Habeck 1997). Possibly host-alternating to Crataegus in parts of USA, but spring populations have not been observed. The only morphs recorded from Crataegus (praeformosa, vicana) are alate ?fundatrigeniae with brown thorax and yellowish brown to brown abdomen, BL 1.0-1.5 mm (Nielsson & Habeck 1971, as tissoti), and sexual morphs found on Crataegus sp. in October (Stoetzel et al. 1999). Secondary hosts are mainly Onagraceae (Ludwigia spp., Isnardia intermedia), but it may colonise various other aquatic or semi-aquatic plants.

Hyalomyzus mitchellensis Smith Apterae are pale to dark green, or light amber to orange, with black siphunculi; BL 1.3-1.9 mm. On Hypericum mitchellianum in North Carolina, USA, living at bases of leaf axils and on flower buds, and on stems under rocks. Also on Hypericum sp. in Pennsylvania (Nielsson & Habeck 1971, as H. sensoriatus) and Maine (aphidtrek.org). Monoecious holocyclic, with oviparae and apterous males in October (original description).

Hyalomyzus monardae (Davis) Apterae are yellow-brown; BL 1.2-1.6 mm. Alatae have secondary rhinaria distributed III 7-9, IV 0-5, V 0-1. On Monarda fistulosa, twisting and curling the leaves (Hottes & Frison 1931, as Phorodon monardae). Widely distributed in USA. [Note: ICZN (Opinion 1818, Sept. 1995) ruled in favour of an application to suppress scrophulariae Thomas in favour of monardae Davis.]

Hyalomyzus orphnophlebos Stoetzel, Jensen & Miller Colour of apterae in life is unknown; BL c.1.6-1.8 mm. Alatae have dark-bordered wing veins, and secondary rhinaria distributed III 37-50, IV 10-17, V 0-4. On Collinsonia serotina, and also collected from Crataegus aestivalis, the presumed primary host (Stoetzel et al. 1999). In USA (Mississippi).

Hyalomyzus pocosinus Stoetzel, Jensen & Miller Apterae are red, with tip of cauda covered in light fluffy wax; BL 1.0-1.7 mm. secondary rhinaria distributed III 22-34, IV 10-21, V 5-10. On lower stem and rootlets of Triadenum virginicum in Maryland, USA. Monoecious holocyclic with oviparae and apterous males in October (Stoetzel et al. 1999).

Hyalomyzus raoi Hille Ris Lambers Colour of apterae in life is unknown; BL 2.3-2.8 mm. Alatae have secondary rhinaria distributed III 35-42, IV 7-11, V 0-1. On Rubus spp. in India. Unlike those of the North American species, alatae have a dark dorsal abdominal patch which may justify its placement in a separate genus, Neohyalomyzus (R.C. Basu et al. 1976; Stoetzel et al. 1999). However because alatae of the two other Indian species are unknown their position is also uncertain, so for the present we retain all three in Hyalomyzus. 2n=8 (Khuda-Bukhsh & Kar 1990).

Hyalomyzus sensoriatus (Mason) Only alatae (?fundatrigeniae) were described from the primary host, Crataegus crusgalli, in Washington D.C. (Mason 1940). They have secondary rhinaria distributed III 45-62, IV 27-43, V 11-17. The secondary host was originally recorded as Hypericum sp., based on collections in Pennsylvania (Nielsson & Habeck 1971), but Stoetzel et al. (1999) identified this material as H. mitchellensis. Alate males and gynoparae collected on Lycopus americana on Long Island, New York (BMNH collection, leg. M.D. Leonard) agree well with sensoriatus, but the life cycle requires confirmation. Also recorded from Quebec, Canada.

Hyalomyzus triangulatus Voegtlin Apterae are pale yellow to greenish yellow in spring, with pale yellow appendages, the siphunculi darker yellow with dusky apices, but the abdomen is darker yellow-green in autumn; BL 1.1-1.6 mm. In leaf axils on Hypericum spp. in eastern USA (Illinois, Maryland). Monoecious holocyclic, with oviparae and alate males in October-November (Voegtlin 1984a).

Hyalopteroides Theobald

Aphidinae: Macrosiphini

One species with the morphological adaptations often associated with grass feeding; elongate body, short blunt RIV+V and short appendages, as well as very short, flangeless siphunculi.

