THE APHIDS

 

 

         SYSTEMATIC TREATMENT OF APHID GENERA

        (in alphabetical order)

 

D

Daktulosphaira

Dasyaphis

Davatchiaphis

Decorosiphon

Defractosiphon

Delphiniobium

Dermaphis

Dimelaphis

Dinipponaphis

Diphyllaphis

Diprociphilus

Distylaphis

Diuraphis

Doraphis

Drepanaphis

Drepanosiphoniella

Drepanosiphum

Durocapillata

Dysaphis

 

 

Daktulosphaira Shimer

Phylloxeridae

 

A genus for the grape phylloxerid, which differs from Phylloxera by having alatae in which the distal sensorium on the antenna is not greatly enlarged. The generic classification of the Phylloxeridae is likely to remain unsatisfactory until the 34 species of Phylloxera described from Carya in North America are better known.

Daktulosphaira vitifoliae (Fitch)  Grape Phylloxera    Apterae are yellow; BL 0.7-1.4 mm. In hairy, scabrous galls on the undersides of vine leaves (but opening on the upper surface of the leaf); or on vine roots causing bird's head‑like swelling and blackening of rootlets. Of North American origin, now in Europe, the Mediterranean, the Middle East, Africa, Korea, China, Australia, New Zealand, and Central and South America. The holocycle takes two years and involves a sexual phase and leaf‑ and root‑feeding stages on American vine species, but on European vine (Vitis vinifera) it is normally anholocyclic on the roots. Leaf galls occur in Europe on cultivars derived from hybrids between vinifera and American vine species. Witiak (2006) made detailed histological and molecular investigations of the development of the leaf galls. There has been much study of genetic variation in and between clonally-structured populations (e.g.  Downie et al. 2001; Vorwerk & Forneck 2006, 2007; Sun et al. 2009; Forneck et al. 2015), and of differences in virulence between clonal lineages (Herbert et al. 2010; Powell et al. 2013).  Benheim et al. (2012) reviewed alternative pest management options. For further general information see Blackman & Eastop (2000, as Viteus vitifoliae). 2n=10.

 

 

Dasyaphis Takahashi

Calaphidinae: Panaphidini

 

A genus for two oriental species associated with Juglandaceae, the apterae of which have fused head and prothorax, reduced eyes and antennae and long hair-bearing processes; although in D. mirabilis an apterous morph without such tubercles can also occur in colonies. Quednau (2003) provided a key couplet to distinguish the two species. Sorin (2009) redescribed both species from Japan and provided keys to apterous and alate viviparae and alate males (in Japanese).

Dasyaphis mirabilis (Tseng & Tao)  Apterae are dorso-ventrally flattened, pale yellowish green, BL c. 1.3 mm.  Apterae both with and without finger-like dorsal processes occur. On undersides of leaves of Pteryocarya stenoptera in China, and redescribed from Juglans ailanthifolia in Japan by Sorin (2009). Alate males occur in October-November (Tseng & Tao 1938, as Sinocallis mirabilis; Sorin 2009). Quednau (2003) illustrated the alata and three variants of the apterous morph.

Dasyaphis rhusae (Shinji)  Apterae are pale yellowish-green, yellow or white; BL 1.2-1.4 mm.  On undersides of mature leaves of Juglans spp. (ailanthifolia, mandshurica, regia) in Japan (Takahashi 1965, Higuchi 1972; as Dasyaphis onigurumi), Korea (Paik 1972), Siberia (Quednau & Shaposhnikov 1988). and China (Qiao et al. 2005b). Alate males were found in early November by Sorin (2009).

 

Davatchiaphis Remaudière

Aphidinae: Macrosiphini

A genus for one very long-bodied species on Phragmites, somewhat resembling Brachycolus but with Phorodon-like processes on the antennal tubercles.

Davatchiaphis persica Remaudière  Plate 14e   Aptere are pale green; BL 2.1-2.4 mm. On upper sides of leaves of Phragmites australis (var. stenophyllus) in Iran (Remaudière 1964). Monoecious holocyclic with oviparae and apterous males in November.

 

Decorosiphon Börner

Aphidinae: Macrosiphini

One distinctive moss-feeding species with long stiff hairs on appendages, spinulose head and swollen siphunculi, as well as the pointed antennal terminal process and characteristically-shaped cauda of a moss-feeding aphid.

Decorosiphon corynothrix Börner   Plate 17a  (Fig.44c)   Apterae are brownish green to brownish yellow; BL 1.4-1.9 mm. On basal parts of Polytrichum spp. and other mosses of family Polytrichaceae growing in damp, shady situations. It is also recorded from Rhytidiadelphus squarrosus. In Europe and eastern North America. Anholocyclic; no sexual morphs are known. Heinze (1953) provided full descriptions of apterae and alatae, and Müller (1973b) discussed its biology.

 

Defractosiphon Börner

Aphidinae: Macrosiphini

Two palaearctic species on Umbelliferae/Apiaceae, related to Hyadaphis but with distinctive siphuncular morphology.

Defractosiphon brevisiphon Mamontova  Plate 13c   Apterae are mottled green with slight whitish pulverulence, head yellowish; BL 1.6-1.9 mm. Alatae are undescribed. On Seseli spp. (campestre, elatum, osseum) in Europe (specimens in BMNH collection from Czech Republic, France, Hungary, former Yugoslavia, Ukraine), and also recorded from Ptychotis saxifraga (Leclant 1978).

Defractosiphon franzi Börner  (Fig.41a)   Apterae are shining green on dorsal abdomen, anteriorly yellowish brown: BL 1.8-2.8 mm. Described from Seseli austriacum, and subsequently found on Peucedanum spp., in rows on upper sides of leaflets with heads directed towards the petioles (Müller 1973a, as D. rugosus). Infested leaves turn yellow In Germany and Ukraine. Monoecious holocyclic on Peucedanum, with oviparae and alate males in October.

 

Delphiniobium Mordvilko

Aphidinae: Macrosiphini

Eleven large species mostly associated with Aconitum, tending to have aposematic coloration and probably sequestering toxins from the host plant. They have remarkably large thoracic spiracles (Hille Ris Lambers 1947a). Qiao & Zhang (2000d) reviewed and keyed the Chinese species.

Delphiniobium aconitifoliae Zhang & Qiao    Apterae are red and yellow with mainly black appendages: BL c. 3.6 mm. On young shoots and upper sides of leaves of Aconitum kusnezofii in China. Monoecious holocyclic, with oviparae and alate males in October (Qiao & Zhang 2000d).

Delphiniobium bogdouli Szelegiewicz    Apterae are undescribed; alatae are shining brown, BL 3.7 mm. On Aconitum barbatum in Mongolia.

Delphiniobium canadense (Robinson)    Apterae are lemon yellow (immatures whitish); BL 2.1-2.6 mm. On Lonicera involucrata in western North America. Monoecious holocyclic, with oviparae and apterous males in June-July (BMNH collection, leg. C.-k. Chan). 2n=20.

Delphiniobium carpaticae Mamontova    Apterae are whitish-green, with cauda and tips of siphunculi dark; BL c.3.2 mm. On Aconitum spp. in Ukraine. Not clearly distinct from D. lycoctoni, except in pigmentation.

Delphiniobium. gyamdaense Zhang    Apterae are yellow with black appendages; BL c. 3.0-3.1 mm. On an Aconitum sp. in China and Tibet (Qiao & Zhang 2000d).

Delphiniobium hanla Paik  (Fig.6k)   Apterae are yellowish orange with mainly black appendages, siphunculi yellowish orange basally and black distally; BL 3.1-3.8 mm. On Aconitum spp. and Delphinium spp. in Korea and east Siberia, colonising upper parts of flower stems (Pashchenko 1988a). It is now also reported from Kazakhstan (Kadyrbekov 2012a).

Delphiniobium junackianum  (Karsch)  Plate 25e  (Fig.6h,i)   Apterae are bluish green, with mainly dark antennae legs, siphunculi and cauda (see influentialpoints.com/Gallery); BL 2.9-4.7 mm. On Aconitum and Delphinium spp., forming colonies mainly on upper parts of stems and between flowers. In north-west and central Europe, and eastward to west Siberia. Monoecious holocyclic, with oviparae and alate males in late September-October. Aphidius sussi Pennacchio & Tremblay is a possibly specific parasitoid.

Delphiniobium lycoctoni Börner  (Fig.6j)   Apterae are yellowish, with pale antennae, legs and siphunculi, cauda dark grey; 3.0-4.2 mm. On Aconitum lycoctonum, curling and spotting leaves. In Austria, France, Italy, and also reported from A. leucostomum in eastern Kazakhstan (Kadyrbekov 2009a). Very similar to D. carpaticae except for pigmentation.

Delphiniobium ussuriense Pashchenko    Apterae are green, shiny, with dark appendages; BL c.3.6 mm. On upper parts of stems of Artemisia sylvatica in east Siberia (Primorskii) (Pashchenko 2000b). The generic placement of this species is uncertain.

Delphiniobium violisuctum Zhang & Qiao    Aptera is dark green, BL c.3.1 mm. Described from one aptera, one alata and one immature found on the upper side of a leaf of Viola verecunda, which is unlikely to be the true host plant. In Shanxi province, China (Qiao & Zhang 2000d).

Delphiniobium yezoense Miyazaki    Apterae are green or bluish green, with mainly pale, black-tipped  appendages and a black cauda; BL 2.9-3.5 mm. On Aconitum spp. in Japan, China and west Siberia (Stekolshchikov et al. 2008a). Monoecious holocyclic with oviparae and alate males in October (Miyazaki 1971). 2n=12 (Chen & Zhang 1985a).

