The Aphids

A genus for one species related to Aploneura and Slavum but with more normal distribution of rhinaria on the antennae of the alatae, which fold their wings roof-like inrepose. The large characteristic horn-like galls can be formed on several species of Pistacia.

Baizongia pistaceae (Linnaeus) Plate 1h Producing large, elongate horn-like galls 15-22 cm long (fig. 126C) on Pistacia spp., especially P. palaestina, in the Mediterranean area, and also in north-west India. Large numbers of alatae, BL 1.8-2.0 mm (fig. 81G), emerge in September-November and fly to roots of Poaceae (e.g. Agrostis, Avena, Dactylis, Festuca, Poa, Triticum). Apterae on grass roots are whitish or pale yellow, producing wax in threads; BL 1.6‑2.3 mm. They are invariably attended by ants, especially Lasius flavus, and may overwinter in ants’ nests.

The holocycle takes two years (fig. 2). Anholocyclic populations occur on grass roots more widely in Europe, the Middle East, north Africa, Kenya and the Indian subcontinent. Roberti (1983) and Wertheim (1954) gave accounts of this aphid in Italy and Israel respectively. Wool (1990, 2002), and Lamb et al. (2012), analysed data from a 20-year population study on Pistacia in Israel.

Histology of the galls on Pistacia terebinthus in Spain was studied by Álvarez (2012). Wool & Kurzfeld-Zexer (2008) studied colonies on cereal roots in culture, rearing them successfully on oats (Avena sativa and A. sterilis) but not on wheat or barley.

Wool (2011/12) provided a comprehensive review of this species. 2n=24.

Berberidaphis Narzikulov

Aphidinae: Macrosiphini

One species in Central Asia superficially resembling Liosomaphis but with better developed antennal tubercles, longer antennae and a short rounded cauda.

Berberidaphis lydiae (Narzikulov) Apterae are shiny pale yellow, BL 1.7-2.7 mm. On undersides of leaves of Berberis spp. in mountainous regions of Tajikistan, eastern Kazakhstan (Kadyrbekov 2009a) and northern Pakistan (Naumann-Etienne & Remaudière 1995). Oviparae and small apterous males appear in October-November (Narzikulov 1957, as Liosomaphis lydiae).

Betacallis Matsumura

Calaphidinae: Calaphidini

Six species are known, all from east and south-east Asia. All viviparae are alate, active insects (behaving like Euceraphis), and have a characteristic transverse dark brown to black bar across the ventral side of the head between the eyes. Most of the species are recorded from Betulaceae, which are also the hosts of several related genera. One species (prunicola) has apparently become associated with Prunus. The Indian species are reviewed by Chakrabarti (1988) and A.K. Ghosh & Quednau (1990), the Japanese species by Higuchi (1972), and the Chinese species by Qiao et al. (2005b).

Betacallis alnicolens Matsumura Alatae are pale green with dark tibiae, large (about 3.5 mm), very active, on leaves of Alnus spp. (hirsuta, japonica, matsumurae) in Japan, Korea and China, and also collected from Corylus mandshurica in Korea (BMNH collection). 2n=22.

Betacallis luminifera Zhang Alatae yellow; BL 2.6-3.5 mm, on leaves of Betula luminifera in Zhejiang, China. Very similar to, and possibly synonymous with, B. sikkimensis, but with more secondary rhinaria.

Betacallis odaiensis Takahashi Alatae are pale lemon-yellow to apricot, with black basal, sensoriated, part of third antennal segment, black tibiae, two black bars on the dorsal abdomen, and black siphunculi; BL 2.4-3.1 mm. On leaves of Betula spp. (ermannii, maximowicziana, platyphylla var. japonica) in Japan. (Indian records of odaiensis refer to other species.) 2n=22.

Betacallis prunicola R.C. Basu, A.K. Ghosh & Raychaudhuri Alatae light yellowish green with dark tibiae, dark transverse bars on ABD TERG 3-5 or 4-5 and dark siphunculi; BL 1.9-2.2 mm (A.K. Ghosh & Quednau 1990). Described from adults and immatures on Prunus cerasus in Meghalaya, India (R.C. Basu et al. 1975), and subsequently collected from other Prunus spp. so it now seems likely that this is a valid host association. This species is also recorded from Arunachal Pradesh, Manipur and West Bengal, and from China and the Philippines. It can be distinguished from other Betacallis species by the long ultimate rostral segment (R IV+V 1.5-2.0 × HT II).

Betacallis querciphaga R.C. Basu, M.R. Ghosh & Raychaudhuri Alatae light grenish yellow with dark markings like B. odaiensis except for the dorsal longitudinal head stripe and differences in wing vein pigmentation; BL 2.4-3.2 mm. In India (Manipur, Meghalaya, Sikkim, West Bengal), the Philippines (Chakrabarti 1988) and Thailand (BMNH collection). It was described from Quercus sp., and alatae and immatures are also recorded from Castanospermum sp.; nevertheless, specimens from Betula alnoides in Thailand, from B. luminifera in China (Qiao et al. 2005b) and from B. utilis in West Bengal (BMNH collection) indicate that the normal host association is probably with Betula. Parasitised by Trioxys (Betuloxys) assamensis in India (Raychaudhuri 1990).

Betacallis sikkimensis R.C. Basu, M.R. Ghosh & Raychaudhuri BL of alata 2.4-3.2 mm, appearance in life unrecorded but presumably pale, with third antennal segment and tibiae darkened only at apices (cf. other species). The siphunculi are black on distal half only and there is often a consolidated or broken dark central patch on abdominal tergites 4 and 5. Described from Quercus sp. – which is probably only a reserve host – in Sikkim, India (R.C. Basu et al. 1974); since collected many times in Uttar Pradesh from Betula alnoides and Betula sp. (Chakrabarti 1988, and BMNH collection). Chakrabarti (1988) described the alate male and apterous ovipara from Betula. 2n=20 (Khuda-Bukhsh & Pal 1983b).

Betulaphis Glendenning

Calaphidinae: Calaphidini

A genus of 6 or 7 species rather evenly distributed through the holarctic on Betula. They are small, rather flat oval aphids with short 6-segmented antennae, the PT being about as long as the base of ANT VI. Viviparae have a short, conical cauda and a bilobed anal plate. Oviparae have dark dorsal markings and the posterior abdomen is extended as an “egg-laying organ”. Males are apterous with extensive dark dorsal markings. Fundatrices are unusual in being either apterous or alate (at least in B. quadrituberculata). Stroyan (1977) gave a detailed generic diagnosis, and accounts are available for North America (Richards 1961), Canada (Richards 1969b), Fennoscandia and Denmark (Heie 1982), India (Chakrabarti 1988, A.K. Ghosh & Quednau 1990) and China (Qiao et al. 2005b). There has been much confusion about the number of species in the brevipilosa/quadrituberculata group, due to seasonal variation in hair lengths. Here, B. helvetica Hille Ris Lambers is regarded as a synonym of B. brevipilosa, although a study of haemolymph proteins by Karwanska (1987) suggests that it may be a distinct species. Parasitised by Aphidius aquilus, which also attacks other related aphids feeding on Betula (Stary 1973).

Betulaphis brevipilosa Borner (= helvetica Hille Ris Lambers) (fig. 31B) Apterae are pale to grass-green aphids, BL 1.5-2.0 mm, with the tips of the antennae and the tarsi dark. Alatae almost always have a dark dorsal abdominal patch. This species is frequently confused in the literature with B. quadrituberculata. It seems to feed mostly on Betula pendula, usually on the upper surfaces of the leaves, occurring in north, central and eastern Europe (but not the UK), Central Asia (Kazakhstan; Kadyrbekov 2017a), and introduced into North America and New Zealand. However, Buga (1995) reports its occurrence on several other native and introduced Betula spp. in Belarus. Accounts are given by Cottier (1953, as B. quadrituberculata) and Heie (1982). Ecology of introduced populations in northern California was studied by Hajek & Dahlsten (1988). B. viridis Richards 1969 seems to be this species. 2n=20.

Betulaphis hissarica Narzikulov Appearance in life is unknown. BL of aptera about 1.7-1.9 mm. Chakrabati (1988) redescribed both apterous and alate viviparae, and also apterous oviparae and males. Recorded from various Betula species (hissarica, alnoides, turkestanica, utilis) in Kazakhstan, Tajikistan, Pakistan and India (Himachal Pradesh, Uttar Pradesh).

Betulaphis japonica Takahashi Appearance in life and biology unrecorded, and alatae are unknown. BL of aptera 1.1-1.6 mm. Found on various Betula species in Japan, Korea, and Mongolia. For further details see Higuchi (1972). Similar to the short-haired form of B. quadrituberculata, but with a relatively shorter R IV+V.

Betulaphis longicornis Quednau & Chakrabarti Apterae yellow: BL 2.3-2.9 mm. The alatae lack a dorsal abdominal dark patch, and have numerous (26-35) closely-spaced elongate transverse secondary rhinaria an antennal segment III. Apterous and alate viviparae are described by Quednau & Chakrabarti (1980), and apterous oviparae and males by Chakrabarti (1988). Recorded from Betula alnoides and B. utilis in India (Assam, Bhutan, Himachal Pradesh, Uttar Pradesh, West Bengal) and Pakistan.

Betulaphis pelei Hille Ris Lambers (fig. 31C) Apterae are yellow with dark tarsi and tips of antennae, BL 1.4-1.7 mm. Alatae sometimes have a dark dorsal abdominal patch. On Betula nana with a boreo-alpine distribution in North Europe, Iceland and Greenland (Heie 1982). Aphids occurring on Betula glandulosa, described from Baffin Island as Betulaphis arctosetis Richards 1961, are probably also this species. It has also been reported to occur on B. albosinensis in western China (Qiao et al. 2005b). 2n=20

Betulaphis quadrituberculata Kaltenbach (fig. 31A,D) Apterae have BL 1.5-2.0 mm and are pale yellowish green or pale yellow to almost white, sometimes with patches of darker pigment (especially in autumn), normally feeding on the undersides of leaves (see influentialpoints.com/Gallery). Alatae sometimes have a dark dorsal abdominal patch. Recorded from various Betula species, especially B. pubescens and also occasionally found on Alnus incana. Widely distributed in Europe (including Iceland) and across Asia to Mongolia and China, and introduced into North America. Apterous males and oviparae may be found from September to November. There is much variation in length of the spinal and pleural hairs on the anterior abdominal tergites which causes short-haired specimens of this species, found particularly in the summer months, to be confused easily with B. pelei and B. brevipilosa. B. aureus Richards 1961, described from Betula sp. in Canada, seems to be this short-haired form of B. quadrituberculata. Varty (1964) has studied its biology in New Brunswick, Canada. See also Heie (1982) and Stroyan (1977). 2n=20.

Bicaudella Rusanova

Calaphidinae: Panaphidini

Five species in the Middle East and Central Asia closely related to Therioaphis but specific to Astragalus (and the related genus Astracantha) with a conical dorsal process on the knob of the cauda. Quednau (2003) restored the group to full generic status, and provided illustrations of all available morphs.