Hyalopteroides humilis (Walker) Plate 14f Apterae are pale yellowish, with pale appendages (see influentialpoints.com/Gallery); BL 2.3‑3.0 mm. On upper sides of leaves of Dactylis glomerata, leaves of which are turned brown by heavy attacks. Usually specific to this host, with only single records from grass species in other genera (Bromus, Holcus, Muhlenbergia, Tridens). Throughout Europe, Iran, Kazakhstan (Kadyrbekov 2017a), and introduced to USA (Essig 1953a) and South America (Argentina; Ortego et al. 2004). Monoecious holocyclic with oviparae and alate males in October (Heie 1994). Dransfield (1979) reported on parasitoids. 2n=16.

Hyalopterus Koch

Aphidinae: Aphidini

A genus once used in a much broader sense to include most Aphidinae with small siphunculi, but now restricted to a group of very similar species of elongate-bodied, mealy aphids that alternate between Prunus spp. and Phragmites. In recent years various studies of this group utilising morphological and molecular techniques (Spampinato et al. 1988, Mosco et al. 1997, Poulios et al. 2007, Lozier et al. 2007, 2008, Rakauskas et al. 2013) have all agreed in demonstrating that there are three taxa utilising different Prunus species as their preferred primary host. However more recent work in China (T. Liu et al. 2020) has revealed a more complex situation with at least four putative species (see under H. arundiniformis). It is presumed that all Hyalopterus spp. migrate to Phragmites, Arundo and other graminaceous hosts in watery situations, but the secondary host populations have not been studied in this way and the identity of the anholocyclic populations on Phragmites in the tropics, e.g. along African rivers, needs to be established. Lozier et al. (2008) provided a key to apterae on Prunus, and Rakauskas et al. (2013) and Moreno-González & Nieto Nafría (2018) provided some additional discriminatory characters. Basky (1982) and Ben Halima-Kamel et al. (2013) studied natural enemies in Hungary and Tunisia. Aphidius transcaspicus is a parasitoid specific to Hyalopterus (Lozier et al. 2009).

[Mutillaphis prunisucta Zhang & Zhang, described from Prunus mume in China (L. Zhang & G. Zhang 2001), seems to be based on fundatrices of a Hyalopterus, possibly amygdali.]

Hyalopterus amygdali (Blanchard) (fig. 88B) Apterae are rather elongate-bodied, pale green mottled with darker green, covered with white wax meal; BL 1.7-2.6 mm. On undersides of leaves of Prunus dulcis, and sometimes on P. armeniaca. Records from other Prunus require confirmation. In the Mediterranean area (Spain, Italy, Greece, Israel, Algeria, Tunisia) aqnd Central Asia, and possibly also in North America (Smith & Parron 1978). Records from China may be based on misidentification (T. Liu et al. 2020). There is also a record from Hungary, where colonies were found on the shoots and leaves of a grapevine (Ripka 2011; possibly this record should be referred to H. arundiniformis). Migration occurs to Phragmites and possibly other reeds or grasses, but the secondary host populations have not been distinguished from those of the other Hyalopterus species. 2n=10.

Hyalopterus arundiniformis Ghulamullah (= H. persikonus Miller, Lozier & Foottit; see Favret et al. 2017) Apterae in life presumably very similar to amygdali; BL 1.9-2.9 mm. On undersides of leaves of Prunus persica, and sometimes on P. armeniaca, but molecular work has shown that there may also be cryptic species under this name with other specific primary host associations; for example samples from P. salicina and P. cerasifera f. atropurpurea in China (T.Liu et al. 2020). Apterae of the H. arundiniformis group in Spain have longer median dorsal cephalic hairs than those of H. amygdali (Moreno-González & Nieto Nafría 2018). Alatae have not been discriminated morphologically from those of H. amygdali, but Phragmites and Arundo have been confirmed as secondary hosts by molecular work (T. Liu et al. 2020). Confirmed records are from Spain, Italy, Greece, Iran, Iraq, Georgia, Afghanistan (including type locality) and Algeria (Peréz Hidalgo et al. 2012b, as H. persikonus), and members of the H. arundiniformis species group are now established to occur commonly and widely in China (T. Liu et al. 2020). Distribution is otherwise uncertain, but apparently this species group is not in North America. Studies of H. amygdali on peaches, by Darwish (1992) in Egypt and Huang et al. (1986) in China, for instance, may need to be referred to the arundiniformis group. Aphids identified as H. persikonus have been the subject of studies in China on differences in gene expression between populations on primary and secondary host plants (Cui et al. 2017).