 

Dermaphis Takahashi

Hormaphidinae: Nipponaphidini

Three species related to Thoracaphis, with apterous viviparae on Fagaceae that have a heavily sclerotised and sculptured dorsal cuticle and long pointed marginal prosomal hairs.  Siphunculi are lacking and the tarsi are often without claws.  Descriptions and available specimens are inadequate to include a key to species here.  We follow A.K. Ghosh & Raychaudhuri (1973) and Sorin (2006) in including three species in this genus, although this is mainly for convenience as these species may not be very closely related. Sorin (2006) reviewed and keyed all three species in Japan.

Dermaphis crematogastri (Takahashi)  Apterae are black, with some wax.  Body oval, convex dorsally, extended somewhat at posterior end, sometimes asymmetrical and irregular in outline; BL (0.8-) 1.0-1.1 mm.  The prosoma has many long stout spinal as well as marginal hairs.  Tarsi usually have claws.  On branches of Quercus sp. in Thailand, attended by ants (Crematogaster sp.), and often in their shelters (Takahashi 1941; as Thoracaphis). Recorded also from Quercus spp. Japan (Sorin 2006), although with discrepancies from the original description.

Dermaphis japonensis Takahashi   Apterae are black, with some greyish-white wax.  Body oval, convex dorsally, extended slightly at posterior end; BL 1.1-1.8 mm.  Even the hind legs lack claws, and are often concealed beneath body. Dorsal median area of prosoma with only 4 pairs of hairs, which are shorter than the marginal hairs (Takahashi 1958).  On branches of Quercus spp. (gilva, glauca) and Lithocarpus sp. (Shinji 1941; as Thoracaphis takahashii) in Japan, and on Quercus sp. in Korea (Paik 1972).  Ant attendance is not recorded. Sorin (2006) described the alata from Japan, and apparently observed migration to Distylium racemosum, and larvae deposited on the undersides of leaves moving to the bases of the buds, but the life cycle could not be followed further.

Dermaphis takahashii (Strand)  Apterae are black, with white wax around margin.  Body subcircular, densely convex; BL c. 1.2 mm.  The hind legs project behind the body and the tarsi always have claws.  In small groups on branches and stems of Quercus glauca, Quercus sp. and Lithocarpus sp., usually in ant shelters made by Crematogaster, in Taiwan, China and Japan (Takahashi 1935d, as Thoracaphis; Tao 1966; Sorin 2006).

 

Dimelaphis Zhang

Eriosomatinae: Fordini

 

One species, D. obtusifalcata Zhang, only known from two alate sexuparae, presumably vagrants, collected on Juglans regia in China. The genus is distinguished by the shape of the pterostigma and the head with a medial suture (G. Zhang & Qiao 1998a).

 

Dinipponaphis Takahashi

Hormaphidinae: Nipponaphidini

 

A genus for one species related to Nipponaphis, living without host alternation on Distylium.  The alate sexuparae have clear wings and characteristic sensoriation of the antennae.  Ghosh & Raychaudhuri (1973) revised the genus.

Dinipponaphis autumna (Monzen)  In small reddish galls, projecting on both sides of leaves, of Distylium racemosum in Japan (fig. 130A and Moritsu 1983).  There is no host alternation.  Alate sexuparae emerging the galls in early December have black head and thorax, clear wings with pale veins and pterostigma, and yellow abdomen; BL 1.5-1.7 mm (Takahashi 1962).  They deposit males and oviparae of very different sizes (Sorin 1987) on the undersides of the leaves.  Adult sexuales overwinter on the buds, eggs are laid in March and fundatrices hatch in April (Sorin 1960). Aoki et al. (1999) reported on defensive behaviour of first instar larvae within the galls.

 

Diphyllaphis Takahashi

Phyllaphidinae

 

A genus of four palaearctic and one nearctic oak-feeding species, related to the wholly Nearctic genus Stegophylla, but the siphunculi are small pores without encircling hairs.  Higuchi (1972) revised the Japanese species, and Quednau (2010) provided keys to the world fauna and illustrations of all available morphs.

Diphyllaphis alba Takahashi   Appearance in life not recorded, presumably entirely white with flocculent wax; BL 1.1-1.4 mm.  On Quercus phillyraeoides in Japan (Honshu, Kyushu; Takahashi 1960, Higuchi 1972).  Apterous viviparae were collected in April (BMNH collection, leg. R. van den Bosch) and September-October, and ovipara and small apterous males were found in December (Quednau 2010).

Diphyllaphis konarae (Shinji) Apterae elongate pear-shaped, light red, orange or white, with white flocculent wax; BL 1.2-1.7 mm.  On Quercus spp. in Japan (Honshu), on rather mature leaves that curl up and may be bound together by the wax secreted by the aphid (Shinji 1924, Takahashi 1960).  Only apterous viviparae are known, collected in October. In Japan (Honshu) and South Korea (Quednau & Lee 2001).

Diphyllaphis microtrema Quednau   Apterae are broadly oval, pale yellowish-green, with dense white flocculent wax; BL 1.7-2.2 mm.  In dense woolly masses on undersides of leaves of Quercus spp. (rubra, velutina, virginiana) in eastern USA (Connecticut, North Carolina, Florida).  Holocyclic in Connecticut; oviparae and alate males occur in late September (Quednau 1971, 2010).

Diphyllaphis mordvilkoi (Aizenburg)  Apterae are elongate pear-shaped, yellowish-white, with much flocculent white wax; BL 1.2-1.5 mm.  On undersides of mature leaves of Quercus spp., causing yellowing and necrosis (Binazzi & Roversi 1988).  In Spain, Italy (Tuscany, Sicily), Turkey, Lebanon (BMNH collection) and Caucasus.  Holocyclic, with apterous males (Roversi & Binazzi 1990). [Contrary to Hales et al. (1997), oviparae are also apterous.] The male genitalia were described and illustrated by Wieczorek et al. (2011).

Diphyllaphis (Nymphaphis) quercus (Takahashi)  Apterae are elongate pear-shaped, whitish with flocculent wax; BL 1.4-1.6 mm.  On Quercus spp., curling the tips of the leaves and causing necrosis.  In Japan, China and Korea.  Moritsu (1983) described and illustrated the alate viviparous female.  Oviparae occur in Korea in early November (Paik 1972, as Nymphaphis quercus), and Quednau (2010) illustrated an ovipara and a small apterous male collected in late October- November. [Note: Aplonervoides erythrocereus Zhang was synonymised with D. quercus by Remaudière & Remaudière (1997), but is described as bright red in life with long wax filaments (Zhang et al. 1992a).]

 

Diprociphilus Zhang & Qiao

Eriosomatinae: Pemphigini

One species in China related to Prociphilus, but with a much longer rostrum than any known species of that genus.

Diprociphilus allivorus Zhang, Qiao & Zhao   Apterae are yellowish white in life, presumably secreting abundant wax; BL c.1.9 mm. On Allium victorialis in Shanxi Province, China (Qiao et al. 1999b). The part of the plant colonised is not mentioned, and other morphs and life cycle are unknown.   

 

Distylaphis Noordam

Hormaphidinae: Nipponaphidini

 

One species producing galls on Distylium in Java, but also having free-living generations on the same host.  First instar soldiers occur in the galls (Noordam 1991, Fukatsu & Antonius 1998, Fukatsu et al. 2005).

Distylaphis foliorum (van der Goot)  Galls on Distylium are spherical, 5-12 mm in diameter, green-reddish-brown in colour and formed on the leaves (Fukatsu & Antonius 1998).  Alatae emerging from galls in late January have BL 2.6-3.3 mm, long forewings with a broad black pterostigma and the wing membrane fuscous basad of Cu1b, and secondary rhinaria distributed III 20-26, IV 8-12, V 7-10.  Apterae are bright greenish, margins of body with thick powdery or flocculent wax, and with two longitudinal rows of wax patches or a thick coat of wax on dorsum; BL c.1.5 mm.  On undersides of leaves of Distylium stellare in Java.  Apparently monoecious holocyclic, but the life cycle needs further clarification.  A morphologically differentiated 1st instar soldier caste was demonstrated by Fukatsu et al. (2005). Oviparae are present on Distylium leaves in April-August; males are unrecorded (Noordam 1991).

 

Diuraphis Aizenberg

Aphidinae: Macrosiphini

Ten species of narrow-bodied wax-dusted with very short siphunculi, living without host alternation on Gramineae/Poaceae, usually rolling or otherwise distorting and/or discoloring the leaves.  The group is possibly entirely of palaearctic origin but was little known until D.  tritici was described from Colorado, USA, in 1911, and there were serious outbreaks of  D. noxia on barley in Russia in 1912.  Subgenus Holcaphis differs from Diuraphis s.str. by lacking a process on abdominal tergite 8.  Accounts are available for north-west Europe (Heie 1992, Blackman 2010) and China (Zhang et al. 1991b), and G.L. Miller et al. (2005) reviewed the genus. Kovalev et al. (1991) provided a key to apterae, and Puterka et al. (2010) reviewed and illustrated the species in the Rocky Mountain region of the USA.

Diuraphis (Holcaphis) agrostidis (Muddathir)    Apterae are yellowish green, densely wax powdered, with head and appendages blackish; BL 1.7-2.6 mm. In unfurled leaves of Agrostis spp., especially A. stolonifera. In Europe and west Siberia (Stekolshchikov et al. 2008a). Monoecious holocyclic with oviparae and apterous males in late September-October (original description). 2n=12.