Bicaudella astragalensis Rusanova (Fig.16a) Apterae are pale brownish or dirty yellow, BL 1.1-1.5 mm. On Astragalus spp. in Afghanistan, Azerbaijan and Iran. Oviparae and alate males occur in October (Remaudière 1989b).

Bicaudella denaensis (Remaudière) Apterae aresquat-bodied, yellowish with dark dorsal markings and black siphunculi; BL 1.1-1.5 mm. On Astragalus sp. in Iran. Alatae are unknown. Oviparae and alate males occur in September (Remaudière 1989b).

Bicaudella farsiana (Remaudière) (Fig.16b,d) Apterae are yellow, rather pale, BL 1.6-1.9 mm. On an Astragalus sp. of the microthrix group in Iran. Oviparae occur in October (Remaudière 1989b).

Bicaudella laurestanica (Remaudière) (Fig.16c,e) Apterae are yellowish with dusky head and thorax and black dorsal abdominal spots; BL 1.3-1.6 mm. On an Astragalus sp. in Iran. Alatae are unknown. Oviparae and alate males occur in September-October (Remaudière 1989b).

Bicaudella rostrata (Remaudière) Apterae are yellowish, rather pale; BL c.1.2-1.4 mm. On Astracantha gossypina in Iran. Alatae and sexual morphs are unknown (Remaudière 1989b).

Bipersona Hottes

Aphidinae: Macrosiphini

A genus for two nearctic species on Cirsium related to Artemisaphis and Obtusicauda but with a distinctive cauda and anal plate. Miller & Jensen (2006) reviewed the genus, removing one species from synonymy. They are active aphids, and seem invariably to be ant-attended.

Bipersona ochrocentri (Cockerell) Plate 28h Apterae have a pale pink head and thorax, brownish purple abdomen, with last two segments pale green, and black antennae, black legs except for yellow basal parts of femora, black siphunculi and black cauda; BL 2.5-3.5 mm. In ant-attended colonies on leaves and stems of Cirsium spp. (ochrocentrus, arvense) in Arizona and New Mexico, USA. Miller & Jensen (2006) provided a redescription of apterae and alatae. Sexual morphs have not been described.

Bipersona torticauda Gillette (Fig.22e) Apterae are brick red to ochre red with black antennae, black legs except basal parts of femora, black siphunculi, and anal plate and cauda reddish to dusky; BL 2.5-3.8 mm. In ant-attended colonies on leaves and stems of Cirsium spp. (there are records from Carduus and Cnicus, but these are probably referable to this genus). In western USA and Canada. Monoecious holocyclic, with oviparae and apterous males in September-October. Miller & Jensen (2006) provided a redescription of all available morphs.

Blackmania Kanturski & Wieczorek

Aphidinae: Macrosiphini

A genus for one species closely resembling Acaudella in many features such as shape of siphunculi and lack of an evident cauda, but with a much longer processus terminalis.

Blackmania eastopi Kanturski & Wieczorek Apterae are red-brown in life (V.F. Eastop, field notes); BL of aptera c.1.6 mm. Alata has 21-31 secondary rhinaria on ANT III. On Polygonum equisetiforme in Israel and Cyprus (Kanturski & Wieczorek 2015b). The life cycle is unknown.

Boernerina Bramstedt

Calaphidinae: Calaphidini

A genus of 4-5 species associated with Alnus (including Duschekia). The apterae are dorso-ventrally flattened, rather elongate oval, and apply themselves closely to the leaf surface. They have paired anteriorly-directed rectangular projections on the head, lateral abdominal hairs arising from asymmetric tubercles that give the abdominal margin a stepped appearance, and typically 5-segmented antennae. The alatae are more normal-looking Calaphidini. One species (occidentalis) differs in that the apterae often have 6-segmented antennae, the frontal projections are only weakly developed and the abdominal margin is more indented than stepped. Hille Ris Lambers & Hottes (1962) revised the genus.

Boernerina alni Takahashi (fig. 29A,B) Apterae are pale yellowish-white, dorso-ventrally flattened, elongate oval, BL 1.7-2.6 mm. Alatae are whitish and waxy. Sorin (1970a) described the apterous male and ovipara, collected in mid-August in Japan, on Alnus spp. Pashchenko (1983b) described a subspecies (insularia) in east Siberia, which is very likely to be the aphid from the Irkutsk region described by Rupais (1989b) as B. sibirica. This subspecies is also now reported from the Nenets region of north-west Russia (Stekolshchikov 2017), and specimens with characters of both insularia and alni s.str. have been collected in west Siberia (Stekolshchikov & Khureva 2020).

Boernerina depressa Bramstedt Apterae are whitish, flattened and elongate oval, BL 1.5-2.5 mm, on undersides of leaves of Alnus viridis in the Alps and Carpathians, and also reported from A. maximowiczii in the Kuril islands (see Holman 2009). An alata recorded from west Siberia by Ivanoskaya (1977) has been re-identified as B. alni (Stekolshchikov & Khureva 2020). Oviparae and very small apterous males are produced in July-August (Hille Ris Lambers & Hottes 1962).

Boernerina occidentalis Hille Ris Lambers & Hottes Apterae are pale, flattened, oval, BL 1.7-2.4 mm, with broad sclerotic bars on abdominal tergites, the anterior ones being sometimes brownish pigmented. Alatae are variably pigmented, darker specimens have blackish-brown sclerotic bars on abdominal tergites. On Alnus sinuata in Alaska, also collected in Oregon (BMNH collection; leg. D. Hille Ris Lambers), and now also recorded from A. alnobetula ssp. fruticosa (=A. fruticosa) in north-east Siberia (Chukotka; Stekolshchikov & Khruleva 2015). Oviparae and small, slender apterous males are produced in July in Alaska (Hille Ris Lambers & Hottes 1962).

Boernerina variabilis Richards Apterae are pale to bright yellow with black on distal parts of antennae and legs, flattened, elongate oval, BL 1.9-2.6 mm. Alatae vary in pigmentation; when well-pigmented they have broad dark bars dorsally and ventrally on the abdomen. On undersides of leaves of Alnus spp. (sinuata, rugosa, alnobetula) in east and west Canada (British Columbia, Northwest Territory, Ontario, Quebec, Labrador). Oviparae and small, apterous males occur in August in Northwest Territory (BMNH collection, R. O’Doherty), but oviparae were collected in October in northern Ontario (BMNH collection, leg. J. Sÿpkens). 2n=16*.

Brachycaudus van der Goot

Aphidinae: Macrosiphini

This genus of c.50 mainly palaearctic species is characterised by the large round spiracular apertures, short often semicircular cauda, and siphunculi with a subapical annular incision. Brachycaudus is primitively heteroecious with Prunus as primary hosts, and about 14 species still host-alternate. There are species groups associated with secondary hosts in Ranunculaceae (subgenus Brachycaudina), Caryophyllaceae (subgenus Acaudus), Boraginaceae and Compositae (subgenus Prunaphis), and plants formerly in Scrophulariaceae but now placed in Orobanchaceae (subgenus Scrophulaphis). For an introduction to the genus and more detailed accounts of seven of the Prunus-feeding species see Blackman & Eastop 2000, p.246. Heie (1992) reviewed the species in Fennoscandia and Denmark, and Blackman (2010) reviewed the British species (but there are errors in the subgeneric classification in the latter work). Burger (1975) reviewed European species then placed in Acaudus. Andreev (2004) revised the subgeneric classification (but his synonymy of subgenus Thuleaphis with subgenus Mordvilkomemor is not accepted here – see Blackman & Eastop 2011). Cocuzza et al. (2007) made a morphological and molecular study of subgenus Appelia. Coeur d’acier et al. (2008) and Jousselin et al. (2009, 2010) studied the molecular phylogenetics of Brachycaudus species and their bacterial symbiont, which largely confirm the subgeneric groupings used here. Jousselin et al. (2010) analysed their data further to reconstruct ancestral character states, and reached the rather surprising conclusion that several species may have reacquired the primary host after periods of monoecy on secondary hosts. This interesting finding requires further investigation.

Brachycaudus (Thuleaphis) acaudata (Hille Ris Lambers) Colour of apterae in life unknown, probably greenish with extensive blackish dorsal sclerotisation; BL 1.5-1.7 mm. Alatae have 10-14 secondary rhinartia on ANT III. The host of type specimens from Greenland is unknown, but it has subsequently been collected from stems of Persicaria viviparum in arctic Canada (Baffin Island; Richards 1963a). Records of B. acaudata from Rhodiola (= Sedum) rosea in Iceland and Wales are referable to B. sedi. Monoecious holocyclic with apterous males (original description).

Brachycaudus (Brachycaudina) aconiti (Mordvilko) Apterae are shining black, BL 2.0-2.6 mm. Apterae have 0-8 secondary rhinaria on ANT III, and alatae have 7-13. On Aconitum spp. and Delphinium spp. colonising flower stems and leaf petioles, and rarely on undersides of leaves. In central, south and south-east Europe, Central Asia and Siberia. Monoecious holocyclic, with apterous males (Nosyrev 1968). Closely related to B. napelli in western Europe; differing rhinarial numbers on ANT III are often given as a discriminant between the two, but may not always be reliable due to alatiformity.

Brachycaudus (Appelia) almatinus Nevsky Apterae according to the original description are pale brown with black lateral spots, black siphunculi, femora, tibial apices and tarsi, and dusky antennae. BL about 1.8 mm. The cauda is very short and broad. Alata have a pale brown abdomen, presumably without a dark dorsal patch. Found in Kazakhstan on several Prunus spp., rolling leaves longitudinally (Nevsky 1951, Kadyrbekov 2014d). Biology is unknown.

Brachycaudus (Thuleaphis) amygdalinus (Schouteden) Host-alternating between Prunus spp., especially P.dulcis or P. persica, and Polygonaceae. Apterae are green with variably developed dark dorsal markings, squat-bodied with rather short pale legs and antennae (see influential points/gallery); BL 1.6-2.1 mm. Alatae have a dark central abdominal patch and secondary rhinaria distributed III 12-21, IV 0-1 (spring migrants) or III 9-16, IV 0 (gynoparae). Colonies in spring cause rolling of young almond or peach leaves and stunting of new growth. The leaves are rolled somewhat obliquely with respect to the mid-rib. rather than perpendicularly to the mid-rib as in B. helichrysi. Migration occurs to Polygonum (particularly small-leaved) species, Persicaria maculosa and Fagopyrum cymosum. On Polygonum aviculare they feed hidden under the thin bracts surrounding the nodes (Talhouk 1977). It is also frequently anholocyclic on Polygonaceae, even where primary hosts are available. In Europe, the Middle East, North Africa (Tunisia: Boukhris-Bouhachem et al. 2007, Algeria: Laamari et al. 2010), Central Asia, Pakistan and South Africa. Darwish (1983) described the morphology of the spring generations. See also Blackman & Eastop 2000, p.246. 2n=12.