Hyalopterus pruni (Geoffroy) Plate 9d (fig. 88A) Apterae rather elongate-bodied, pale green with darker green mottling, covered with white wax meal; BL 1.5-2.6 mm. On undersides of leaves of Prunus domestica, and sometimes on other Prunus spp., especially P. armeniaca, but not P. dulcis. Infested leaves do not curl (influentialpoints.com/Gallery). Alatae have a green abdomen with white wax patches on each segment (see aphidtrek.org). Migration occurs to Phragmites, or sometimes to Arundo donax; for its appearance on Phragmites see influentialpoints.com/Gallery. Widely distributed in Europe and Asia and introduced to North America, but records from Africa and Australia are based on secondary host populations and are possibly another species. Records from China (except north of the Tianshan mountains) need to be referred to the H. arundiniformis (T. Liu et al. 2020). 2n = 10.

Hybothoracaphis Chen, Jiang & Qiao

Hormaphidinae: Nipponaphidini

A genus for one species described from Quercus in Tibet with a prominent spinal ridge, a single cluster of pustules on the head, and paired and single clusters of pustules mid-dorsally on the otherwise rather smooth prosoma.

Hybothoracaphis laevigata Chen, Jiang & Qiao Apterae are dark brown, flattened, oval: BL 0.8-1.3 mm. Along veins on undersides of leaves of Quercus aquifolioides in Tibet (Chen et al. 2016). Other morphs and life cycle are unknown.

Hydaphias Börner

Aphidinae: Macrosiphini

A palaearctic genus on Galium and related Rubiaceae, characterised by the thin, often inwardly curved, flangeless siphunculi. Heie (1992) provided an account of the species in Scandinavia.

Hydaphias carpaticae Mamontova-Solukha Apterae are dirty yellow with pale green spinal stripe; BL c.1.9-2.0 mm. Alatae have c.26-28 secondary rhinaria on ANT III and c.5-6 on IV, whereas alatae of other described species have 10-20 on III and 0-4 on IV. Colonising stems of Galium intermedium in Ukraine, and also reported from G. ruthenicum, (= verum) in Kazakhstan (Kadyrbekov 2012a).

Hydaphias molluginis Börner Plate 13g (Fig.29k) Apterae are dirty green or greenish yellow with dark appendages; BL 1.4-2.0 mm. In ant-attended colonies close to ground, on shoots and flower stems that become deformed and stunted. Described from Galium mollugo in Germany. The name H. hofmanni Börner is commonly applied to very similar aphids on G. verum that tend to have smaller marginal abdominal tubercles, but the development of such tubercles is notoriously variable and environmentally dependent. None of the other characters that have been used in the literature were consistent in separating the specimens from G. verum and G. mollugo in the BMNH collection, and it seems probable that experimental work may show that one variable species is involved. H. helvetica Hille Ris Lambers, described originally from G. verum and completely lacking marginal abdominal tubercles, is also likely to be a synonym. Aphids of the H. molluginis group occur throughout Europe, and across Asia to east Siberia, Korea and China (Jiang et al. 2005, as H. hofmanni). Oviparae occur in September.

Hydaphias mosana Hille Ris Lambers (Fig.29j) Apterae are dull, pale yellowish green to dark green with dusky appendages; BL 1.0-1.6 mm. On subterranean parts of Galium mollugo and G. verum, attended by ants. Throughout Europe, including England and Spain (BMNH collection, leg. J.H. Martin), eastward to Turkey, Russia (Volgograd) and Kazakhstan (Kadyrbekov 2017a). Oviparae and apterous males occur in September (original description).

Hydronaphis Shinjii

Aphidinae: Macrosiphini

Five Myzus-like species in east Asia with spinulose head, long dorsal hairs and a short cauda, resembling Cryptaphis but the hairs are not apically expanded and the siphunculi are slightly clavate. One species alternates between Corylopsis and the roots of Impatiens in Japan, while the others are recorded from various presumed secondary host plants, usually feeding on the roots. Miyazaki (1971) reviewed the Japanese species, but in Japan there are also possibly undescribed species from Hamamelidaceae, Hydrangeaceae, Marantaceae, Myrtaceae and Orchidaceae (M. Miyazaki, pers. comm.).