Diuraphis (Holcaphis) bromicola (Hille Ris Lambers)    Colour of apterae is undescribed, probably as oviparae which are pale green, powdered with grey wax, with head darker; BL 1.6-2.0 mm. On Bromus spp., especially B. inermis, in eastern Europe (Germany, Russia). Monoecious holocyclic with oviparae and apterous males in September-October (original description). The aptera was described for the first time by G.L. Miller et al. (2005).

Diuraphis (Holcaphis) calamagrostis (Ossiannilson)    Apterae are pale yellowish with dark head and appendages, white wax-powdered; BL 1.5-2.5 mm. In swollen leaf sheaths and longitudinally-rolled leaves of Calamagrostis spp. In northern Europe (Sweden, Finland, Poland, Lithuania). Samples have also been collected from two Carex spp. (Heie 1992), although these were probably vagrants. Monoecious holocyclic with oviparae in early September (original description).

Diuraphis (Holcaphis)  frequens (Walker)    Apterae are pale green, wax-powdered, with head darker, and blackish appendages; BL 1.3‑2.1 mm. Usually colonising Elymus repens, bunching the still unfurled leaves and stopping growth.  Also recorded from Echinochloa, Hordeum, Leymus, Lolium and Triticum.  Widespread in Europe, Asia (Turkey, Mongolia) and North America.  Monoecious holocyclic on Elymus with oviparae and apterous males in October (Hille Ris Lambers 1939b). Diuraphis (Holcaphis) elymophila G. Zhang 1991 is a synonym (G.L. Miller et al. 2005). 2n = 14.

Diuraphis (Holcaphis) holci (Hille Ris Lambers)    Apterae are green, wax-powdered, with dark green head and appendages; BL 1.3-2.1 mm. Usual hosts are Holcus spp., but there are also records from Agropyron, Hordeum and Poa. In Europe and North America. Monoecious holocyclic on Holcus, with oviparae in October. 2n=14.

Diuraphis mexicana (McVicar Baker)    Apterae are pale yellow-brown to pale green with pale brown head and appendages, wax-powdered; BL 1.4-2.0 mm. In rolled leaves of Bromus spp., and also found on Dactylis glomerata (Richards 1959, as Brachycolus nodulus). In Mexico, USA (Colorado, New Mexico) and Canada (British Columbia). Peña-Martinez & Vera-Castello (1998) and Puterka et al. (2010) compared this species with D. noxia. Mainly anholocyclic in Mexico, perhaps monoecious holocyclic at higher latitudes. 2n=8*.

Diuraphis muehlei (Börner)    Apterae are dirty yellowish, wax-powdered, with brownish head and appendages; BL 1.3-2.0 mm. In rolled, yellowed leaves of Phleum pratense. Widely distributed in Europe. Monoecious holocyclic with apterous males (original description), oviparae occurring in October (BMNH collection, paratypes, leg. C. Börner). This species was synonymised with D. noxia by G.L. Miller et al. (2005), but is morphologically and biologically distinct.

Diuraphis noxia (Mordvilko ex Kurdjumov)  Russian Wheat Aphid   Plate 13e   Apterae are pale yellow- green or grey‑green lightly dusted with white wax powder; BL 1.4‑2.3 mm. On grasses and cereals (Agropyron, Anisantha, Andropogon, Bromus, Elymus, Hordeum, Phleum, Triticum; see Kindler & Springer 1989). Very injurious to wheat and barley; infested leaves are rolled into tubes and desiccated, and infested ears become bent. Of palaearctic origin, now widespread; southern Europe, Middle East, Central Asia, Pakistan, North Africa, Kenya, South Africa, Chile, Argentina, North America (first record 1986) and South America (Chile 1988, Argentina 1992; Clua et al. 2004). Liu et al. (2010) provided DNA evidence of at least two separate introductions into the USA. Monoecious holocyclic (with apterous males) in cold temperate parts of Asia, probably mainly or entirely anholocyclic in warmer regions, but some introduced populations have the ability to reproduce sexually where environmental conditions permit (e.g. in Argentina, Ricci et al. 2011, and USA, Puterka et al. 2012). Lengthy periods of cold in winter caused heavy mortality of anholocyclic populations in the Czech Republic (Havelka et al. 2015). There is a large literature; see Botha (2013) for a recent review. Puterka et al. (1993) studied worldwide genetic variation, B. Zhang et al. (2012) studied microsatellite and mitochondrial DNA variation in holocyclic populations in north-west China, and B. Zhang et al. (2014) used a multigenome approach to map the putative invasion route of one clone from the Middle East via Africa to the New World. Now that sexual reproduction is known to occur in US populations the pervasive use of the term ”biotype” in the economic literature about this species should be reconsidered, as genetic recombination has the potential to generate an unlimited number of “biotypes” (Puterka et al. 2014, 2015). The complete mitochondrial genome has been sequenced (Bo Zhang et al. 2013; de Jager et al. 2014). Earlier Russian language literature was reviewed by Kovalev et al. (1991). 2n = 10.

Diuraphis (Holcaphis)  tritici (Gillette) (= Diuraphis (Holcaphis) agropyronophaga Zhang; synonymy by Eastop & Blackman 2005)    Apterae are pale yellowish‑green with a powdery white coating of wax; BL 1.1‑2.2 mm. In curled leaves of grasses, especially Agropyron occidentale and Elymus spp., and also sometimes on Triticum.  Described from North America (Colorado, Montana, Illinois); also in Peru (Wille 1940) and east Asia (China, Tibet, Mongolia), where it possibly originated (Zhang et al. 1991b, as D. agropyronophaga). Zhang et al. (1991b) noted that  it was able to colonise wheat (Triticum sp.) in fields treated with 2,4-D herbicide, because this curls wheat leaves and thus provides a suitable microclimate for the aphid. Monoecious holocyclic with oviparae and apterous males in October (Palmer 1952).  See Parker (1916) for an early account of the biology.

 

Doraphis Matsumura & Hori

Hormaphidinae: Hormaphidini

 

A genus for one or two poplar-feeding species with no obvious close relatives, the large aleyrodiform apterae of which were originally described as a scale insect.  A cladistic study (Chen et al. 2014) has placed it in a basal position in the Hormaphidini. Alatae have an unbranched media in the forewing and a single oblique vein in the hind wing.  Tao (1969) gave an account of the genus (in Chinese).

Doraphis populi (Maskell)  Apterae dark, flattened, scale-like, almost circular, secreting wax from a ring of marginal glands; BL 2.2-3.0 mm. On bark of Populus spp. (including ciliata, maximowiczii, sieboldii).  Alatae are purplish-brown, with rather short 4- or 5-segmented and heavily sensoriated antennae; BL 2.2-2.6 mm (Matsumura & Hori 1929).  Immature alatae form groups on the undersides of the leaves (Takahashi 1931).  In Japan, China, Korea, Siberia and India.  Anholocyclic populations on P. tremulae in the region of Vladivostock have only two parthenogenetic generations a year, one apterous and one alate (Mordvilko 1929, as Paracerataphis tremulae; made a subspecies of populi by Matsumura & Hori 1929).  Elsewhere the life cycle is unknown.  Ceratoglyphina populisucta Zhang & Zhong (1985) is possibly a previously undescribed morph of this species.

 

Drepanaphis Del Guercio

Drepanosiphinae

 

Active, rather robust alate aphids on leaves of nearctic Aceraceae.  Apparently related to the oriental genus Yamatocallis, and with similar flask-shaped siphunculi, but these are always  without any subapical reticulate ornamentation.  Dorsal abdominal tubercles are variably developed on abdominal tergites 1-4 (usually largest on 3), and these are often conspicuously pigmented.  Holocyclic, with males and all viviparous females alate, oviparae apterous.  Trioxys ameracis is a specialised parasitoid.  Smith & Dillery (1968) revised the genus in a very accessible publication, so only a summary treatment is provided here, with emphasis on the recognition of the living aphids.

Drepanaphis acerifoliae (Thomas)  (Plate 7b; fig. 26F)  Alatae have head and thorax reddish-brown, abdomen pale with dorsal tubercles dark at least on abdominal tergites 3-4, dark siphunculi, and wing veins distinctly dark-bordered.  There are three longitudinal white wax stripes on head and pronotum, white wax markings on meso- and metathorax and anterior abdomen, and a covering of wax on abdomen posterior to siphunculi;  BL 1.9-2.3 mm.  On Acer saccharinum and A. rubrum, occasionally on A. saccharum and other maples. Apterous oviparae and alate males occur in September-October (Hottes & Frison 1931). Common and widely distributed in North America, and introduced (on A. saccharinum) into Europe - Italy, Spain; Pérez Hidalgo et al. (2008) and Hungary (Ripka 2010).  2n=38.

Drepanaphis carolinensis Smith   (fig. 26E)  Alatae similar to acerifoliae in life, but the wing veins are not dark-bordered; BL 1.6-2.2 mm.  On Acer saccharum, and occasionally on A. rubrum.  Widespread in North America east of Rocky Mountains.

Drepanaphis choanotricha Smith & Dillery   Alatae black with pale legs.  Head and pronotum striped with white wax, meso- and metanotum and dorsal abdomen with wax dots, denser posteriorly; BL 1.1-1.7 mm.  On low branches of Acer saccharum (always within 50 cm of ground), in Illinois and North Carolina, USA.

Drepanaphis granovskyi Smith & Knowlton   (fig. 26D)   Alatae almost entirely unpigmented, without conspicuous wax; BL 1.2-2.0 mm.  On Acer grandidentatum in Idaho and Utah, USA.