Brachycaudus (Nevskyaphis) ballotae (Passerini) Colour of apterae in life unknown, probably shiny dark green to black; BL c.1.7-1.9 mm. On shoots of Ballota nigra, attended by ants (Szelegiewicz 1981a). In continental Europe (Germany, Czech Republic, Hungary, north Italy and Poland), and also recorded from Iran (Hodjat 1993). Monoecious holocyclic. Molecular data (Jousselin et al. 2010) indicate that B. lamii is probably a synonym.

Brachycaudus (Nevskyaphis) bicolor (Nevsky) (Fig.9b) Apterae are shining yellowish tinged with pink, to pale green, with a large shiny black dorsal abdominal patch (see influential points/gallery); BL c.2.1-2.4 mm. Alatae have secondary rhinaria distributed III 28-52, IV 0-8, V 0. Ant-attended colonies occur on root collars, and at bases of leaves near ground level, of Boraginaceae (e.g. Anchusa, Cerinthe, Cynoglossum, Echium, Lindelofia, Myosotis) and also recorded from a species of Codonocephalum = Inula (Compositae). In UK (Stroyan 1955, Baker 2009), north-west France (Cocuzza et al. 2010), Madeira, Portugal, Spain, Italy, Greece, Egypt, Iran (Rezwani et al. 1994), Caucasus, west Siberia, Tajikistan, Uzbekistan, Kazakhstan, Afghanistan, and alatae have been trapped in Pakistan. Sexual morphs are unrecorded, and it is apparently anholocyclic in Italy (Barbagallo & Patti 1998), but spring colonies have been observed on Amygdalus scoparius in Iran, probably the primary host (Naumann-Etienne & Remaudière 1995).

The disjunct distribution and paler siphunculi of British populations suggest that they may be a distinct taxon, possibly specific to Cynoglossum (Blackman 2010). 2n=12.

Brachycaudus (Thuleaphis) brevirostratus Pashchenko Apterae are dirty green to pale brown; BL c.1.3 mm. On Polygonum spp. in east Siberia. Possibly this is a synonym of B. rumexicolens.

Brachycaudus (Prunaphis) cardui (L.) Plate 10a Apterae on herbaceous plants are shiny black dorsally, light green to yellowish or reddish ventrally (immatures are green or reddish) (see influentialpoints.com/Gallery); BL 1.9-2.3 mm. Alatae have extensive dark dorsal pigmentation, and secondary rhinaria distributed III 12-39, IV 0(-4). B. cardui lives in dense ant-attended colonies on stems and leaves of many species of Compositae (e.g. Arctium, Carduus, Cirsium, Cynara, Chrysanthemum, Tanacetum, Matricaria) and Boraginaceae (e.g Borago, Cynoglossum, Echium, Symphytum), and also frequently on other plants, e.g. Capsella. In Europe, Asia, north Africa and North America. Heteroecious holocyclic in continental climates with a sexual phase on Prunus, esp. P. domestica or P. spinosa (or sometimes P. avium or P. armeniaca). Apterae in rolled leaves of plum in spring, are usually green with the dorsal cuticle sclerotic and often brownish, but not shiny black like the summer generations on secondary hosts. Spring migrant alatae have secondary rhinaria distributed III 20-30, IV 0(-1). According to Börner (1952), the aphids which use P. spinosa as primary host and migrate to basal parts of stems of Compositae are a distinct species, B. lateralis (q.v.), but further work seems necessary to confirm this. Central Asian populations with longer hairs are regarded as a subspecies, B. cardui ssp. turanica Mordvilko (differentiated by Andreev 1999). See also Blackman & Eastop 2000, p.247. 2n=10.

Brachycaudus cerasicola Mordvilko ex Nevsky (= phlomicola Nevsky) Apterae are shiny yellowish brown to dark brown or black due to extensive dorsal sclerotic shield; BL 1.7-2.1 mm. Immatures are dirty brown. Alatae have secondary rhinaria distributed III 21-50, IV 2-23, V 0(-5). Spring colonies roll and twist the leaves of Prunus spp. and turn them red. There is apparently a facultative host alternation to Lamiaceae (Dracocephalum, Eremostachys, Nepeta, Perovskia, Phlomis, Stachys) and Orobanchaceae (Pedicularis) (Shaposhnikov 1964b, as phlomicola; Narzikulov & Daniyarova 1990). Kadyrbekov (2014d) records it also from plant genera in several other families, including Rhodiola (Crassulaceae) and Gentiana/Gentianella (Gentianaceae), where there is possible confusion respectively with B. sedi and B. gentianae. In Central Asia, and there is a redescription from Iran by Sedighi et al. (2018). This species was erroneously synonymised with B. pilosus by Blackman & Eastop (1994).

Brachycaudus (Appelia) cerinthis Bozhko Apterae are shining blackish, BL 1.6-2.1 mm. Alatae have secondary rhinaria distributed III 8-12, IV 0-2, V 0. On stems, flower stalks and bracts, and deforming flower buds, of Cerinthe spp. In France, Italy, Czech Republic, Poland, Hungary and Ukraine. Monoecious holocyclic on Cerinthe (Cocuzza et al. 2007).

Brachycaudus crassitibiae Nevsky Apterae are dark brown with paler appendages; BL c.1.3-1.4 mm. Alatae have 4-6 secondary rhinaria on ANT III. On Turgenia (~Caucalis) latifolia in south Kazakhstan.

Brachycaudus (Acaudus) divaricatae Shaposhnikov Host-alternating between Prunus cerasifera (sometimes P. domestica or P. spinosa) and Caryophyllaceae (usually Silene latifolia s. lat.) in east Europe and the Middle East. Apterae are shining dark brown to black, BL 2.0-2.2 mm, rolling leaves of Prunus in spring. Spring migrant alatae have secondary rhinaria distributed III 16-31, IV 0, but later generations have III 30-50, IV 3-4, V 0. The migration to Silene is apparently facultative (Shaposhnikov 1962). In recent years this species has spread north-westward to the eastern Baltic coast, where it has an abbreviated life cycle, staying on P. cerasifera and producing sexual morphs and eggs from late June (Rakauskas & Juronis 2006, Rakauskas & Turčinavičiene 2006). It has also now reached the Czech Republic, where its morphology and DNA have been compared with eastern Baltic populations (Bašilova et al. 2012). Havelka et al. (2017) have compared its DNA with that of the closely related B. lychnidis. The male genitalia were described and illustrated by Wieczorek et al. (2011). Native predators attacking B. divaricatae in Lithuania were studied by Danilov et al. (2016).

Brachycaudus divaricatellus Shaposhnikov Apterae are shiny yellowish-rose, elongate oval, BL c.2.0 mm. Alatae have secondary rhinaria distributed III 30-50, IV 15-22, V 0. On undersides of leaves of Prunus cerasifera, causing slight curling of tips and discoloration. In south-west Turkmenistan (Shaposhnikov 1964a). Life cycle is unknown. Shaposhnikov (1964b) described the ovipara.

Brachycaudus (Thuleaphis) eurotiae (Mamontova-Solukha) Apterae are shiny yellow, without wax; BL 1.1-1.6 mm. On Krascheninnikovia spp., living in galls formed by curling and thickening of the upper sides of the leaves. Alatae have 5-7 secondary rhinaria on ANT III only, and dark dorsal markings. In Ukraine (Mamontova-Solukha, 1968, as Cryptosiphon eurotiae), Kazakhstan and western China (Kadyrbekov et al., 2002). Kadyrbekov (2002) erected a new genus (Scythaphis) for this species which has an unusually setose abdominal tergite 8 and lacks marginal tubercles (and has a somewhat distinctive biology), but Stekolshchikov (2007) transferred it to Brachycaudus (Thuleaphis).

Brachycaudus gentianae Daniyarova Apterae are shining brown; BL c. 1.8 mm. On Gentiana olgae in Tajikistan, and on Gentiana tianshanica in Kazakhstan (Kadyrbekov 2017a). Possibly this is a synonym of B. cerasicola, which is also recorded from Gentiana/Gentianella by Kadyrbekov (2014d).

Brachycaudus helichrysi (Kaltenbach) Plate 9i Apterae are very variably coloured (but always pale); pale green, pale yellow, whitish or pinkish (see influentialpoints.com/Gallery); BL 0.9-2.0 mm. Alatae have secondary rhinaria distributed III 13-46, IV 0-18. On stems and in flowerheads of numerous plant species, esp. Compositae/Asteraceae (e.g. Achillea, Ageratum, Aster, Bidens, Chrysanthemum, Cineraria, Erigeron, Gnaphalium, Helianthus, Matricaria, Senecio) and Boraginaceae (e.g. Anchusa, Cynoglossum, Myosotis, Symphytum), but also many others including Rumex, Saxifraga, Trifolium, Veronica. World-wide, and a major pest (for further information see Eastop & Blackman 2000, p.247). Heteroecious holocyclic with sexual phase on Prunus spp. (especially P. domestica, P. insititia, P. spinosa) in colder climates. Apterae in spring populations on Prunus are green, brownish or yellowish, shiny and slightly waxy, causing leaves to roll up tightly perpendicular to the mid-rib. BL is 1.4-2.0 mm on Prunus, down to 0.9 mm on secondary hosts. Darwish (1984) described the morphology of spring generations.

Anholocyclic populations occur in warmer regions and in glasshouses, and are sometimes found on new growth of various trees. Molecular studies have now revealed that populations identified as B. helichrysi throughout the world comprise two forms that are sufficiently distinct to be regarded as sibling species, genetically isolated from each other (Pifaretti et al. 2012, 2013a,b and possibly Rebijith et al. 2013). Host alternation to Prunus domestica and related species only occurs in one of these forms (H1), in regions with a continental climate. The second form (H2) is mainly anholocyclic, and is mostly absent from the coldest regions, but has now been shown to have a sexual phase on Prunus persica in northern India (Pifaretti 2013b). Mixed populations of the two forms often occur on herbaceous hosts, and no clear differences in host preference on herbaceous plants between these two forms have so far been detected. Further studies have revealed a third form (H3) restricted to East Asia, associated with apricots and Prunus species endemic to that region (Popkin et al. 2017). 2n=12.

Brachycaudus (Prunaphis) iranicus Davatchi & Remaudière (Fig.9a) Apterae are greenish yellow to pale green with shiny brown dorsal markings (a transverse bar on the mesonotum, a central abdominal patch and presiphuncular sclerites) and brown siphunculi; BL 1.6-2.0 mm. On undersides of leaves of Anchusa azurea (= italica) and A. strigosa, feeding close to the main veins, in the Middle East (Iran, Turkey, Israel, Lebanon and Syria). The alata (hitherto undescribed) has dark antennae, 13-16 secondary rhinaria on ANT III, and a central abdominal sclerite on abdominal tergites 5-6, fused each side with presiphuncular sclerites, and with a large central lacuna. Monoecious holocyclic with alate males (Davatchi & Remaudière 1953).

Brachycaudus (Prunaphis) jacobi Stroyan Apterae are black and shiny dorsally, olive green ventrally; BL 1.4-2.0 mm. Alatae have secondary rhinaria distributed III 6-16, IV 0-2. At base of stem or on roots of Myosotis and Pulmonaria spp., ant-attended. In Europe (UK, Netherlands, Germany, Italy). Monoecious holocyclic with apterous males (Müller 1975a). 2n=12.