Hydronaphis calanthes Sorin Apterae are reddish orange with blackish brown siphunculi and a pale cauda; BL c.2.1-2.3 mm. On Calanthe discolor in Japan, attacking the plant at ground level. Alatae and other morphs are undescribed.

Hydronaphis colocasiae Raychaudhuri, Raha & Raychaudhuri Colour and size of apterae are unknown. BL of al. c. 2.7-2.9 mm. Alatae have secondary rhinaria distributed III 9-13, IV 0, and a black dorsal abdominal patch (Singh et al. 1980). On Colocasia sp. in India (Nagaland).

Hydronaphis impatiens Shinji Plate 17d In early-spring colonies living on undersides of leaves of Corylopsis spp., without causing any leaf deformation. Apterae (fundatrices) are pale yellow, with dark appendages and black-tipped siphunculi; BL c. 2.4 mm. Progeny of fundatrices are all alate, migrating in April-May to found colonies on roots and subterranean stems of Impatiens spp.; the emigrant alatae have a yellow abdomen with a black dorsal patch. Apterae on Impatiens are black with black antennae, siphunculi and cauda; BL c.2 mm. Alate males and gynoparae return to Corylopsis in October (S. Takahashi 1986). Only known from Japan and Korea (Kim et al. 2008).

Hydronaphis laporteae Miyazaki Apterae are dull yellow with dorsum dark brown to black, siphunculi are pale with black apices, and cauda is pale; BL 1.6-1.8 mm. Alatae have secondary rhinaria distributed III 28-37, IV 8-14, V 2-6. On undersides of leaves (original description), or perhaps more commonly on roots (S. Takahashi 1986), of Laportea bulbifera in Japan.

Hydronaphis liriopes Aida Apterae are whitish yellow, with a dark green to blackish central abdomen patch, black siphunculi and a pale cauda; BL 1.6-1.9 mm. Alatae have secondary rhinaria distributed III 33-43, IV 10-18, V 0-4. On roots of Liriope platyphylla (= L. muscari) in Japan.

Hyperomyzus Börner

Aphidinae: Macrosiphini

About 20 species associated with Ribes as primary hosts, and mostly having liguliflorous Compositae/Asteraceae as secondary hosts, but some (subgenus Hyperomyzella) migrating to Orobanchaceae. Except for the swollen siphunculi they resemble Nasonovia, which have similar biology. Hyperomyzus s.str. (6 species) and subgenus Hyperomyzella (1 species) are of palaearctic origin, while 9 of the 14 known species of subgenus Neonasonovia are nearctic. Hille Ris Lambers (1949), Heinze (1961), Müller (1969b), Heie (1994) and Blackman (2010) gave accounts of the European species, and Nieto Nafría et al. (2017) reviewed and keyed North American species of subgenus Neonasonovia. Many of the North American species were originally described in the genus Amphorophora.

Hyperomyzus carduellinus (Theobald) Plate 21e Apterae are pale green with brown apices to segments of antennae and legs (see aphids of Karnataka website); BL 1.8-2.7 mm. Alatae generally have a more solid and compact central abdominal patch than in the very similar H. lactucae, and secondary rhinaria distributed III 34-64, IV 14-31, V 5-13. On various Compositae/Asteraceae (Bidens, Gynura, Ixeridium, Lactuca, Prenanthes, Reichardia, Senecio, Sonchus, Taraxacum, Tridax). In Africa south of the Sahara, India, Japan, Korea, south-east Asia, Australia, New Zealand (?), Fiji, Hawaii, New Caledonia (Mille et al. 2020), and more recently recorded from Florida (Halbert et al. 2000) and Argentina (Ortego et al. 2004). Apparently entirely anholocyclic. Rebijith et al. (2013) found CO-I barcode differences indicative of two cryptic species in Indian samples of H. carduellinus, but the possibility needs to be considered that H. lactucae was also present in these samples. 2n=12.

Hyperomyzus (Neonasonovia) gansuensis (Zhang, Chen, Zhong & Li) Apterae are green, BL 2.0-2.4 mm. On Zanthoxylum nitidum (unlikely to be the true host) in Gansu Province, China (G. Zhang 1999, as Liosomaphis).