Drepanaphis idahoensis Smith & Dillery  (fig. 26B)   Alatae entirely covered with white wax except for a dark, U-shaped mark between the dorsal tubercles an abdominal tergite 3 and the siphunculi.  Legs pale, BL 1.4-2.0 mm.  On Acer grandidentatum in western USA. For colour photographs of alate vivipara, ovipara and male see aphidtrek.org.

Drepanaphis kanzensis Smith   (fig. 26L)  Alatae yellowish-white, with a profuse covering of white wax, except on thoracic lobes and the prominent dark  tubercles on abdominal tergite 3.  Legs pale, wing veins not dark-bordered.  BL 1.2-2.0 mm.  On Acer saccharum in North America east of the Rocky Mountains, but with a more northerly range than D. carolinensis.

Drepanaphis keshenae Granovsky   (fig. 26H)  Alatae have body profusely covered with white wax, through which project the large dark tubercles on abdominal tergite 3. Wing veins distinctly bordered with fuscous.  BL 2.0-2.5 mm.  On Acer saccharum in eastern USA and Ontario, Canada.

Drepanaphis knowltoni Smith & Dillery   (fig. 26A)  Alatae white with wax, very like idahoensis except for dark legs; BL 1.6-2.4 mm.  On Acer grandidentatum in Idaho and Utah USA, and also recorded from other Acer spp. in eastern USA (Smith & Dillery 1968).

Drepanaphis monelli Davis   Alatae have body entirely covered with white wax powder except for dark thoracic lobes and a brownish-yellow U-shaped line between the tubercles on abdominal tergite 3 and the dark siphunculi; BL 1.9-2.3 mm.  On Aesculus glabra, especially on yellowed leaves (the only Drepanaphis species not associated with Acer).  In eastern USA and Canada (New Brunswick).

Drepanaphis nigricans Smith   Alatae have black body with 2-3 longitudinal wax stripes on head and pronotum and small spots or flecks of wax on thorax and abdomen.  Legs pale, wings clear except for dark spots at ends of veins.  On Acer rubrum, especially on terminals of rapidly growing shoots (water-shoots), in eastern USA, west to Michigan and Tennessee, and Canada (New Brunswick).  Closely related to, and possibly conspecific with, D. tissoti.

Drepanaphis parva Smith   (fig. 26K)  Alatae have reddish-brown head and thorax, the head and pronotum having 5 longitudinal white wax stripes, and greenish-grey-brown abdomen with white posterior tergites and longitudinal rows of white wax dots; BL 1.4-2.6 mm.  Legs pale, fore wings somewhat cloudy with veins diffusely bordered.  On Acer rubrum in eastern USA and Canada.

Drepanaphis sabrinae Miller   (fig. 26I)  Alatae are orange-brown to orange-yellow with small flecks of white wax, and 2-3 longitudinal stripes on pronotum; BL 2.0-2.6 mm.  On Acer saccharum in eastern USA, west to Minnesota and Kansas.

Drepanaphis saccharini Smith & Dillery  (fig. 26G)  Alatae very similar in life to parva (q.v.), but with clear wings; BL 1.5-2.3 mm.  On Acer saccharinum (but not cultivated varieties) in eastern and mid-western USA; only occasionally on A. rubrum.

Drepanaphis simpsoni Smith   (fig. 26J)  Alatae have reddish-brown head and thorax with short fine longitudinal white wax stripes, and light yellow to white abdomen with the finger-like tubercles on abdominal tergite 1 reddish-brown; BL 1.5-2.3 mm.  On Acer saccharum in eastern USA as far south as North Carolina, and in eastern Canada.  D. pallida Richards seems to be a pale form of D. simpsoni.  2n=30*.

Drepanaphis spicata Smith   Alatae in life closely resemble those of knowltoni (q.v.), but are larger and have fore femora much paler ventrally than dorsally; BL 2.4-3.2 mm.  On Acer spicatum in eastern USA (mountainous areas west to Tennessee and south to North Carolina) and Canada (New Brunswick).  Records from A. grandidentatum in western USA (eg. Palmer 1952) are probably referable to D. knowltoni and/or D. idahoensis.

Drepanaphis tissoti Smith   Alatae black with pale legs, strongly resembling nigricans (q.v.); BL 1.3-2.0 mm.  On Acer rubrum in eastern USA, usually on growing terminals of young shoots (water-shoots); one record from A. saccharum.  Doubtfully distinct from D. nigricans.

Drepanaphis utahensis Knowlton & Smith   (fig. 26C) Alatae yellow, with head and pronotum edged with black, thoracic lobes brown, tubercles on abdominal tergite 3 dark, and entire body frosted with white wax; BL 1.5-2.0 mm.  A very pale form occurs in midsummer.  On Acer grandidentatum in western USA (a record from A. saccharum in Nova Scotia, Canada, seems likely to be another species).  2n=30.

       

Drepanosiphoniella Davatchi, Hille Ris Lambers &

 Remaudière

Drepanosiphinae

 

A genus for three palaearctic species related to the nearctic genus Drepanaphis, but the primary rhinaria have no fringe of hairs and dorsal body hairs are long and stiff as in Chaitophorinae.  All three species are generally found at altitudes more than 1000 m.a.sl. and primarily assocated with A. monspessulanum.  Both apterous and alate viviparae occur. Males are apterous. Wieczorek et al. (2015) fully revised the genus and provided keys to all available morphs.

Drepanosiphoniella aceris Davatchi, Hille Ris Lambers & Remaudière   (fig. 19B)  Apterae grey to dark brown, lightly dusted with wax powder; BL 1.7-2.5 mm.  Femora are brown conspicuously mottled with darker-brown, and siphunculi are dark brown.  The siphunculi of alatae are often much longer than those of apterae, and are usually markedly curved; alatae also have a very short wide pterostigma with two black patches surrounding a pale central area.  On undersides of leaves of Acer spp. especially A. monspessulanum and its subspecies (cinerascens, microphyllum), and also recorded from A. velutinum, in south-west Asia (Iran, Iraq, Turkey).  Holocyclic, with oviparae and apterous males in October; Wieczorek et al. (2015) described all morphs. A subspecies, D. aceris ssp. caucasica, was described from Acer ibericum in Armenia (Mamontova 1982), and redescribed by Wieczorek et al. (2015); it has no wax powder, long antennal hairs and straight siphunculi of similar length in both apterae and alatae.   

Drepanosiphoniella fugans Remaudière & Leclant   Apterae are dull grey to light brown, lightly dusted with wax powder; BL 2.0-2.5 mm. General appearance is similar to D. aceris, and it was originally described as a subspecies, but given separate species status by Wieczorek et al. (2015). On undersides of leaves of Acer monspessulanum (incl. ssp. microphyllum) at high altitudes in southern France, Spain and Italy, and also in the Middle east (Lebanon). Holocyclic, with oviparae and apterous males in October (originally these were recorded as collected from A. campestre, but there is some doubt about this - see Wieczorek et al. 2015, who provided descriptions of all morphs). Stary & Mackauer (1971) described a new species of parasitoid from this aphid.

Drepanosiphoniella remaudierei Wieczorek    Colour of apterae in life unknown; BL 2.2-2.9 mm. On Acer monspessulanum at c.1700 m.a.sl. in Morocco (Wieczorek et al. 2015). Sexual morphs and life cycle are unknown.

 

Drepanosiphum Koch

Drepanosiphinae

 

A genus for nine species of medium-sized to large elongate aphids with long antennae and tubular siphunculi, all associated with Acer.  All viviparae are alate, forming "spaced-out aggregations" on undersides of leaves in shade (Kennedy et al. 1967).  Males are alate, oviparae are large and apterous with the end of the abdomen extended like an ovipositor.  The genus has several specialised parasitoids; Discritulus planiceps (which pupate like Praon under the aphid but in a flat disc rather than a tent), Monoctonus pseudoplatani and Trioxys cirsii (which emerge from pale mummies),and Aphelinus thomsoni (which emerge from black mummies).  Stroyan (1977) provided a generic diagnosis and keys to British species, and Heie (1982) gave an account of the species of north-west Europe. Wieczorek et al. (2016b) provided descriptions, morphometric data and keys to all known sexual morphs (seven species), and discussed taxonomic status and host relationships of all the species.

Drepanosiphum acerinum (Walker)  Alatae whitish-green or yellow, with thoracic lobes darker, a conspicuous brown-black spot in front of the base of each siphunculus, siphunculi either entirely or distally dark, and often a short brown bar or bars on abdominal tergites 4-5 only (see influentialpoints.com/Gallery); BL 2.1-3.3 mm.  On Acer pseudoplatanus, under lower leaves, usually in shade.  In Europe, except the Baltic region.  Sexuales occur in September-October.  Records of D. acerinum from other Acer spp. are probably all referable to other Drepanosiphum species, eg. aceris, caucasicum.  Sutakova (1984) studied associations with microbial organisms in relation to transmission of maple leaf perforation virus, and Keller (1987) recorded a fungal infection.

Drepanosiphum aceris Koch   (fig. 21A)  Alatae yellowish or pale whitish-green with dark thoracic lobes, siphunculi brown with black apices, and often with rather narrow cross-bands on abdominal tergites 4 and 5, plus lateral spots in front of bases of siphunculi, and dark spots at wing-tips (see influentialpoints.com/Gallery); BL 2.7-4.2 mm.  On Acer campestre, usually under leaves near ground, in Europe and east to the Caucasus and Georgia (Barjadze et al. 2010a).  Sexuales occur in September-November.

Drepanosiphum braggii Gillette   (fig. 21B)  Alatae pale yellow to greenish with yellow-brown thoracic lobes and dark-tipped siphunculi; BL 2.7-3.6 mm.  On Acer negundo, undersides of leaves, in western USA and Mexico (BMNH collection, leg. R. Peña).  Sexuales occur in September-November (Palmer 1952).  2n=30.