Brachycaudus (Acaudus) klugkisti (Börner) Apterae are shining black dorsally, red-brown ventrally; BL 1.5-2.5 mm. Alatae have secondary rhinaria distributed III 11-31 (in a row), IV 0-8. On upper parts of Silene (incl. Melandrium) spp. In Europe (UK, Scandinavia, Finland, Germany, Poland, France, Italy). Monoecious holocyclic with apterous males (Müller 1987). 2n=10.

Brachycaudus (Nevskyaphis) lamii Koch Apterae have a shining black dorsum; BL 1.8-2.2 mm. On Lamium spp., especially L. album, in Europe (Germany, Poland, Switzerland, Italy, Slovakia). Monoecious holocyclic, with oviparae in October (BMNH collection, leg. D. Hille Ris Lambers). Molecular data (Jousselin et al. 2010) indicate that this is probably a synonym of B. ballotae.

Brachycaudus (Prunaphis) lateralis (Walker) Apterae with a shining black dorsum, ventrally green or reddish; BL 1.6-2.6 mm. On stems and leaves, usually close to ground, of plants in numerous genera of Compositae including Arctium, Anthemis, Carduus, Cirsium, Chrysanthemum, Galinsoga, Leucanthemum, Matricaria and Senecio. Also found on Capsella. In Europe, eastward to Ukraine. Mainly anholocyclic, although a sexual phase may sometimes occur on Prunus (Müller & Horatschek 1979). It is closely related to B. cardui, often treated as a subspecies, and DNA studies (Jousselin et al. 2009, 2010) indicate that it may be a variant form of that species.

Brachycaudus (Scrophulaphis) linariae Stroyan Apterae are deep blackish green, shiny, immatures being paler green (see influential points/gallery); BL 1.4-1.9 mm. Alatae have secondary rhinaria distributed III 11-37, IV 5-10, V 0-2. On basal parts of Linaria spp., attended by ants. In UK, Denmark, Sweden, Finland, France, Germany, Italy and most of eastern Europe (Andreev & Mamontova 1998). Monoecious holocyclic; Patti & Barbagallo (1997) described oviparae and apterous males collected from L. purpurea in Sicily in October.

Brachycaudus (Nevskyaphis) lucifugus Müller Apterae are yellowish green with shiny dark brown to black dorsum; BL 1.3-2.2 mm. Alatae have secondary rhinaria distributed III 15-30, IV 0-9, V 0-1(-5). On roots and at leaf bases of Plantago lanceolata, in Europe (UK, Germany, Hungary, Italy). Ant-attended. Monoecious holocyclic with apterous males (original description and Stroyan 1964a).

Brachycaudus (Acaudus) lychnicola Hille Ris Lambers Apterae are reddish brown with shining black dorsal shield; BL 1.9-2.3 mm. Alatae have secondary rhinaria distributed III (15-) 23-36, IV 0-4. On roots, stem-bases and lower leaves of Lychnis flos-cuculi and Silene (incl. Melandrium) spp., attended by ants. In north-west Europe (Netherlands, Sweden). Monoecious holocyclic with apterous males (original description). Molecular data (Jousselin et al. 2010) indicate that, if the specimens from Czech Republic used in their analysis are correctly identified as this species, then it is probably a synonym of B. lychnidis.

Brachycaudus (Acaudus) lychnidis (L.) Apterae are reddish brown with shining brown-black dorsal shield (see influentialpoints.com/Gallery); BL 1.8-2.9 mm. Alatae have secondary rhinaria distributed III 15-34, IV 0-4. On stems, leaves and flowers of Lychnis and Silene (incl. Melandrium) spp. Europe, eastward to west Siberia, Turkey, Iran, Caucasus and Kazakhstan. Monoecious holocyclic with apterous males (Heie 1992). 2n=12.

Brachycaudus (Nevskyaphis) malvae Shaposhnikov Apterae are shining blackish green (see influential points/gallery); BL 1.8-2.3 mm. Alatae have secondary rhinaria distributed III 17-31, IV 0-3, V 0. On Malva spp., in ant-attended colonies at base of stem and on lower leaves. In England, Spain, southern Russia, Ukraine, and also China, if Brachycaudus atuberculatus Zhang 1981 is a synonym (see Andreev 2004). 2n=12.

Brachycaudus (Scrophulaphis) mimeuri Remaudière Apterae are shining pale to dark brown; BL 0.7-1.4 mm. Alatae have secondary rhinaria distributed III 12-33, IV 1-10, V 0-1. On roots of Euphrasia and Odontites spp. in southern and eastern Europe; for distribution see Jörg & Lampel 1988, but also in Denmark (Heie 1992) and Sicily (Barbagallo & Stroyan 1982), and possibly also in South Australia on Parentucellia latifolia (Eastop 1966, as ?B. persicaecola). Not attended by ants (Hille Ris Lambers 1967). Presumably anholocyclic, and closely related to B. persicae (q.v.).

Brachycaudus (Prunaphis) mordvilkoi Hille Ris Lambers Apterae are green with shiny black dorsal shield; BL 1.7-2.3 mm. Alatae have secondary rhinaria distributed III 12-21, IV 3-6, V 0-1. On certain Boraginaceae (Anchusa, Echium, Solenanthus, Symphytum), with one record from Compositae (Hieracium pilosella). Widely distributed in Europe (but not recorded from British Isles, Scandinavia or Iberian peninsula) and in Middle East (Turkey; Şenol et al. 2017). Monoecious holocyclic with apterous males (Müller 1975a).

Brachycaudus (Brachycaudina) napelli (Schrank) (Fig.6b,c) Apterae are shining black with end of abdomen red, immatures are brown; BL 2.0-2.9 mm. Alatiform apterae occur with 3-25 secondary rhinaria on ANT III, alatae have 17-37 on III and 0-4 on IV. On Aconitum spp., esp. napellus, forming colonies on stems, flowers and fruits, and sometimes on leaves. Only occasionally on Delphinium spp. In west and north-west Europe (not UK), south to Spain, and also recorded from Austria and Czech Republic (Cocuzza et al. 2010). Closely related to B. aconiti (q.v.). Monoecious holocyclic with apterous males.

Brachycaudus (Acaudus) pallidus Andreev Apterae are shining dark brown; BL 1.5-2.2 mm. On roots of Silene nutans in Moldova.

Brachycaudus (Scrophulaphis) persicae group Apterae have a shiny dark brown or black dorsum, and black siphunculi (see influential points/gallery); BL 1.5-2.2 mm. Alate spring migrants have secondary rhinaria distributed III 23-51, IV 9-21, V 1-6. B. persicae (Passerini) s.str. is most commonly found in large spring colonies on young stems of Prunus persica or P. armeniaca, often persisting into the summer on root suckers. Alatae from Prunus have been transferred successfully to Euphrasia sp. and Rhinanthus glaber and subsequently to Melampyrum pratense (Burger 1975), so it seems probable that there is a facultative host alternation from Prunus to Orobanchaceae (Euphrasia, Melampyrum, Rhinanthus), at least in Europe, but anholocyclic overwintering is also common on Prunus roots.

On Orobanchaceae it lives on the above-ground parts (Heie 1992). Aphids commonly regarded as this species are also widely distributed on peach outside Europe, in the Middle East, Central Asia, southern Africa, Australia, New Zealand, North and South America. Similar aphids also occur commonly in Europe on P. domestica and P. spinosa, and occasionally on Pyrus communis and Chaenomeles japonica. These all form a complex of which the host relationships and life cycles need further clarification. Müller & Steiner (1988a) applied the name B. semisubterraneus Börner to anholocyclic populations on plum in Germany, and suggested that the form distributed world-wide on peach was also an anholocyclic species, to which they applied the name B. persicaecola (Boisduval). However, aphids of the B. persicae group are recorded from Orobanchaceae in both Australia and Chile (BMNH collection), indicating that some host alternation occurs in southern continents, and Burger (1975) showed that not all populations on plum and peach are anholocyclic, so the application of the names semisubterraneus and persicaecola is still unclear. There are also records from Odontites, but these may be referable to B. mimeuri (q.v.), a closely-related anholocyclic form on roots of Orobanchaceae that has also become widely distributed. 2n=10.

Brachycaudus (Mordvilkomemor) pilosus (Mordvilko ex Nevsky) Apterae are black with pale appendages and cauda; BL 1.1-1.7 mm. Alatae have secondary rhinaria distributed III 16-23, IV 6-8, V 0-1. On various Prunus spp., especially P. ulmifolia (= P. triloba), damaged leaves of which develop gall-like thickenings and red spots, and P. bucharica, the leaves of which are only slightly curled and reddened (Narzikulov 1965c). Possibly there is no host alternation; populations on P. prostrata in Pakistan included apterous males and oviparae in May together with fundatrices, and apterous and alate viviparae, and additional sexual morphs were collected in August (Naumann-Etienne & Remaudière 1995). In mountainous regions of central Asia (Kazakhstan, Kirgizia, Tajikistan, Turkmenistan), Iran, India and Pakistan (BMNH collection).

Brachycaudus (Nevskyaphis) plantaginis Holman & Szelegiewicz Apterae are yellowish brown with shiny dark dorsum; BL c.1.9-2.1 mm. Alatae are undescribed. Ant-attended, on root collar of Plantago depressa and P. major in Mongolia and east Siberia (Pashchenko 1988a).

Brachycaudus (Acaudus) populi (del Guercio) Apterae are red-brown with shiny black dorsal shield; BL 1.6-2.9 mm. Alatae have secondary rhinaria distributed III 19-36, IV 0-3. On upper parts of Silene spp., widely distributed in Europe, eastward to Crimea, Russia (Asltai) and Kazakhstan. Monoecious holocyclic with apterous and alate males. Closely related to B. lychnidis, and confused with that species prior to Burger (1975). 2n=12.

Brachycaudus (Appelia) prunicola (Kaltenbach) Apterae are shiny black, dark green or deep yellow-green with black dorsal cross-bands, immatures grey-green (see influential points/gallery); BL 1.4-2.4 mm. Alatae have secondary rhinaria distributed III 23-43, IV 7-16, V 0-3. In spring on new growth of Prunus (in Europe usually P. spinosa, sometimes P. domestica or P. insititia), causing severe leaf curl and discoloration. Host-alternating between Prunus and Tragopogon, but part of the population – possibly a subspecies – may live without alternation on P. spinosa. In continental Europe, Siberia (Stekolshchikov et al. 2008) east to Irkutsk (Stekolshchikov & Shaposhnikov 1998), Iran (Rezwani & Radjabi 1986), Kazakhstan (Kadyrbekov 2005c) and Pakistan (Baluchistan; Naumann-Etienne & Remaudière 1995). [Note: the sample (no. 992) identified as B. prunicola from Brittany used in DNA analysis by Jousselin et al. (2010), and branching separately from the rest of the prunicola/schwartzi/tragopogonis group in their phylogeny, was B. prunifex (q.v).] 2n=12.