Hyperomyzus (Neonasonovia) hieracii (Börner) Apterae are shining brownish black, with black siphunculi; BL 1.7-2.0 mm. On upper parts of stems and in inflorescences of Hieracium spp. in Europe (UK, Denmark, Germany, Austria, Switzerland). Oviparae and apterous males have been reared in Scotland in October (Blackman 2010) and oviparae have been found in northern Russia (Murmansk) in early August. Stekolshchikov & Buga (2006a) provided a full description of the sexual morphs, and of a fundatrix.

Hyperomyzus (Neonasonovia) inflatus (Richards) Colour of apterae in life not noted; BL 1.6-2.9 mm. Alatae have a black dorsal abdominal patch and secondary rhinaria distributed III 52-70, IV 10-18, V 2-5. The aptera was described from Solidago sp. in Minnesota, USA, but subsequent records (Maine, North Carolina, Pennsylvania: BMNH collection) have all been from Hieracium, and this is probably the true host, on which it is apparently monoecious holocyclic. Nieto Nafría et al. (2017b) provided a redescription.

Hyperomyzus lactucae (L.) Plate 21d Apterae are opaque green with pale appendages (see influentialpoints.com/Gallery); BL 2.0-3.2 mm. Alatae have a rather fragmented dark dorsal abdominal patch, pale brownish siphunculi, and secondary rhinaria distributed III 30-62, IV 4-22, V 0-8. Heteroecious holocyclic in temperate regions with Ribes spp., esp. nigrum, as primary hosts, feeding on undersides of young leaves which curl slightly and acquire yellow spots. Migrating in late May-June to form colonies on upper parts of stems and flowerheads of Sonchus spp, and occasionally other related Compositae/Asteraceae. Common and almost world-wide, except southern Africa (but it does occur in the highlands of Kenya). It is anholocyclic on Sonchus in warmer regions, but in the tropics it tends to be replaced by H. carduellinus (q.v.). See also Blackman & Eastop (2000). The subspecies status (as Hyperomyzus lactucae ssp. asiatica) given to some Central Asian populations by Narzikulov & Umarov (1969) does not seem to be justified. 2n=12.

Hyperomyzus lampsanae (Börner) Apterae are ivory white, with dark apices to segments of antennae and legs (see influentialpoints.com/Gallery); BL 2.3-3.0 mm. Alatae have secondary rhinaria distributed III 45-75, IV 18-32, V 3-12. On undersides of leaves of Lampsana communis. In north-west, north and central Europe, and recorded also from Turkey (Özdemir et al. 2005) Monoecious holocyclic with oviparae and alate males in October (Hille Ris Lambers 1953). 2n =12.

Hyperomyzus luteus (Mordvilko) Apterae are “pale greyish, greenish yellow” (Shaposhnikov 1964); BL has apparently not been recorded. On Ribes alpinum in spring, migrating to Orobranchaceae (Pedicularis sp., Rhinanthus major). The life cycle was established by Börner (1952). In central and eastern Europe, and Central Asia (Kirghizia). This species has not been properly described, and the specific name is based on a proposed synonymy with Rhopalosiphum luteum Mordvilko 1929 by Börner (1952). This is regarded as a nomen nudum in catalogues, but in the absence of alternative evidence we follow Shaposhnikov (1964) and Holman (2009) in applying the name and accepting the synonymy.

Hyperomyzus (Neonasonovia) nabali (Oestlund) Apterae are shiny green, with siphunculi brown on distal half; BL 2.0-3.2 mm. Apterae have secondary rhinaria distributed III 3-69, IV 0-24, V 0-4 (higher numbers presumably are alatiform specimens), and alatae have them distributed III 31-130, IV 28-53, V 0-15. On stems, flowerheads and undersides of leaves of Prenanthes (= Nabalus) alba and P. altissima in eastern North America (but there are also two alatae from Utah in BMNH collection, leg. G.F. Knowlton, that appear to be this species, and a possible record from Hieracium albiflorum in Oregon). Heteroecious holocyclic, with sexual phase on Ribes gracile and R. glandulosum. Spring forms were described by Mason (1925) as Amphorophora hayhursti and A. pergandei). Nieto Nafría et al. (2017b) provided full descriptions of apterous exules, gynoparae and oviparae.