Drepanosiphum caucasicum Dzhibladze   Alatae pale yellow with dark dorsal abdominal markings on abdominal tergites 4 and 5, or 5 only, very like D. acerinum but with siphunculi whitish transparent, only the extreme apices being sometimes slightly dusky; BL 2.4-2.9 mm.  On Acer trautvetteri in Georgia (Dzhidbladze 1962), and on A. tauricolum in Turkey (Remaudière et al. 2006). D. asiaticum Narzikulov, described from A. regelii and A. turkestanicum in Tajikistan as a subspecies of D. platanoidis (Narzikulov 1968), seems likely to be a synonym.

Drepanosiphum dixoni Hille Ris Lambers   (fig. 22A)  Alatae greenish-ivory with brown head, dark brown to black thoracic lobes, pale brown to black siphunculi, and two extensive blackish sometimes coalescing cross-bars on abdominal tergites 4-5, plus dark sclerites in front of siphuncular bases; BL 2.5-3.3 mm.  Brachypterous alatae are common in midsummer (Dixon 1972).  On Acer campestre, under lower leaves in shade, sometimes on water-shoots growing from lower parts of trunk.  Recorded from England, Netherlands, Yugoslavia and northern Italy (van Harten & Coceano 1981).  Sexuales in October (Hille Ris Lambers 1971).

Drepanosiphum iranicum Hille Ris Lambers   (fig. 22B)  Colour of alata in life not recorded, probably pale green, with variably pigmented head and thorax, and with a conspicuous black spot in front of base of each siphunculus (even in pale specimens); BL 3.1-4.1 mm.  On Acer cappadocicum and A. velutinum in Iran (Hille Ris Lambers 1971).  Oviparae in October (BMNH collection, leg. S. Hodjat); both sexual morphs were described by Wieczorek et al. (2016b).  Described as a subspecies of, and closely related to, D. platanoidis, but the constancy of abdominal pigmentation and the different host relationship indicate that this is a good species.  2n=30 (Blackman & Eastop 1994).

Drepanosiphum latifemorum Wieczorek, Junkiert & Kanturski  Colour in life unknown, probably pale green without any dark pigmentation; BL 2.4-3.2 mm. On Acer monspessulanum in Morocco; described from specimens collected in 1938, and not recorded since (Wieczorek et al. 2016b).

Drepanosiphum oregonense Granovsky   (fig. 22C)  Alatae have orange-brown thoracic lobes and pale green abdomen; BL 1.9-2.6 mm.  Darker specimens may have a blackish-brown mesosternum and dark spots in front of bases of the rather thick, swollen, dark-tipped siphunculi.  The black stripe on the fore femur and black basal section of ANT III are distinctive.  On various Acer spp. in southern, central and eastern Europe, Mediterranean region, south-west Asia, Kazakhstan (Kadyrbekov 2014d), India (Chakrabarti 1988), Pakistan (Naumann-Etienne & Remaudière 1995). It was introduced to western USA (California, Oregon) some time before 1939, and much more recently to Argentina (Ortego et al. 2014).  Sexuales occur in October in California, but oviparae were collected in July in Germany (BMNH collection, leg. W. Quednau).  Michelena Saval & Gonzalez Funes (1988b) recorded a parasitoid in Spain.

Drepanosiphum platanoidis (Schrank)  Plate 7a, and fig. 6  Alatae have yellow-brown head and thorax with darker brown markings, and a pale green abdomen with or without variably developed dark cross-bars (but if present these are never restricted to abdominal tergites 4-5) (see influentialpoints.com/Gallery); BL 3.2-4.3 mm.  On undersides of leaves of Acer pseudoplatanus; recorded from many other Acer spp., but these are apparently only visited on a casual basis.  It seems to be common on sycamores wherever they are grown in Europe, central Asia, north Africa, Australia, New Zealand, USA and Canada (British Columbia).  Sexuales occur in September-November.  There has been much work on this well-known aphid, including studies of its population dynamics (eg. Chambers et al. 1985, Wellings et al. 1985), distribution on leaves (Dixon 1976), seasonal variation (Dixon 1972, 1974, 1975), function of siphunculi (Dixon & Stewart 1975), alarm pheromone (Fan et al. 2011), effects on wood formation (Dixon 1971a), flight behaviour (Dixon & Mercer 1983), sexual morph determination (Dixon 1971b), effects of predation by ants (Warrington & Whittaker 1985) and chemical control (Parry et al. 1989). The male genitalia were described and illustrated by Wieczorek et al. (2011). 2n=30.

 

Durocapillata Knowlton

Aphidinae: Macrosiphini

One western North American species of uncertain affinities with stiff capitate hairs and distinctive siphuncular morphology.

Durocapillata utahensis Knowlton  Plate 15b  (Fig.21i)   Apterae are shining yellow-green with black tips to antennae, legs and siphunculi; BL 1.3-1.5 mm. Curling leaves and distorting young growth of Chrysothamnus viscidiflorus in Idaho and Utah, USA. Monoecious holocyclic with oviparae and apterous males in late September-October (Palmer, 1952).

    

Dysaphis Börner

Aphidinae: Macrosiphini

 

A palaearctic genus of about 110 species, many of which are heteroecious with spring colonies distorting and discolouring the leaves of their primary hosts in the Pyroideae (Amelanchier, Crataegus, Malus, Pyrus, Cotoneaster, Sorbus), before migrating to secondary hosts, especially in Umbelliferae/Apiaceae but also several other families (Campanulaceae, Compositae, Valerianaceae, Plantaginaceae, Polygonaceae). There are also many species that are monoecious either on the original primary or secondary hosts and some that have adapted to entirely new hosts, e.g. Liliopsida/monocots.  On the primary host they are generally medium-sized, rather plump-bodied dark-legged aphids, greenish, bluish or pinkish grey in colour and covered in wax meal.  Characteristic features are the short, often helmet-shaped cauda and usual presence of spinal tubercles on head and posterior abdominal tergites.  Colonies are almost always attended by ants.  In Dysaphis s. str. on Malus, the fundatrix forms a small apical primary gall on the young leaf, but her progeny soon leave this to form and develop in marginal leaf-rolls.  Migration of many heteroecious species occurs in the second and/or third generation , so the spring colonies are short-lived and may comprise only the immediate descendants of the fundatrix; but in subgenus Pomaphis colonies may persist for several generations on the primary host.  On their secondary hosts they usually live concealed at stem bases inside sheathing leaves, where they are attended and sheltered by ants. Some species are monoecious holocyclic on herbaceous plants. In subgenus Pomaphis the secondary host apterae differ substantially from those on the primary host, to some extent resembling apterae of small Myzus. Shaposhnikov (1956, 1986, 1990) and Stroyan (1957, as Sappaphis; 1963, 1985) have published extensively on Dysaphis. Stroyan (1985) provided notes on several undescribed species on named host plants, most of which (indica, oreoselini, peucedani, selinumi, sharmai) have subsequently been described. Shaposhnikov & Moralev (1979) reviewed and keyed the Crataegus-feeding species, Stekolshchikov (2006) reviewed and keyed the species on Polygonaceae, and Stekolshchikov (2008) reviewed the species feeding on Liliopsida. Regional reviews are available for Fennoscandia and Denmark (Heie 1992), UK and Ireland (Blackman 2010), the Indian subcontinent (Chakrabarti & Medda 1993) and Japan (Sugimoto 2003). The common apple- and pear-feeding species are reviewed in Blackman & Eastop 2000, pp. 264-269. The numbers of secondary rhinaria given in the accounts below refer to alatae (and in some cases apterae) produced in colonies on secondary hosts.

Dysaphis acroptilidis Daniyarova    Apterae are greenish brown, wax-dusted; BL c.2.6 mm. Alatae have secondary rhinaria distributed III 35-40, IV 18-20. On Acroptilon (= Rhaponticum) repens in Tajikistan.

Dysaphis affinis (Mordvilko)  Living without host alternation in red leaf galls on Malus spp. in Asia (west Siberia, Turkey, Armenia, Georgia, Kazakhstan, Tajikistan, Pakistan, north-west China), and also recorded from Latvia (probably introduced on seedlings or root-stocks).  Injurious to apple tree nurseries, rolling and blistering the leaves (Shaposhnikov 1964).  The second generation includes many alatae as well as apterae. Naumann-Etienne & Remaudière (1995) found galls containing apterous and alatiform viviparae Pakistan (Passu) in September, although their population differed in certain respects from typical affinis. Shaposhnikov (1986) reported sexuales in Russia in October. 2n=12.

Dysaphis allii Daniyarova    Apterae are pale green, dusted with greyish wax; BL c.1.7 mm. In an ant-attended colony on roots and onions of a wild Allium sp. in Kazakhstan. Stekolshchikov (2008) redescribed the apterae (possibly they are fundatrices) of this species, only known from the original collection.

Dysaphis angelicae (Koch)  Forming cherry-red curled-leaf galls on Crataegus.  Fundatrix is greenish-grey.  All or nearly all of the second generation are alatae, greyish-pink with black dorsal markings and secondary rhinaria distributed III 58-79, IV 19-32, V 2-17, and migrate to Angelica spp., forming ant-attended colonies on lower leaf bases (Stroyan 1963). Apterae on Angelica are yellowish grey or greenish grey, wax-powdered; BL 1.7-2.2 mm (see influentialpoints.com/Gallery). Alatae produced on Angelica have secondary rhinaria distributed III 46-87, IV 15-37, V 0-13. In Europe (England, France, Netherlands, Sweden, Germany, Hungary, Belarus). Records from North America are uncertain (Footit et al. 2006). (Note: spring migrant alatae of a closely-related species that migrates to Laserpitium, D. laserpitii (q.v.), may key out to angelicae.) 2n=12.