Brachycaudus (Appelia) prunifex (Theobald) Apterae are shiny dark green to black (see influential points/gallery); BL 1.5-2.4 mm. Alatae have secondary rhinaria distributed III 22-41, IV 2-13, V 0(-2). Colonies occur all-year-round on Prunus spinosa, in spring causing severe leaf curl and discoloration to new growth. There is also a record from P. cerasifera var. atropurpurea. Suggestions of a facultative host alternation to Tragopogon do not appear to be substantiated. Alate males have been collected in September (BMNH collection), but oviparae are not yet recorded. In England, Wales, Ireland, and northern France. This species has been synonymised previously with B. prunicola (Kaltenbach), which occurs on the same host throughout the rest of Europe and in central Asia, but it can be distinguished from that species (and from B. schwartzi) by the longer hairs on the antennae and anterior abdominal tergites (Blackman 2010). 2n=12.

Brachycaudus (Acaudus) rinariatus Andreev Apterae have a shining black dorsal shield; BL 1.7-2.2 mm. Alatae have secondary rhinaria distributed III 36-48, IV 16-19, V 1-6. On Linaria vulgaris in eastern Europe (Russia, Moldova, Ukraine), and on L.× incompleta in Kazakhstan (Kadyrbekov, 2017a, as B. himalayensis).

Brachycaudus (Brachycaudina) rociadae (Cockerell) (Fig.6a) Apterae are shining reddish brown to black, fading to hazel brown anteriorly and posteriorly; BL c. 2.0 mm. Immatures are bright coral red. Alatae have 2-13 secondary rhinaria on ANT III only. On stems and leaves of Delphinium spp.; feeding on leaves causes them to curl tightly (Palmer 1952) . Widely distributed in North America. Monoecious holocyclic, with oviparae and apterous males in mid-July (Ontario; BMNH collection, leg. S.F. MacDonald) or early October (Colorado and Manitoba).

Brachycaudus (Thuleaphis) rumexicolens (Patch) Apterae are reddish with variably developed dark dorsal markings; BL 1.3-2.1 mm. Alatae have more-or-less fused cross-bands on posterior abdominal tergites, 6-22 secondary rhinaria on ANT III, and 0(-1) on IV. In inflorescences of Rumex spp., especially acetosella, and sometimes on other Polygonaceae (Fagopyrum, Persicaria, Polygonum. Rheum). In Europe, north Africa (Tunisia: Boukhris-Bouhachem et al. 2007), west and Central Asia, north-east Siberia (Chukotka: Stekolshchikov & Khruleva 2015), India, Australia, North and South America. Monoecious holocyclic, with apterous males (Tuatay & Remaudière 1964). B. brevirostratus in east Siberia might be this species. 2n=12.

Brachycaudus salicinae Börner Apterae are shining grey-black; BL 1.4-1.9 mm. Alatae have secondary rhinaria distributed III 16-38, IV 3-13, V 0. In rolled leaves of Inula spp., especially salicina, and also Pulicaria dysenterica (Holman 2009); in Europe (Germany, France, Poland, Czech Republic, Slovakia), Iran (Rezwani & Parvizi 1990) and Central Asia (Kazakhstan; Kadyrbekov & Aoitzhanova 2005). Monoecious holocyclic (Börner 1952).

Brachycaudus (Appelia) schwartzi (Börner) Apterae of BL 1.4-2.1 mm, shiny yellow-brown to dark brown with extensive dorsal black sclerotization; immatures are yellow-brown. Alatae have secondary rhinaria distributed III 23-36, IV 6-17, V 0(-1). Living without host alternation on Prunus persica; spring colonies cause severe curling and distortion of peach leaves. In Europe, Iran, Kazakhstan, North Africa (Tunisia, Egypt), India, South America and California. Records from other Prunus spp. with the possible exception of P. serotina, may be referable to other species. Records of B. tragopogonis from peach in the literature are referable to this aphid. Darwish (1983) gave descriptions of all developmental stages. A morphometric and molecular study by Cocuzza et al. (2007) indicated that this species should perhaps be treated as a subspecies of B. prunicola. (See also Blackman & Eastop 2000, p.249). 2n=12.

Brachycaudus (Thuleaphis) sedi (Jacob) Apterae are yellowish green, with dark cross-bands on posterior abdomen; B: 1.0-2.1 mm. Alatae have secondary rhinaria distributed III 11-19, IV 0-5, V 0(-1), VI base 0(-1). On stem apices and flowers of Rhodiola (= Sedum) rosea, causing distortion. There is also a record from Sedum anacampseros (Cocuzza et al. 2010). In Europe (including Iceland, as Thuleaphis acaudata; Prior & Stroyan 1960) and west Siberia (Ivanoskaya 1975, as Rhodiolaphis cholsunensis). Monoecious holocyclic, with oviparae and apterous males appearing in August (original description). 2n=8*.

Brachycaudus shaposhnikovi Narzikulov Apterae are shining yellowish green with black dorsal sclerotic markings, siphunculi pale with tips darker; BL 2.1-2.7 mm. Alatae have 15-18 secondary rhinaria on ANT III. Monoecious holocyclic on Atraphaxis spp. in Tajikistan (Narzikulov & Daniyarova 1990) and Kazakhstan (Kadyrbekov 2014e).

Brachycaudus spiraeae Börner Apterae are light green to greyish brown with variable dark dorsal markings; BL 1.2-1.8 mm. Apterae have secondary rhinaria distributed III 0-11, IV 0-1, and alatae have them distributed III 19-34, IV 4-13, V 0-2. On Spiraea spp., in leaves rolled and curled into narrow “pods”. In Europe, west Siberia, Iran (Rezwani 2010) and Central Asia. Generally regarded as monoecious holocyclic on Spiraea, but molecular work has revealed a close relationship with H3 populations of B. helichrysi, possibly involving introgression, and implying some degree of migration from Spiraea to herbaceous hosts (Popkin et al. 2017). 2n=12.

Brachycaudus (Appelia) tragopogonis (Kaltenbach) Apterae are shining grey-brown to dark brown with dark dorsal sclerotic markings (see influential points/gallery); BL 1.4 –2.3 mm. Alatae have secondary rhinaria distributed III 27-49, IV 6-22, V 0-4. On aerial parts of Tragopogon spp. and Scorzonera spp. in Europe, west Siberia, south-west and Central Asia (Kazakhstan; Kadyrbekov 2005c, 2009a), and introduced to western USA and South America (Brazil, Chile). Monoecious holocyclic. In western Europe there is possible confusion with the morphologically similar B. prunicola which has a sexual phase on Prunus spinosus. Hille Ris Lambers (1948) described a subspecies with long dorsal hairs, B. tragopogonis ssp. setosus, on T. longirostris in Israel, and collections from south-west and Central Asia and Pakistan may be of this form (Mostafawy 1967), which is also reported from Hungary (Basky 2015). 2n=12 (2n=11 and 12 for samples of ssp. setosus from Israel and Iran respectively).

Brachycaudus umbelliferarum Nevsky Apterae are pale brown or yellowish green with dark brown pleural and marginal spots, or entirely dark brown; BL c. 2.4 mm. On roots of an umbelliferous plant, probably Heracleum sp. south-east Kazakhstan. With marginal tubercles on prothorax, abdominal tergites 1, 2 and 7, this species is unlikely to be a Brachycaudus, and is regarded by Andreev (2004) as a nomen dubium.

Brachycaudus (Nevskyaphis) virgatus Shaposhnikov Colour of apterae in life is unknown, probably greenish with darker head and dorsal markings; BL 1.6-1.8 mm (2 specimens). On stems and leaves near base of host plants, which are Boraginaceae (Anchusa, Symphytum) and the composite Senecio fluviatilis (= nemorensis). Alatae have not been described. Only known from southern Ukraine.

Brachycaudus viridanus (Nevsky) Apterae are pale green, BL 1.3-1.7 mm. On stems of Cousinia sp. in Uzbekistan. Alatae are also pale (brownish head and thorax and pale green abdomen), and have about 7 rhinaria on ANT III. This species was synonymised by with B. cardui (presumably ssp. turanica), and could possibly be a dwarf form of that species lacking dark sclerotisation.

Brachycolus Buckton

Aphidinae: Macrosiphini

Four or five species with very short broad-based conical or barrel-shaped flangeless siphunculi and triangular cauda associated with Caryophyllaceae. Heie (1992) and Blackman (2010) reviewed the species in western Europe.

Brachycolus aestivus Zhang, Qiao & Zhang Colour of apterae in life unrecorded; BL c.1.5 mm. On Bupleurum chinense in Shanxi province, China (L. Zhang et al. 2008). To judge from the host association this species might belong in Semiaphis.

Brachycolus brachysiphon Richards Colour of apterae in life unobserved; BL c. 1.8 mm. On Stellaria longipes in northern Canada.

Brachycolus cerastii (Kaltenbach) (Fig.51b) Apterae are dirty green, wax-powdered; BL 1.3-1.7 mm. Alatae have 3-6 secondary rhinaria on ANT III. On Cerastium spp., esp. C. arvense, causing shoots to be stunted and deformed into gall-like structures. Apparently widely distributed in Europe, but rarely collected. Also recorded from Kazakhstan on C. davuricum and C. tianschanicum (Kadyrbekov 2014e, 2017a). Monoecious holocyclic (but sexual morphs are apparently undescribed). 2n=14.

Brachycolus cucubali (Passerini) Plate 13d Apterae are whitish yellow to pale green, wax-powdered; BL 1.4-2.2 mm. Alatae have secondary rhinaria distributed III 9-16, IV 0-2. On Silene spp., rolling leaves into elongate pseudogalls (Heie 1992, as Hayhurstia). Widely distributed in Europe, in eastern Siberia and Kazakhstan and in north Africa. In Kazakhstan it is also recorded from Oberna and Cerastium (Kadyrbekov 2017a). Monoecious holocyclic; Casiraghi et al. (2020) studied the life cycle on Silene vulgaris in northern Spain, redescribed the aptera and alata, and described oviparae and apterous males found in the pseudogalls in November to January. Apterous viviparae also live through the winter in the galls along with the overwintering eggs.

Brachycolus stellariae (Hardy) Apterae are pale green, wax-powdered; BL 1.1-2.0 mm. Alatae have secondary rhinaria distributed III 4-9, V 0-1. On upper sides of leaves of Stellaria spp., which are rolled into oblong pseudogalls. Also recorded from Moehringia trinervia. In Europe. Monoecious holocyclic with oviparae and very small apterous males produced in September (Blackman 2010).

Brachycorynella Aizenberg

Aphidinae: Macrosiphini

Two palaearctic species with different biologies, similar to Brachycolus but with shorter antennae and terminal processes, and very short conical siphunculi.

Brachycorynella asparagi (Mordvilko) Apterae are green, covered with grey mealy wax; BL 1.2-1.8 mm. On Asparagus spp., causing stunting of shoots and “rosetting” of leaves, which turn blue-green. In Europe (not yet in UK, but recently expanding its range), North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), south-west and Central Asia (Kazakhstan; Kadyrbekov & Aoitzhanova 2005), China, and introduced into North America (see also Blackman & Eastop 2000). Monoecious holocyclic with alate males. 2n=10.