Hyperomyzus (Neonasonovia) niger (McVicar Baker) Apterae are very shiny black; BL 1.3-1.8 mm. Alatae have secondary rhinaria distributed III 31-39, IV 15-17, V 5-9 (2 specimens). On Stevia spp. in Mexico. Apterae were redescribed by Hille Ris Lambers (1974), and additional data for both apterae and alatae were provided by Nieto Nafría et al. (2017b).

Hyperomyzus (Neonasonovia) nigricornis (Knowlton) Colour of apterae in life unknown; BL 2.3-3.3mm. Host-alternating between Ribes spp. and one or more composite secondarey hosts. Apterae from Ribes have 6-16 secondary rhinaria on ANT III only, and those from secondary hosts have 17-44 on III and 0-3 on IV. Alatae have secondary rhinaria distributed III 38-63, IV 2-11 (from Ribes) or III 62-98, IV 7-31 (from secondary host(s). In western USA (Idaho, Oregon, Utah, Washington). The secondary hosts require confirmation; hosts identified to species are Hieracium albiflorum (see aphidtrek.org) and Hemizonella (= Madia) minima. Descriptions of apterae and alatae from primary and secondary hosts were provided by Nieto Nafría et al. (2017b). It is very similar to H. ribiellus, but with darker tibiae and siphunculi.

Hyperomyzus pallidus Hille Ris Lambers Apterae are opaque greenish or yellowish white with pale appendages (see influential points/Gallery); BL 2.3-3.5 mm. Alatae have a more-or-less solid black dorsal abdominal patch, and black siphunculi (cf. lactucae). In spring colonies at shoot tips and on undersides of young curled leaves of Ribes uva-crispa, and sometimes R. alpinum). Heteroecious holocyclic, migrating to Sonchus spp., where it colonises undersides of lower leaves (Hille Ris Lambers, 1953). In Europe, Iran, Turkey, Central Asia, east and west Siberia, and (introduced to) North America, and now also in South America (Nieto Nafría et al. 2018).

Hyperomyzus (Neonasonovia) petiolaris (Knowlton & Allen) Apterae are green, siphunculi are darker on distal halves; BL 2.1-2.4 mm. Apterae have 12-15 secpndray rhinaria on ANT III only, alatae have them distributed III 50-77, IV 2-8. On foliage of Ribes spp. (nigrum, hudsonianum var. petiolare) in western USA (Washington, Utah). Probably monoecious holocyclic on Ribes (oviparae in late July; Palmer 1952, as Amphorophora fronki).

Hyperomyzus (Neonasonovia) picridis (Börner & Blunck) Plate 21g Apterae are shining yellow-green with brownish dorsal intersegmental markings and dusky siphunculi (see influential points/Gallery); BL 2.4-3.0 mm. Heteroecious holocyclic with Ribes alpinum as the primary host, migrating to Picris spp., Crepis spp. and Rhagadiolus stellatus. Also recorded from Hieracium sp. (Theobald 1926, as Amphorophora hieracioides), but the host plant may have been misidentified. Colonies on Picris are usually fairly small and occur just beneath the inflorescences. Widespread in Europe, in North Africa (Tunisia; Boukhris-Bouhachem et al. 2007) and also in Ontario, Canada (BMNH collection) and Australia (Tasmania 1999; Brumley 2020). Recently also reported to occur in Kazakhstan (Kadyrbekov 2012a), but on primary and secondary hosts (Ribes nigrum and Hieracium umbellatum) different from those used by European populations, so further confirmation of identity seems to be required. Oviparae and alate males appear in October (Hille Ris Lambers 1949). 2n=12.

Hyperomyzus (Neonasonovia) pullatus Hall & Garraway Apterae are shining black, BL 1.2-2.4 mm. Described from a Eupatorium sp. in Jamaica (Hall & Garraway 2009); previously it had been collected on a plant of the same genus in Costa Rica, and in quite large numbers from plants of several composite genera (Ageratina, Alchemilla, Bidens, Eupatorium, Galinsoga, Simsia) in Mexico. Nieto Nafría et al. (2017b) described the alata, and studied morphological variation between samples from different host genera. The life cycle is unknown.