Dysaphis angelicophaga Zhang, Chen, Zhong & Li    Apterae are ivory-white, wax-dusted; BL c.1.8 mm. Alatae have secondary rhinaria distributed III 61-67, IV 7-11, V 1-2. On roots of Angelica sinensis, ant-attended (and stated to be an important pest). In Gansu Province, China (Xinglong mountains, 2170m; G. Zhang 1999). Not a typical Dysaphis; the hairy cauda suggests that it may be placed in the wrong genus.

Dysaphis (Pomaphis) anisoidis Barbagallo & Stroyan    Apterae are shining black dorsally and pruinose ventrally; BL 1.0-1.8 mm. At base of stem and on root collar of Pimpinella anisoides (Barbagallo & Stroyan 1982), ant-attended. In France and Italy. Presumably monoecious holocyclic; Stekolshchikov (1998) distinguished populations in Armenia on Pimpinella tripartita and P. rhodantha as a subspecies, D. anisoidis ssp. nairi, collected fundatrices from this plant, and obtained oviparae and alate males in laboratory cultures.

Dysaphis annulata (Börner)    Only known from the holotype aptera, described as from Ranunculus, but belonging to the brancoi group which is usually associated with Valeriana, so the original host record may have been in error (see Stroyan 1963).

Dysaphis anthrisci (Börner)  A complex of sibling species and subspecies rolls the margins of the leaves of apple in spring in Europe, the separate taxa being only reliably distinguished by their life cycles and/or secondary host relationships.  D. anthrisci s. str. occurs throughout Europe and migrates in the second generation to form ant-attended colonies at the leaf bases of Anthriscus sylvestris, gynoparae and males returning to Malus in October. Spring migrant alatae have secondary rhinaria distributed III 32-61, IV 4-23, V 0-5. Apterae on Anthriscus are bluish grey coated with wax powder; BL 1.9-2.3 mm. Alatae produced on Anthriscus have secondary rhinaria distributed III 42-67, IV 13-26, V 0-7. Populations migrating from Malus to Anthriscus nemorosa in Russia are regarded as a subspecies, D. anthrisci ssp. majkopica (Shaposhnikov 1965), although aphids from A. nemorosa in Armenia show intermediate morphology (Stekolshchikov & Lobanov 2002). A closely-related species, D. ussuriensis,  lives all-year-round on Anthriscus aemula (= nemorosa) in east Asia and will key to anthrisci in the Anthriscus aphid key; it can only be distinguished by use of  a complex discriminant function (Shaposhnikov & Stekolshchikov 1989). There are other very closely related species in Europe and Asia, differing more in biology than in morphology: D. chaerophylli, D. chaerophyllina, D. brachycyclica and D. caucasica going to Chaerophyllum spp.; D. bunii to Bunium spp.; D. physocaulis to Physocaulis (=Myrrhoides) nodosus; and D. flava and D. zini with unknown secondary hosts.  D. devecta and D. radicola are also very closely related. This group was studied by Shaposhnikov (1956, 1959, 1965, 1990), discussed further by Stroyan (1985), and more recently reviewed by Stekolshchikov & Lobanov (2002). Other members of the anthrisci group also occur in east Asia, e.g. in Korea on Malus baccata (see Stroyan 1985), and China (Zhang & Zhong 1984).  2n=12 (incl. ssp. majkopica).

Dysaphis apiifolia (Theobald)    Apterae are yellowish to greenish grey, wax-powdered; BL 1.6-2.4 mm. They form dense colonies at the leaf bases of various Umbelliferae/Apiaceae (e.g. Anethum, Ammi, Apium, Conium, Foeniculum, Laserpitium, Levisticum, Petroselinum, Peucedanum, Ptychotis, Smyrnium), attended and sheltered by ants. Alatae produced on these hosts have secondary rhinaria distributed III 45-82, IV 16-34, V 0-7. Populations in northern Europe have a sexual phase on Crataegus (“ssp. petroselini”). Cherry-red to crimson curled-leaf galls of this form occur on Crataegus in spring. The fundatrix is greenish-grey, and all its progeny are alate, with secondary rhinaria distributed III 58-99, IV 19-42, V 0-13; the immature alatae are greenish. This form may preferentially utilise Petroselinum as secondary host (Stroyan 1963). The more polyphagous, widely-distributed form (“D. apiifolia s.str.”) occurs in Middle East, Central Asia, Africa, Mauritius, Australia, North and South America, and is a world-wide pest of celery and parsley (Blackman & Eastop 2000). It is mostly anholocyclic, but red galls agreeing with apiifolia s. str. have been found on Crataegus azarolus in Italy (Stroyan 1985), indicating that this form may also have at least a partial host alternation.  Shaposhnikov (1987) provided a key to the species of the apiifolia/petroselini group. 2n=12.

Dysaphis (Pomaphis) ariae (Börner)  In leaf-roll galls on Sorbus aria in alpine Europe (Austria, France).  Apterous fundatrigeniae were straw-white (Börner 1950).  Stroyan (1963) considered that this might be a starvation form of plantaginea, but it appears to be a distinct, shorter-haired species (see also Shaposhnikov 1963).  It is possible that D. gallica on Cymbalaria muralis is the secondary host form of D. ariae, although host transfers were so far unsuccessful (Stroyan 1985; see also under D. plantaginea).

Dysaphis armeniaca Shaposhnikov   Rolling and reddening the lateral edges of the leaves of Malus domestica and M. orientalis in Armenia.  Apterae are greenish-pinkish brown, slightly shiny on dorsal surface; BL c. 1.5 mm. Immatures are pinkish-grey (cf. D. foeniculi malidauci).  Almost all the second generation are alatae, migrating to an unknown secondary host; of numerous attempted transfers to various Umbelliferae/Apiaceae only Ligusticum alatum elicited a settling response, so the true secondary host may be a related species (Shaposhnikov 1986).

Dysaphis atina A.K.Ghosh, R.C. Basu & Raychaudhuri    Colour of apterae in life is not recorded; BL 1.8-2.6 mm. On Cirsium spp. and Carduus spp. in northern India (Himachal Pradesh, Uttar Pradesh, Jammu and Kashmir), in Pakistan (Naumann-Etienne & Remaudière 1995), and also in Iran on Cirsium arvense (BMNH collection, leg. R. van den Bosch). The alata was described by Chakrabarti & Medda (1993).

Dysaphis (Pomaphis) aucupariae (Buckton)  Spring generations live in conspicuous rolled or twisted-leaf galls on Sorbus torminalis, which are initially pale yellow-green, becoming tinged with red and later golden yellow.  Despite Buckton's name for this aphid, it occurs on Sorbus aucuparia only rarely.  Fundatrices  (BL 2.7-3.1 mm) and their apterous progeny (BL 2.2-2.5 mm) are mottled greyish and blue-green underneath a thick coat of mealy white wax (see influentialpoints.com/Gallery). Migration (of alatae with secondary rhinaria distributed III 62-79, IV 24-33, V 1-7) occurs in the third and subsequent generations to Plantago spp., especially P. lanceolata (Stroyan 1957), where colonies occur on the undersides of leaves, in grooves between veins, attended by ants. Apterae on Plantago are pinkish or reddish yellow, and have siphunculi with dark tips and brownish patches around bases; BL 1.1-2.0 mm. Alatae produced on the secondary host have secondary rhinaria distributed III 42-68, IV 10-28, V 0-9. In Europe, east to Crimea and Caucasus, and the Azores. Also, presumably anholocyclic populations are found on Plantago in Australia (Carver & Hales 1983), New Zealand (Sunde 1984) and Argentina (Ortego 1998a), and alatae have also been trapped in Washington USA (BMNH collection; leg. S.H. Halbert).  2n=12.

Dysaphis bonomii (Hille Ris Lambers)    Apterae are pale to dull greyish green with dark dorsal bands or patches, wax-dusted (see influentialpoints.com/Gallery); BL 1.2-2.5 mm. The usually alatiform apterae have secondary rhinaria distributed III 0-35, IV 1-12, V 0-1, and alatae have them distributed III 27-43, IV 6-13, V 0-3 (III and IV sometimes fused in both morphs). On basal parts of Pastinaca sativa, attended by ants. In Europe (southern England, Sweden, Germany, Austria, Italy). Monoecious holocyclic with apterous males.

Dysaphis brachycyclica Shaposhnikov   Rolling leaves of Malus spp. in eastern Europe (North Caucasus, Crimea, Armenia) in spring, but the galls are yellow rather than red (cf. chaerophyllina, etc.).  All of the second generation are alatae, migrating in May to form ant-attended colonies on basal parts of Chaerophyllum bulbosum. Apterae on the secondary host are probably pinkish grey, wax-dusted; BL 2.6-3.0 mm. The return migration is in mid-July to August.  It is a member of the D. anthrisci species complex. Shaposhnikov (1990) provided a detailed comparison with the closely related D. chaerophyllina; the characters given to separate these two species in the key to Chaerophyllum aphids may not be reliable for all specimens and/or all populations. Stekolshchikov & Lobanov (2002) regard it as a subspecies of D. flava, which seems to differ only in general body size across all morphs..