Brachycorynella lonicerina (Shaposhnikov) (Fig.38c) Apterae are green, dusted with greyish wax powder, BL c.1.8 mm. On Lonicera spp., feeding on upper sides of leaves, which are folded in half (Shaposhnikov, 1964, as Semiaphis). In Central Asia, southern Russia, Ukraine, Iran and Pakistan (Naumann-Etienne & Remaudière 1995). Heteroecious holocyclic but with with an unknown secondary host (Remaudière & Remaudière 1997, p. 298). Oviparae and alate males appear on Lonicera in late September-October (Narzikulov 1965b; Mukhamediev & Akhmedov 1982). 2n=10*.

Brachymyzus A.N. Basu

Aphidinae: Macrosiphini

One species in India with a Myzus-like head but with thick truncate siphunculi and an unusual cauda.

Brachymyzus jasmini A.N. Basu Apterae have head, thorax and antennae blackish brown, abdomen greenish, BL 1.6-2.1 mm. On leaves and young shoots of Jasminum humile in north-east India (West Bengal). Oviparae were described from Sikkim on Nellia (Neillia?) sp. (Rosaceae), and an alate male was found on Pilea microphylla (Mondal et al. 1978). There is possibly host alternation between Neillia and Jasminum.

Brachysiphoniella Takahashi

Aphidinae: Macrosiphini

One or two species of grass-feeding aphids in east and south-east Asia, with distinctive very short black siphunculi and a long, dark, rather hairy cauda.

Brachysiphoniella apiaca Zhang & Zhang Colour of apterae in life is unrecorded; BL c.1.6 mm. On Apium graveolens in China (L. Zhang & G. Zhang 2000a). The host needs confirmation, as this aphid is very similar to the grass-feeding B. montana.

Brachysiphoniella montana (van der Goot) Plate 14d Apterae are brownish green, densely covered in mealy wax (see aphids of Karnataka website); BL 1.1-1.9 mm. On grasses, well-adapted to aquatic species such as Leersia hexandra due to its waxy coat. Also recorded from Cynodon, Eleusine, Microstegium, Miscanthus, Oryza (as Melanaphis vandergooti Raychaudhuri & Banerjee), Panicum, Pennisetum and Phragmites. India, Bangladesh, Nepal, China, Taiwan, Vietnam, Korea, Japan, Philippines, Malysia, Indonesia, and Australia (NSW). Presumably mainly anholocyclic, but at least in Korea there may be host alternation to Pyrus spp. (Lee et al. 2002c).

Brachyunguis Das

Aphidinae: Aphidini

About 40 species mainly on xerophytic plants, mostly Amaranthaceae. Related to Aphis but with a very short antennal terminal process (usually shorter than in Aphis subg. Protaphis) and very short, often pale siphunculi. The clypeus is often globose, but the extent of its enlargement varies considerably, and subgeneric separation of Xerophilaphis on grounds of greater enlargement of clypeus (Remaudiere & Halbert 1996) does not seem warranted. Many palaearctic species and one described from South America are coated in mealy wax in life, but wax production is not reported for North American species. Kadyrbekov (1999c) reviewed and keyed the species in Central Asia, Kadyrbekov (2001c) summarised the characters of the genus and transferred several species to it, and Kadyrbekov (2014c) described several more species from Kazakhstan. Nieto Nafría et al. (2005a) reviewed the two species on the Iberian peninsula and provided lists of synonymies. Despite this the genus is still poorly known; many of the nominal species are very similar to one another and their separate identity needs experimental confirmation.

Brachyunguis agriphylli Bozhko Apterae are green, wax-powdered, with dark green pleural markings; BL c.1.5-1.6 mm. In large colonies on Agriophyllum squarrosum (= A. arenarium) in Ukraine. Not clearly distinct from B. harmalae.

Brachyunguis armelliae Kadyrbekov Apterae are greenish with brown head, without wax powdering; BL 1.5-1.7 mm. The single known alata has 7-8 secondary rhinaria on ANT III. On inflorescences of Nitraria schoberi, ant-attended, in south-east Kazakhstan (Kadyrbekov 2014c).

Brachyunguis atraphaxidis (Nevsky) Apterae are dark green, with paler appendages, body wax-dusted; BL c.1.6-1.7 mm. In small colonies on leaves and flowers of Atraphaxis spp., attended by ants. Recorded from Iran, Georgia, Turkmenistan, Uzbekistan, Tajikistan, Kyrgyzstan, and Kazakhstan (Kadyrbekov 2017a).

Brachyunguis bahamondesi Remaudière & Halbert Apterae are pale, with apices of tibiae, tarsi and siphunculi black, body colour in life unrecorded, BL 1.0-2.2 mm. On Amaranthaceae (Chenopodium, Atriplex) in Argentina.

Brachyunguis berezhkovi (Ivanoskaya) Apterae are pale green, pruinose, with pale appendages; BL c.1.4-1.6 mm. In small ant-attended colonies at shoot terminals of Salsola richteri in Central Asia (Turkmenistan).

Brachyunguis bicolor Ivanoskaya Apterae are grey with darker patches laterally, wax-dusted; BL c. 1.3 mm. On Cynanchum acutum, singly or in small colonies on stems, flowers and flower stalks, attended by ants. In Ukraine (as B. cynanchiacuti Bozhko), Turkmenistan and Kazakhstan.

Brachyunguis bishopi Remaudière & Halbert Apterae are pale, colour in life unrecorded, BL 1.2-2.3 mm. On terminal parts of Sarcobatus vermiculatus in western USA. Oviparae and apterous males occur in October (original description).

Brachyunguis blanchardi Remaudière & Bahamondes Apterae are dark grey, wax-dusted, BL 1.6-2.0 mm. On Chenopodium (=Allenrolfea) vaginata in Argentina. The clypeus is greatly enlarged in this species, indicating specialisation on Amaranthaceae.

Brachyunguis bonnevillensis Knowlton Apterae are pale bluish green, with pale appendages (see aphidtrek.org); BL 1.0-2.1 mm. Alatae have a shiny black head and thorax. In small colonies on leaves, flower stems and new shoots of Sarcobatus vermiculatus. It is well camouflaged on lower surfaces of leaves, and has a dwarf form in summer (Palmer 1952). Also recorded from Atriplex canescens (Remaudière & Halbert 1996), although there is possible confusion with B. tetrapteralis (q.v.). In western USA and Mexico.

Brachyunguis brevisiphon Kadyrbekov, Renxin & Shao Apterae are greenish with a slight grey wax film; BL c. 1.2 mm. In ant-attended colonies on shoots of Tamarix spp. in China (Xinjiang-Uygur region; Kadyrbekov et al. 2002), and subsequently recorded from Kazakhstan (Kadyrbekov 2017a). The description does not provide any clear differences from B. tamaricophila, and the key character that we have used to distinguish this species may not be reliable.

Brachyunguis cahuille (Dickson) (Fig.36g) Apterae are dull dark green, BL 0.9-1.4 mm. On leaves of Suaeda spp. in southern California, Arizona and Mexico (Remaudière & Halbert 1996; as B. tetrapteralis).

Brachysiphon calligoni (Nevsky) Apterae are pale green, covered in wax meal; BL 1.3-1.7 mm. On young shoots of Calligonum spp. in Uzbekistan and Kazakhstan (Kadyrbekov 2017a).

Brachyunguis calotropicus Menon & Pawar Apterae are pale to muddy green, with sparse wax powder, BL 1.2-1.5 mm. In dense aggregations on upper or undersides of younger leaves of Calotropis spp. (gigantea, procera), sometimes attended by ants. The life cycle is unknown. First recorded from the Delhi region of India without a proper description (Menon & Pawar 1958). B. ushari Müller & El Tigani 1986 seems to be a synonym, and the present account is based on that assumption. There is also a possibility that B. calotropicus may prove to be a synonym of B. harmalae. In India, south-west Asia (Saudi Arabia) and Africa (Algeria, Egypt, Sudan). 2n=8 (Kurl 1978).

Brachyunguis convolvulisucta Zhang, Chen, Zhong & Li Appearance in life is unrecorded; BL c.1.5-1.6 mm. On Convolvulus arvensis in Xinjiang province, China (G. Zhang 1999).

Brachyunguis cuscutae (Nevsky) Apterae are green, densely covered with wax; BL 1.2-1.5 mm. Found in immense numbers on flowers and stalks of Cuscuta spp. in early November (Ivanoskaya 1960). In Uzbekistan, and also recorded from Iran (Rezwani et al. 1994) and Kazakhstan (Kadyrbekov 2014e).

Brachyunguis cynanchi (Nevsky) Apterae are dull red to dark violaceous, coated with fine mealy wax; BL 1.2-1.8 mm. On Cynanchum acutum in Uzbekistan, found in large numbers on apical parts of stems, flower stalks and upper sides of leaves (Ivanoskaya 1960). Also recorded from Iran (Rezwani 2010) and Kazakhstan (from C. sibiricum; Kadyrbekov & Aoitzhanova 2005).

Brachyunguis dendrostellerae Kadyrbekov Apterae are greenish without wax powdering, with siphunculi variably pigmented and dark reddish eye: BL 0.9–1.8 mm. On flowers and stems of Diarthron arenarium and D. ammodendron (= D. stachyoides) in south-east Kazakhstan, and of D. lessertii in Iran, ant-attended (Kadyrbekov 2014c, Barjadze et al. 2022).

Brachyunguis flexosiphon Kadyrbekov Apterae are dark green with red eyes; BL 1.5-1.8 mm. Described from the inflorescence of an unidentified umbelliferous plant (Kadyrbekov 1999c), and subsequently recorded from Zosima tordyloides (= Semenovia rubtzovii), Ferula spp., Schrenkia ?golickeana and Bunium setosum (= setaceum?) (Kadyrbekov 2014d, 2017a). In Kazakhstan.

Brachyunguis harmalae Das (= plotnikovi Nevsky) Plate 7h, fig.36h Apterae are mid- to deep green anteriorly, more yellowish on posterior abdomen, thickly dusted with wax powder, and with two longitudinal rows of submarginal dark green spots; BL 1.2-2.0 mm. Alatae have 3-7 secondary rhinaria on ANT III. Often ant-attended. Monoecious holocyclic on Peganum harmala in Pakistan, where Das (1918) studied its life cycle in detail, and on Calligonum in Central Asia, where damaging populations have been reported from cotton (Nevsky 1928a, as plotnikovi). Both apterous and alate males occur. The populations that occur in the Middle East, Africa (Tunisia, Eritrea, Kenya, Sudan, Senegal) and on the Iberian peninsula (Nieto Nafría et al. 2005a), colonising various, mostly xerophytic plants including Calligonum and Calotropis, are presumably anholocyclic. Eastop & Raccah (1988) listed 12 hosts in 7 botanical families in the Arava valley, Israel. An outbreak is also recorded on Citrus paradisi (Swirski 1963). B. flavidus (Ivanoskaya), described from Peganum harmala in Turkmenistan, is not clearly distinct. 2n=8.

Brachyunguis kaussarii Remaudière & Davatchi Colour of apterae in life is unknown; BL 1.3-1.6 mm. Alatae have 7-9 secondary rhinaria on ANT III. On Stocksia brahuica (recorded as “bralmica” in error; G. Remaudière, pers. comm.) in Iran.