Hyperomyzus (Hyperomyzella) rhinanthi (Schouteden) Plate 21f Apterae are yellowish green, green or dark green with extensive shiny black dorsal sclerotisation includng a large abdominal patch, antennae and legs mainly black, siphunculi and cauda black (see influential points/Gallery); BL 2.4-3.0 mm. Spring colonies feed on young growth of Ribes spp. (aureum, rubrum), curling and distorting leaves into leaf-nests. Heteroecious holocyclic, migrating to flowers of certain Orobanchaceae; Rhinanthus spp. (esp. major), Euphrasia spp. (Heie 1994) and Pedicularis verticillata (Austria, BMNH collection, leg. H.L.G. Stroyan). It seems to prefer cool moist habitats. In Europe, including Iceland and Faroes, western Siberia, Transcaucasia, Kazakhstan (Kadyrbekov 2017a) and Canada (Quebec: Cloutier et al. 1986). 2n=12.

Hyperomyzus (Neonasonovia) ribiellus (Davis) Apterae are yellow-green to green, with pale siphunculi that are only very slightly swollen on distal half; BL c.1.9-2.1 mm. Apterae have 7-37 secondary rhinaria on ANT III, alatae have them distributed III 30-43, IV 0(-1). In ant-attended colonies on shoot tips and undersides of young leaves of Ribes spp. in North America. Monoecious holocyclic with alate males (Palmer 1952, as Amphorophora ribiella). Davis (1909) provided a full description of this aphid under the name Macrosiphum cynosbati. 2n=12.

Hyperomyzus (Neonasonovia) sandilandicus (Robinson) Apterae are green, BL 2.2-2.9 mm. Alatae have markedly tuberculate secondary rhinaria distributed III 39-63, IV 0 (-2). On Crepis tectorum in Canada (Manitoba).

Hyperomyzus sinilactucae Zhang Apterae are unknown. Alatae (4 specimens) are similar to those of H. lactucae but with more secondary rhinaria (III 61-94, IV 19-32, V 7-8). On Cirsium segetum (which may not be the true host) in Hebei Province, China.

Hyperomyzus (Neonasonovia) thorsteinni Stroyan Apterae are bright yellow-green, shining; BL 1.5-2.0 mm. On Euphrasia spp., on young growth and in inflorescences which become slightly curved and stunted. In Iceland, Norway and UK. Monoecious holocyclic, with alate males (Stroyan 1972b, as H. boerneri ssp. thorsteinii).

Hyperomyzus (Neonasonovia) zirnitsi Hille Ris Lambers Apterae are shiny green; BL 2.2-2.9 mm (spring), 1.8-2.4 mm (summer). Secondary rhinaria of alata are distributed III 75-101, IV 16-34, V 0-4. In spring colonies on shoot apices, petioles and curling leaves of Ribes spp. (alpinum, rubrum). In Europe, and Mongolia. Heteroecious holocyclic, migrating to Euphrasia spp. (Ossiannilsson 1959) and also Pedicularis sp. A subspecies with a shorter cauda and fewer secondary rhinaria, H. zirnitsi ssp. boerneri Prevost, lives monoeciously on Euphrasia rostkoviana in Switzerland and northern Italy, and its alatae are reported from Finland (Heikinheimo 1990).

Hysteroneura Davis

Aphidinae: Aphidini

One originally North American species in which the cauda is long and pale, the antennal terminal process is long and the hind wing has only one oblique vein.

Hysteroneura setariae (Thomas) Rusty Plum Aphid Plate 9a Apterae are rusty brown with contrastingly pale sections of the antennae and tibiae, black siphunculi and a notably pale cauda (see aphids of Karnataka website); BL 1.3-2.1 mm. In colonies at the bases of the spikelets of many genera and species of Poaceae, sometimes on leaves or unripe seeds, often attended by ants. It occurs sometimes on Cyperaceae and on seedlings of oil palms and coconuts. Heteroecious holocyclic in temperate North America with a sexual phase usually on Prunus domestica. Ant-attended populations may persist on plum throughout the year (Gillette & Taylor 1908). Of North American origin, it was apparently introduced to North Africa in the 1940’s, and anholocyclic populations are now widely distributed throughout the warmer parts of the world after rapid spread in the mid-1960’s. Carver (1976) found sexuales and eggs on P. persica in Australia, but it is not known whether the holocycle was completed. See also Blackman & Eastop (2000, p.282-3). 2n=12.