Dysaphis brancoi (Börner)  In deep purplish-red leaf-roll galls on Malus domestica or M. orientalis in spring, migrating in second and third generations to Valeriana spp., where it lives in ant-attended colonies at bases of stems and on shallow roots. Apterae on the secondary host are dark grey or yellowish-greenish grey dusted with wax powder, imm. pale to pinkish grey; BL 1.9-2.5 mm. The usually alatiform apterae have secondary rhinaria distributed III 0-38, IV 0-30, V 0-8, and alatae have them distributed III 70-106, IV 28-48, V 2-9. Throughout Europe (except Iberian peninsula), and also recorded from Kazakhstan, India (Chakrabarti & Medda 1993), east Siberia (Pashchenko 1988a) and north-west China (G. Zhang 1999).  Populations in Britain tend to be longer-haired and have been separated as a subspecies (D. brancoi rogersoni; Stroyan 1963). The east Siberian populations also host-alternate between Malus and Valeriana but are even more distinct (D. brancoi ssp. malina; Shaposhnikov & Stekolshchikov 1994), and should perhaps have separate species status. In continental Europe there are a group of closely-related taxa that are mon. hol. on Valeriana (see D. leefmansi).

Dysaphis (Pomaphis) brevirostris (Börner)    Apterae are pale brown, with brown siphunculi; BL 1.2-1.6 mm. Alatae have secondary rhinaria distributed III 30-40, IV 8-12,V 4-6. On Campanula spp. and Jasiona montana, usually in moist situations such as rocky or shingly banks of streams (Stroyan 1966). In Europe (only known from UK, Germany and Slovakia). Monoecious holocyclic ; oviparae and alate males are produced in September. However, Prior (1971) found that a population from Jasione in Devon was anholocyclic. D. brevirostris is a member of the D. sorbi group; Barbagallo & Patti (1994) key the three European spp. on Campanulaceae.

Dysaphis bunii Shaposhnikov    Apterae were not described from life but are probably bluish or pinkish grey, wax-powdered, with dark dorsal abdominal cross-bands; BL c.2 mm.  On Bunium sp. in west Turkmenistan. Heteroecious holocyclic, with Malus pumila as primary host (Shaposhnikov 1956).  It is a member of the D. anthrisci group.

Dysaphis candicans (Passerini)    Apterae are whitish, wax-dusted; BL unknown. At base of stem and on root collar of Orobranche ramosa in Italy (Bologna). Not recorded since original description; see Stroyan (1963).

Dysaphis capsellae (Mordvilko ex Nevsky)    Colour of apterae in life is unknown; BL c.1.7 mm. On radical leaves of Capsella bursa-pastoris in Central Asia. A member of the difficult D. foeniculus group, of uncertain status; Stroyan (1985) noted that members of this group as well as other Dysaphis spp. may use Capsella as a “reserve host”. Shaposhnikov (1987a) subsequently provided a key with some discriminants, treating capsellae as a subspecies of foeniculus.

Dysaphis caucasica Shaposhnikov    Apterae are probably greyish pink, wax-dusted, BL not recorded. On a Chaerophyllum sp. (? C. aureum) in Russia (Transcaucasia). Life cycle is  unknown.

Dysaphis centaureae (Börner)    Appearance of apterae in life is not recorded; BL 1.4-1.6 mm. On roots and rhizomes of Centaurea spp. In central and eastern Europe (Austria, Czech Republic, Ukraine), with a record also from Finland (Åland; Albrecht 2010). Monoecious holocyclic according to Börner (1952).

Dysaphis cephalariae Narzikulov    Apterae are dirty green, with greyish-green wax pulverulence; BL c.1.9 mm. On rhizomes of Scabiosa soongorica in Tajikistan (Narzikulov & Daniyarova 1990).

Dysaphis cephalarioides Shaposhnikov    Apterae are dirty greenish brown, bluish grey or bluish green, without dorsal pulverulence; BL c.1.5 mm. On stem bases and root collars of Cephalaria spp. in southern Ukraine and Armenia. Monoecious holocyclic; oviparae and alate males were obtained on C. gigantea in September (Stekolshchikov 1998).

Dysaphis chaerophylli (Börner)  Causing leaf-roll galls on Malus in spring.  Second generation is almost entirely alate, migrating to found anht-attended colonies on roots, stems and leaf bases of Chaerophyllum spp. (aromaticum, hirsutum, temulum, but perhaps not bulbosum; Stroyan 1963).  Apterae on the secondary host are dull greyish pink, wax-dusted, BL 1.5-2.6 mm. Alatae have secondary rhinaria distributed III 47-73, IV 15-25, V 0-4. In west and central Europe (Austria, Czechoslovakia, England, Germany, Switzerland).  A member of the D. anthrisci species group.

Dysaphis chaerophyllina Shaposhnikov   Rolling and reddening the edges of the leaves of Malus domestica and M. orientalis in Russia (north Caucasus, Armenia, Crimea), migrating in second and third generations (May-June) to basal parts of Chaerophyllum maculatum and, in some areas, Ch. bulbosum. Apterae on the secondary host are presumably greyish pink, wax-dusted; BL 2.4-3.2 mm. Return migration from Ch. maculatum is in September-October.  Shaposhnikov (1990) provided a detailed comparison with the closely-related D. brachcyclica, and Stekolshchikov & Lobanov (1990) computed discriminant functions for all morphs of these two species.  A member of the D. anthrisci species complex. 2n=12.

Dysaphis cnidii Shaposhnikov & Stekolshchikov     Apterae are pale greenish grey, wax-dusted; BL 1.4-1.63 mm. In ant shelters at bases of lower leaves of Cnidium ajanense in east Siberia (Shaposhnikov & Stekolshchikov 1989). The life cycle is unknown. It is one of a group of species with long antennal hairs living on Umbelliferae/Apiaceae in east Asia, closely related to D. mordvilkoi and probably with Malus as primary host.

Dysaphis cousiniae Narzikulov    Apterae are wax-powdered, BL 2.2-3.1 mm. On basal parts of Cousinia spp. and Arctium lappa in Tajikistan (Stroyan 1985). Stroyan also identified specimens from Handelia trichophylla as this species, but possibly these were D. handeliae. A variant population was described as a subspecies, D. cousiniae minor Narzikulov.

Dysaphis crataegi (Kaltenbach)  Forming deep cherry-red curled-leaf galls on Crataegus spp. in spring.  The fundatrix is greenish-grey (see influentialpoints.com/Gallery), immature alatae are pinkish.  Second generation is nearly all alate, migrating to form dense ant-attended colonies on ground level parts of Umbelliferae/Apiaceae. Apterae on  secondary hosts are yellowish grey or greenish grey, wax-powdered; BL 1.4-2.5 mm. Alatae have secondary rhinaria distributed III 31-89, IV 4-30, V 0-4. The name D. crataegi is being applied to a complex of closely related species or subspecies with different primary and/or secondary host plant relationships (Stroyan 1963, 1985; Shaposhnikov & Moralev 1978).  In western Europe, D. crataegi s. str. preferentially colonises Daucus carota; D. crataegi ssp. kunzei (Börner) goes to Pastinaca; and D. crataegi ssp. aethusae (Börner) to Aethusa cynapium and Torilis japonica.  In south-west and central Asia, D. crataegi ssp. pallida Shaposhnikov & Moralev forms typical red galls containing greenish-white immatures (cf. incognita) on several Crataegus species, but its secondary host has not been established, and D. crataegi ssp. heraclei Shaposhnikov & Moralev migrates from Crataegus orientalis to Heracleum pastinacifolium.  In fact, all these taxa are part of a larger group of Crataegus-feeding and host-alternating species that includes angelicae, apiifolia, incognita, laserpitii, lauberti, ranunculi and virgata (subgenus Crataegaria of Shaposhnikov 1964).  These are treated separately here; however, the classification as either subspecies of crataegi or as separate species is somewhat arbitrary, not being based on any valid biological criteria.  Populations regarded as crataegi s. str. on Daucus occur in south-west and central Asia and USA, where they are probably anholocyclic on the secondary host plant.  2n=12 (on Daucus).

Dysaphis crithmi (Buckton)    Apterae are grey to greenish grey, wax-powdered; BL 1.7-2.2 mm. In sheathing leaf-bases of  Crithmum maritimum growing in sheltered situtations such as rock clefts, crevices in sea-walls, etc. (Stroyan 1963). In western and southern Europe, including Iberian peninsula and Mediterranean islands  (Corsica, Malta, Cyprus). Apparently it is completely anholocyclic.

Dysaphis deltoidei Shaposhnikov & Stekolshchikov    Galls are formed in spring on leaves of Malus mandshurica. The gall of the fundatrix is greenish yellow and formed by folding and swelling of the upper part of a leaf (Stekolshchikov 1998). Fundatrigeniae feed in a swollen marginal leaf roll, red with greenish yellow spots or entirely red. Emigrant alatae (BL c.1.9 mm) fly in late May-June to found colonies on roots and stem bases of Peucedanum deltoideum ( = P. terebinthaceum var. deltoideum). Apterae on the secondary host are pink-grey with grey-brown cross-bands, dusted with white wax; BL 1.6-1.9 mm. In east Siberia (southern Primorsk). Very like D. mordvilkoi; Stekolshchikov (1998) provided discriminants.