Brachyunguis letsoniae Das Apterae are green, covered in white wax powder, BL c. 1.85 mm. Alatae have dark cross bars on abdominal tergites 6-8, and 6-8 rhinaria on ANT III. Feeding on young growth and inflorescences of “Letsonia scandens” (possibly this is Lettsomia (= Argyreia) splendens?), in Pakistan in October. It has not been reported since its original description.

Brachyunguis lycii (Nevsky) Apterae are pale green coated with mealy wax; BL 1.2-1.6 mm. In large colonies on leaves and fruits of Lycium spp.in Central Asia. Specimens conforming to Nevsky’s description have also been found on Lycium in Cyprus, Egypt and Iran (BMNH colln), although it is unclear how this species differs from B. harmalae colonising various other plants in these regions. 2n=8*.

Brachyunguis monstratus Kadyrbekov Apterae are dark green with dark red eyes; BL 1.0-1.4 mm. On species of Atraphaxis and on Diarthron lessertii in Kazakhstan, Iran and Xinjiang, China (Kadyrbekov 1999c, 2004b, 2014c; Barjazde et al. 2022).

Brachyunguis nevskyi (Kreutzberg) Apterae are dark green; BL c. 1.15 mm. On Pistacia vera in south-west and central Asia, colonising inflorescences in spring and new shoots, petioles and leaves in autumn (Ivanoskaya 1960, as Xerophilaphis).

Brachyunguis nurikamalae Kadyrbekov Apterae are greenish with brown head, without wax powdering: BL 1.5-2.0 mm. On inflorescences of Ziziphora sp. in northern Kazakshtan, attended by ants (Kadyrbekov 2014c). Alatae have 5-9 secondary rhinaria on ANT III.

Brachyunguis peucedani (Nevsky) Apterae are pale green covered with wax; BL 1.6-1.8 mm. On flowers and stems of Peucedanum pseudoreoselinum in Uzbekistan, and recorded from Ferula spp. in Kazakhstan by Kadyrbekov (2003c). To judge from the descriptions by Nevsky and by Ivanoskaya (1960), placement of this species in Semiaphis by Eastop & Hille Ris Lambers (1976) and Remaudière & Remaudière (1997) seems to have been in error.

Brachyunguis poacynophila Zhang, Chen, Zhong & Li Appearance of aptera in life is unrecorded; BL c.1.4 mm. On Apocynum venetum in north-west China (G. Zhang 1999).

Brachyunguis rhei (Nevsky) Apterae are yellowish brown; BL c. 2 mm. On Rheum maximoviczii, forming large colonies on undersides of leaves, and subsequently recorded from several Rumex spp. In southern Kazakhstan and Turkmenistan.

Brachyunguis salsolacearum (Nevsky) Apterae are green, covered with fine pruinose secretion, siphunculi and cauda green, dorsal abdomen with small dark green spots; BL 1.3-1.9 mm. In large numbers on young growth and stems of various Amaranthaceae (Anabasis, Girgensohnia, Halocharis, Salsola) in Central Asia (Narzikulov & Daniyarova 1990).

Brachyunguis saxaulica (Nevsky) Apterae are brownish yellow, greenish or dark brown, covered with fine wax; BL 1.6-2.1 mm. At ends of young shoots of Haloxylon spp., forming “cone-like pseudogalls”, ant-attended. In Kazakhstan, Tajikistan and Turkmenistan (Ivanoskaya 1960, as Xerophilaphis), and also recorded from Iran (Rezwani 1987).

Brachyunguis shaposhnikov Ivanoskaya Apterae are green, tinged with orange near siphunculi, without wax; BL c.2.2 mm. Described from a plant of the Polygonaceae, and recorded from Polygonum aviculare by Kadyrbekov (2017a). In western Kazakhstan.

Brachyunguis skafi Remaudière & Talhouk Apterae are very dark, blackish; BL 1.2-1.9 mm. Recorded from three species of Astracantha (previously all three have been placed in Astragalus) in Turkey, Iran and Lebanon. Monoecious holocyclic with oviparae and apterous males in September-October (Remaudière & Talhouk 2000).

Brachyunguis suaedus (Paik) The aptera described from Suaeda glauca in Korea by Paik (1965, as Hyalopterus) is Clypeoaphis suaedae, and the alatae may be vagrants from another plant. The length of the antennal terminal process and black pigmentation suggest that it does not belong in Brachyunguis, where it is currently placed.

Brachyunguis tamaricis (Lichtenstein) Plates 13a, b, fig. 113B Apterae according to the original description are velvety grey-green, with no mention of wax dust (but development of wax may depend on age and microclimate); BL is 0.9-1.7 mm. Alatae have secondary rhinaria distributed ANT III 2-8, IV 0-2, V 0-1. Colonies on twigs are inconspicuous, resembling small leaves or leaf-scales of the host plant. They may be attended by ants (Monomorium). On Tamarix spp. in south, central and eastern Europe, North Africa, south-west and central Asia east to Pakistan. Monoecious holocyclic. Ivanoskaya (1956) described the fundatrix on T. meyersi in Turkmenistan (as tamaricophila), and specimens from Pakistan on T. kotschyi in the BMNH collection also seem to be fundatrices. Oviparae are recorded from Portugal (Ilharco 1996a) and oviparae and apterous males have been found in France (G. Remaudière, cited in Nieto Nafría et al. 2005a). 2n=8.

[There is much confusion in the literature on the nomenclature of the Tamarix-feeding species of Brachyunguis. B. tamaricophila of Ivanoskaya (1956; nec. Nevsky 1928b) seems to be tamaricis, and B. tamaricivorum Narzikulov) (= tamariciarum Ivanoskaya) is not clearly distinct. B. tamaricifoliae (Hall 1926) is also a synonym of tamaricis. Records of tamaricis from Calotropis procera probably refer to either B. harmalae or B. calotropicus. See also Nieto Nafría et al. 2005a. ]

Brachyunguis tamaricophilus (Nevsky) (fig. 113A) Apterae according to the original description are green, densely covered with mealy wax secretion; BL 0.7-1.7 mm. In large, ant-attended colonies on young shoots, leaves and flowers of Tamarix spp. Monoecious holocyclic in Uzbekistan (Tashkent), oviparae being found, but males are still undescribed (Nevsky 1928b). Ivanoskaya (1956) described the fundatrix on T. karakalensis in Turkmenistan (as tamaricifoliae). It is also recorded from Kazakhstan, Iran, Israel, Libya and Pakistan (BMNH collection). B. tamaricifoliae of Ivanoskaya (1956; nec. Hall 1926) seems to be this species. [The account of this species from Sicily by Barbagallo & Stroyan (1982) applies to B. tamaricis.]

Brachyunguis tausaghyz (Nevsky ex Ivanoskaya) Apterae are greenish yellow, with pruinose covering, and pale siphunculi; BL c.1.6 mm. On Scorzonera tausaghyz and Seriphidium nitrosum in Kazakhstan, colonising flower heads (Narzikulov et al. 1971). Kadyrbekov (2017a) reported it also from Chondrilla brevirostris. B. afghanica, described from Artemisia turanica in Afghanistan (Narzikulov & Umarov 1972), does not seem to be distinguishable from this species on the basis of the published descriptions.

Brachyunguis tetrapteralis (Cockerell) Apterae are pale greyish green to blue-green, with antennae dusky/dark distally; BL 1.0-1.7 mm. On stems and leaves of Atriplex spp in western and south-western USA, and Mexico, and there is also a record from a Chenopodium sp. Usually ant-attended. Monoecious and holocyclic, with sexuales in early October (aphidtrek.org). Records from Suaeda (e.g. Remaudière & Halbert, 1996) should perhaps all be referred to B. cahuille, which has a dark, very swollen clypeus, shorter siphunculi and shorter antennal terminal process. Records from Sarcobatus are possibly all referable to B. bonnevillensis, and records from plants outside the Amaranthaceae also need to be treated warily. The synonymy of Aphis piutapa (Hottes & Wehrle 1951b, on Lycium parviflorum) with B. tetrapteralis is possibly incorrect, as the description of A. piutapa differs in certain respects such as the shape of the cauda. The palaearctic species B. harmalae is very similar, but has slightly longer dorsal body hairs.

Brachyunguis tigrahaudus Kadyrbekov Apterae are greenish with brown head, without wax powdering; BL 1.0-1.4 mm. On flower heads of Helichrysum arenarium and Jurinea cyanoides in Aral region of south-west Kazakhstan (Kadyrbekov 2014c).

Brachyunguis transaralensis Kadyrbekov Apterae greenish, without wax powder; BL 1.5-1.8 mm. Alatae have 5-9 secondary rhinaria on ANT III. On flowerheads of Chondrilla brevirostris, attended by ants, in Aral region of south-east Kazakhstan (Kadyrbekov 2014c).

Brachyunguis zawadovskii (Nevsky) Apterae are green, covered with mealy wax; BL 1.6-1.9 mm. In large colonies on upper sides of leaves of Lycium depressum (= turcomanicum) in Central Asia.

Brachyunguis zygophylli (Nevsky) Apterae are yellow, yellow-green or green to dark green, in life covered with dense mealy wax; BL 1.6-2.0 mm. In large colonies on stems, flowers and undersides of leaves of Zygophyllum fabago. In south-west (Turkey, Iran) and Central Asia. The record from Spain (Gómez-Menor 1951, as Brachyunguis zygophilli) should be referred to B. harmalae (Garcia Prieto et al. 2004, Nieto Nafría et al. 2005a).

Braggia Gillette & Palmer

Aphidinae: Aphidini

About 12 nominal taxa living without host alternation on Eriogonum in western USA. Related to Aphis but characterised by a combination of short antennal terminal process, short siphunculi, and usually a very short broad cauda. The dorsal cuticle is strongly reticulated and often has variably developed dark sclerotisation. Apterae often have secondary rhinaria on ANT III or III-V. K.S.Pike et al. (2009) reviewed the genus and keyed the species, studying mitochondrial DNA sequence variation in six species and describing two new ones. They also provided information on primary and secondary parasitoids. Hille Ris Lambers (1966a) described subspecies of three of the species on the basis of small morphological differences, but further work is needed to assess the significance of the observed variation.

Braggia agathona (Hottes) Apterae are pinkish brown, often with a dark central spot on the abdomen, the whole body covered with white wax; BL 1.4-1.8 mm. On undersides of leaves of Eriogonum corymbosum in Colorado and Utah, USA. Monoecious holocyclic, with oviparae and alate males in October (original description).

Braggia columbiana Pike Apterae are black or blackish brown with a white or whitish grey waxy dorsal pattern (see http://www.flickr.com/photos/sandnine/4606202941/); BL 1.4-2.6 mm. On Eriogonum spp., predominantly E. compositum, in Oregon and Washington, USA (K.S.Pike et al. 2009). Apterous and alate viviparae occur from about mid-May to mid-September, sexual morphs are undescribed.

Braggia deserticola Hille Ris Lambers Apterae are shining black dorsally, grey and wax-powdered ventrally, with black appendages; BL 1.1-1.8 mm. Found in small numbers in flower heads of Eriogonum fasciculatum, incl. ssp. polifolium, in California and Utah, USA, and in Mexico. A sample from Eriogonum sp. at a higher altitude in California with shorter R IV+V and paler tibiae was distinguished as a subspecies, B. deserticola thanatophila Hille Ris Lambers.

Braggia echinata Gillette & Palmer (Fig.27e) Apterae are greyish olive-green, usually with a slight rusty area at base of each siphunculus, and with white dorsal reticulations and pale, thick hairs giving a frosted appearance; BL 1.3-1.9 mm. On leaves and stems of Eriogonum corymbosum and Eriogonum sp. in western USA (Colorado, Utah). Oviparae and alate males in October (Palmer 1952).

Braggia eriogoni (Cowen) Plate 8d (Fig.27a,b,g) Apterae are black or brownish black with a dorsal pattern of white reticulation, accentuated by a wax secretion; BL 1.1-2.1 mm. On leaves, stems and flowerheads of Eriogonum spp. in western USA. Hille Ris Lambers (1966a) distinguished several subspecies on the basis of differences in hair length, siphuncular shape and extent of dorsal pigmentation, but further work is needed to investigate whether any of the observed variation is correlated with host plant, season or geographical distribution. Sexuales have not been described.

Braggia longicauda Pike Apterae are black, blackish grey or blackish brown, with white wax powdering; BL 1.5-2.9 mm. On Eriogonum spp., predominantly E. elatum, in western USA (Washington, Oregon, California). Oviparae and alate males occur in September-October (K.S.Pike et al. 2009).

Braggia uncompahgrensis Hottes (Fig.27f) Apterae are pale grey-green, with frosty appearance due to thick pale dorsal hairs; BL 0.9-1.4 mm. On Eriogonum corymbosum in Colorado and Utah, USA. Oviparae and alate males occur in October (Palmer 1952). Doubtfully distinct from B. echinata.

Braggia urovaneta (Hottes) (Fig.27c,d) Apterae are black, appearing grey due to powdering with wax; BL 0.9-1.9 mm. On flower stems and upper parts of Eriogonum sp(p). in western USA. Oviparae and alate males were collected in Colorado on E. corymbosum in early October (Hottes 1950). Hille Ris Lambers (1966a) distinguished a population with a shorter R IV+V on E. latifolium in California as a subspecies, B. urovaneta ssp. pachysiphon.

Brevicoryne van der Goot

Aphidinae: Macrosiphini

Eight palaearctic and two nearctic species all except one of which live without host alternation on Brassicaceae. The exception is a Salix feeder and should possibly be given its own genus. Brevicoryne are related to Lipaphis and characterised by very low rounded antennal tubercles, short siphunculi and broad triangular cauda. Eastop in Hodjat (1981) gave a key to alate viviparae of Middle Eastern species.

Brevicoryne arctica Richards Apterae are very dark blue or green, with wax; BL 1.7-2.mm. Alatae have secondary rhinaria distributed III 12-13, IV 5. On Lesquerella arctica in northern Canada (Ellesmere Island). Monoecious holocyclic with oviparae and apterous males in late July (original description).

Brevicoryne barbareae Nevsky Apterae are dark green, dark brown to black, with appendages dark except for pale ANT III and tibiae (black at apices); BL 1.5-2.0 mm. Alatae have secondary rhinaria distributed III 17-20, IV 3-5. On undersides of leaves of Barbarea vulgaris in Central Asia. This name has been applied to a species on Barbarea and Nasturtium in north-west India that differs from Nevsky’s species in several respects, having apterae with extensive dorsal sclerotisation, dark tibiae, a much longer cauda, and alatae (brachypterae) with rhinaria distributed III 35-36, IV 0-1 (David & Hameed 1975). Alate males have been described of this Indian species (L.K. Ghosh et al. 1980, as B. barbareae).

Brevicoryne brassicae (L.) Cabbage Aphid or Mealy Cabbage Aphid Plate 12f Apterae are greyish green or dull mid-green with dark head and dark dorsal thoracic and abdominal markings, densely coated in greyish white mealy wax (see influentialpoints.com/Gallery); BL 1.6-2.6 mm. Alatae have a dark head and thorax and dark cross-bands on dorsal abdomen. It occurs on many genera and species of Brassicaceae, and is a major pest of field crops (Blackman & Eastop 2000) in all temperate and warm temperate parts of the world. Feeding is stimulated by the presence of mustard oil glycosides, so other plants with these compounds, e.g. Reseda, are also colonised. Monoecious holocyclic with alate males in colder regions, anholocyclic wherever winters are mild. Mustaţă et al. (2000) studied the large complex of natural enemies of B. brassicae in Romania. Ruiz-Montoya et al. (2003) used enzyme electrophoresis to look for host-related genetic structure in Mexican populations, and Ruiz-Montoya et al. (2005) and Ruiz-Montoya & Núñez-Farfán (2013) studied phenotypic differences between populations on two sympatric Brassica species. The finding by Rebijith et al. (2013) of CO-I barcode differences indicative of possible cryptic species within Indian populations of B. brassicae on Raphanus sativus requires further investigation. 2n=16.

Brevicoryne crambe Bozhko (= Brevicoryne crambinistataricae Bozhko) Apterae are pale green, wax-dusted, BL 1.9-2.8 mm. On terminal shoots and stems of Crambe tatarica, and also recorded from species in other crucifer genera (e.g. Barbarea, Isatis, Diplotaxis, Sisymbrium). In Hungary, Moldova, Armenia, Bulgaria, Turkey, Ukraine and Iran.

Brevicoryne fraterna (Strom) Apterae are pale green to yellow-green; BL 1.2-1.8 mm. On leaves and young twigs of unidentified species of Salix in USA (Illinois, Ohio, Utah, Colorado). Monoecious holocyclic with oviparae and apterous males in October (Palmer 1952).

Brevicoryne jiayuguanensis (Zhang, Chen, Zhong & Li) Apterae are silvery whitish green, powdered with wax; BL 1.5-1.8 mm. On Armoracia rusticana in Gansu province, China (Zhang 1999, as Brachycolus). The broadly triangular cauda and many (15-19) thick hairs on abdominal tergite 8 as well as the host association indicate that the species belongs in this genus.

Brevicoryne lonicerina Mukhamediev & Akhmedov Apterae are pale green with brown appendages; BL 2.3-2.6 mm. On Lonicera spp. in Central Asia.

Brevicoryne nigrisiphunculata Hodjat Apterae are yellow to dusky green, with short black siphunculi; BL 2.1-2.7 mm. On undersides of older leaves of Crambe and Brassica in Turkey, Iran, Armenia (Szelegiewicz 1979, as B. crambinistataricae) and Ukraine.

Brevicoryne shaposhnikovi Narzikulov Apterae are pale green dusted with grey wax; BL c.1.6-1.7 mm. On Lonicera spp. in Central Asia (Mukhamediev & Akhmedov 1982).

Brevicorynella Nevsky

Aphidinae: Aphidini

A genus with two species resembling Brachyunguis but with protruberant eyes lacking ocular tubercles. [Placed in Macrosiphini by Remaudière & Remaudière (1997), perhaps because of the absence of marginal tubercles, but the spiracles on abdominal segments 1 and 2 are widely spaced, which is a more reliable indicator that it should be placed in Aphidini.

Brevicorynella quadrimaculata Nevsky Apterae brown, dark brown or dark green covered with a fine pruinose secretion, antennae pale yellow with articulations and all of last segment black, abdomen with four blackish-brown dorsal spots; BL 1.1-1.6 mm. The aphids cluster on the ends of shoots of Tamarix spp., and are attended by ants (Crematogaster). In central Asia (Kazakhstan, Tajikistan, Uzbekistan) (Nevsky 1928b), Iran (Barkhordari et al. 1981), and also recorded from Tamarix chinensis (= T. ramosissima) in China (Xinjiang; G. Qiao, pers. comm.).

Brevicorynella sexmaculata Qiao & Zhang Apterae are green covered with a fine pruinose secretion, with three pairs of dark patches on dorsal abdomen; BL 1.3-1.6 mm. Alatae are undescribed. On Tamarix chinensis (= T. ramosissima) in Xinjiang province, China (Qiao et al. 2004b).

Brevisiphonaphis Stekolshchikov & Qiao

Aphidinae: Macrosiphini

A genus for one Artemisia-feeding species in north-east China, related to Microsiphum and Macrosiphoniella but with an unusual form of cauda and long fine hairs on body and appendages. Alatae are unknown.

Brevisiphonaphis hirsutissima Stekolshchikov & Qiao Colour of apterae in life is unrecorded, BL 1.6-1.8 mm. On Artemisia argyi in Liaoning prov., China (Stekolshchikov & Qiao 2008a, b). Life cycle is unknown.

Burundiaphis Remaudière

Aphidinae: Macrosiphini

One African species superficially resembling Paczoskia, but with R IV+V having numerous short stiff hairs and very long siphunculi without subapical reticulation. Alatae are unknown.

Burundiaphis autriquei Remaudière Apterae are shining greenish black; BL 2.2-2.9 mm. On Davallia chaerophylloides ( = denticulata) epiphytic on oil-palms, often in mixed colonies with Micromyzella. Only known from Burundi (Remaudière & Autrique 1985).

Byrsocryptoides Dzhibladze

Eriosomatinae: Eriosomatini

Two species described from galls on the primary host (Zelkova), one of which is now known to migrate to Carex. Alate spring migrants, emerging from pseudogalls on Zelkova, have an unbranched media, only one oblique vein in the hind wing, and inconspicuous ring-like siphunculi. Resembling Kaltenbachiella in many characters, but in Byrsocryptoides the secondary rhinaria are less annular and the processus terminalis is relatively longer, and a detailed cladistic study by Sano & Akimoto (2011) placed this genus in a basal position in the tribe Eriosomatini. Dzhibladze (1965) provided keys to fundatrices, apterous fundatrigeniae and spring migrants of the species on Zelkova.

Byrsocryptoides zelkovae Dzhibladze The galls (pseudogalls) on Z. carpinifolia in Georgia are formed by rolling, distortion and blistering of a terminal or subterminal leaf, starting at the base. Alatae (fig. 120D) leaving the galls in June are small (BL about 1.2-1.3 mm), rather long-bodied, with dark brown head, meso- and metathorax and pale yellow to green prothorax and abdomen (Dzhibladze 1960b). They migrate to aerial parts of Cyperaceae; apterae of elongate form with a long last rostral segment collected on Carex riparia in Germany (BMNH collection, leg. F.P. Müller) are provisionally assigned to this species, and Barjadze & Gratiashvili (2009) obtained experimental colonisation of Carex capitata in Georgia.

Byrsocryptoides zelkovaecola Dzhibladze Galls (fig. 125A) are similar to those of B. zelkovae, and occur on Zelkova carpinifolia in Georgia. Alatae (fig. 120C) of BL 1.1-1.4 mm emerge in June-July. The life cycle is unknown (Dzhibladze 1965b).