Dysaphis devecta (Walker)  Rolling and reddening the edges of the leaves of Malus in spring; the galls contain a mixture of normal bluish-grey wax-powdered apterae and dark green to reddish alatiform apterae with different degrees of pigmentation and sclerotisation of head and thorax.  Monoecious, with an abbreviated life cycle of only 3-4 generations; oviparae and alate males are produced within the galls (Hille Ris Lambers 1945, Forrest 1970).  Forrest & Dixon (1975) studied gall formation, Llewellyn & Hargreaves (1986) studied the energetics of gall-feeding, and Alston & Briggs (1977) investigated the genetic basis of resistance to this aphid in apple.  In Europe (not Scandinavia), and also recorded from China (Zhang et al. 1990). Foottit et al. (2006) and Holman (2009) also cite a record of this species from Canada by Quednau (1966), but this was perhaps a misidentification of Dysaphis tulipae (the material is now lost; F.W. Quednau, pers. comm. 2010).  Despite its unusual biology, comparable morphs are very similar to those of D. anthrisci and its relatives, and indicate a close relationship to that group.

Dysaphis emicis (Mimeur) ( =  Sappaphis albocinerea Hille Ris Lambers, = Dysaphis rheicola Daniyarova)    Apterae are grey-green with a dark grey patch on the prothorax and dark cross-bands on posterior abdominal tergites; BL 1.5-3.1 mm. On roots (sometimes deep down) of several genera of  Polygonaceae (Emex, Rheum, Rumex) in Europe, north Africa, Middle East, Central Asia, India, Pakistan, China, Brazil and Argentina. There are also records from Mesembryanthemum spp. (in Madeira; Ilharco 1984) and Pedicularis olgae (in Tajikistan; Stroyan 1985). Monoecious holocyclic on Rumex acetosella in Netherlands, with oviparae and apterous males in October (Hille Ris Lambers 1956a, as Sappaphis albocinerea), but with widely distributed anholocyclic populations in warmer climates. [This species may be synonymous with the eastern palaearctic D. rumecicola (q.v.), as Stekolshchikov (2006) demonstrated that there is an east-west cline in hair length and shape in populations of the emicis/rumecicola group. We retain both names here pending further investigation, in order to avoid hiding potentially useful information. Stekolshchikov used emicis as a subspecies name for anholocyclic populations of this group; however populations that overwinter anholocyclically are not necessarily genetically isolated from holocyclic ones, and anholocycly per se is not a valid reason for subspecies designation.]  

Dysaphis eremuri (Narzikulov)    Apterae are dark brown mottled with grey wax; BL 1.7-2.4 mm. In large colonies on root collars of Eremurus spp. in Central Asia (Afghanistan, Tajikistan, Kazakhstan). Stekolshchikov (2008) redescribed apterae and alatae of this species and compared it with D. alii and D. neostroyani. [The Uzbekistan population with a shorter rostrum described as a subspecies (D. eremuri ssp. baisunensis) by Narzikulov was synonymised with D. tulipae by Stekolshchikov (2008).]

Dysaphis ferulae (Nevsky)    Apterae are pale brown, with small dark brown spots on dorsal abdomen, dusted with grey wax; BL 2.0-2.8 mm. On Ferula spp. in Central Asia. Stroyan (1985) pointed out that this species must be closely related to D. foeniculus; Shaposhnikov (1987b) re-examined type specimens of D. ferulae and provided comparative data.

Dysaphis flava Shaposhnikov   Rolling lateral margins of leaves of Malus domestica in spring. Immatures in the leaf-rolls are pale yellow (Shaposhnikov 1956). Migration occurs in the second generation to found colonies on the root collars of Chaerophyllum bulbosum. Apterae on the secondary host are greyish green or brownish; BL 1.4-2.6 mm. The return migration occurs in mid- to late July as in the closely-related D. brachycyclica, which is regarded as a subspecies by Stekolshchikov & Lobanov (2002). In European Russia and Kazakhstan.   

Dysaphis foeniculus (Theobald)   Plate 11a   Apterae are grey-green lightly dusted with wax; BL 1.6-2.3 mm. In dense colonies on basal parts of various Umbelliferae/Apiaceae, including Anethum, Apium, Daucus, Foeniculum, Ferula, and also recorded occasionally from Rumex. In southern Europe, Mediterranean area, Middle East, Central Asia, India, Pakistan, Japan, Africa, Australia, New Zealand, North and South America. D. foeniculus is apparently anholocyclic everywhere. D. malidauci with a sexual phase on Malus and migrating to a similar range of secondary hosts in Central Asia is closely related and was originally described as a subspecies of D. foeniculus by Shaposhnikov (1987a). Stroyan (1985) and Shaposhnikov (1987a,b) reviewed the complex of species and/or subspecies around D. foeniculus, which includes capsellae, ferulae and pulverina. 2n=12.

Dysaphis (Pomaphis) gallica (Hille Ris Lambers)    Apterae are dark mottled blackish green, usually with a reddish tinge at bases of siphunculi (see influentialpoints.com/Gallery); BL 1.2-1.6 mm. Alatae have secondary rhinaria distributed III 54-92, IV 16-35, V 0-8. Described from Antirrhinum majus in Israel, and subsequently recorded from Cymbalaria  muralis in UK, France, Switzerland and Italy (Stroyan 1957a), and Veronica anagallis-aquatica in Sicily (Patti 1983). Heteroecious holocyclic in northern Germany; the primary host is unconfirmed, but in experimental transfers gynoparae produced numerous oviparae on Cotoneaster tomentosus (Müller 1968). An aphid in Pakistan (Baluchistan) forming crumpled-leaf galls similar to those of D. plantaginea has also been suggested as the possible primary host form of D. gallica (Naumann-Etienne & Remaudière 1995, and see under D. plantaginea).

Dysaphis handeliae Daniyarova    Apterae are pale brown, with wax; BL c. 2.0-2.1 mm. On basal parts of Handelia trichophylla in Central Asia (Tajikistan, Uzbekistan).

Dysaphis (Pomaphis) henrystroyani Barbagallo & Patti    Apterae are pale greenish, with antennae, legs, siphunculi and cauda mostly pale (immatures are salmon-pink); BL 1.4-1.8 mm. Alatae have secondary rhinaria distributed III 27-41, IV 8-14, V 4-8. On leaves, tender stems and flowers of Trachelium coeruleum, leaves becoming curled and twisted. Not ant-attended. Only known from Sicily, where it is possibly anholocyclic. Closely related to D. sorbi and D. brevirostris.

Dysaphis hirsutissima (Börner)    Apterae are bluish green with a dark dorsal pattern of paired sclerites and cross-bands; BL 1.5-2.3 mm. Alatae have secondary rhinaria distributed III 33-74, IV 19-36, V 1-11. On stem bases and in leaf sheaths of Anthriscus sylvestris, ant-attended. Recorded from UK, Denmark, Netherlands, Sweden, Finland, Switzerland and Italy. Monoecious holocyclic, with oviparae occurring in late September (Stroyan 1963). A single apterous male was collected in early October (BMNH collection, leg. V.F. Eastop). 2n=12.

Dysaphis hissarica Daniyarova    Apterae are greenish-yellow tinged with brown; BL c.2.4 mm. On Ferula ovina in Central Asia.

Dysaphis incognita Shaposhnikov & Moralev    Forming red curled-leaf galls on leaves of Crataegus curvisepala and Crataegus sp. in Crimea and Armenia, and also recorded from C. korolkowi in Kazakhstan (Kadyrbekov 2002d).  Second generation is all alate, migrating to an unknown host (transfer experiments to many potential hosts were unsuccessful - see Shaposhnikov & Moralev 1978).  Immatures in the Crataegus galls are yellowish or brownish pink (cf. D. crataegi ssp. pallida).  See also Stroyan (1985).

Dysaphis (Pomaphis) indica Chakrabarti & Medda    Colour in life unrecorded; BL of aptera 2.9-3.1 mm. Host-alternating between Sorbus cuspidata and Saussurea piptathera in Uttar Pradesh Garhwal Hills), India. Described as subspecies of D. pavlovskyana (Chakrabarti & Medda 1993), but given full species status by Stekolshchikov (2014a).

Dysaphis inulae Rezwani    Apterae are greyish brown, wax-dusted, with pale tibiae and dark siphunculi and cauda; BL  1.8-2.3 mm. On Inula brittanica in Iran (Rezwani 2008). Alatae and life cycle are unknown.

Dysaphis lappae (Koch)    Apterae are dirty olive greenish to brownish, sometimes with a purple tinge, and older adults may have yellowish margins to abdomen (see influential points/gallery); BL 1.7-2.5 mm. Alatae have secondary rhinaria distributed III 37-55, IV 9-19, V 0-1. On stem bases, root collars and roots of Arctium spp., ant-attended. Also recorded from Petasites albus. In Europe, Transcaucasia, Central Asia, and Siberia eastward to Irkutsk; also north Africa (Egypt, Eritrea, Tunisia), and (ssp. cynarae) introduced to South America. Monoecious holocyclic on Arctium, with oviparae and alate males in September. Very similar aphids on Cirsium arvense in Europe and on Cynara spp. in the Mediterranean region and Middle East are currently classified as subspecies, D. lappae ssp. cirsii (Börner) and D. lappae ssp. cynarae (Theobald) respectively.  The latter subspecies is also reported from Argentina (Ortego 1998a) and South Africa (Millar 1994). D. lappae cirsii is possibly a synonym of the Central Asian D. radicivorans.  Colonies of D. lappae cirsii are reportedly not ant-attended (Stroyan 1963). In Sicily, anholocyclic populations of D. lappae cynarae have also been found on Cirsium syriacum, Galactites tomentosa and Silybum marianum (Barbagallo 1974).

Dysaphis laserpitii (Börner)    Apterae are yellowish grey or greenish grey, wax-dusted; BL 2.1-2.2 mm. On Laserpitium latifolium in Europe (Germany, France, Sweden, Poland, Austria). Heteroecious holocyclic, with a sexual phase on Crataegus (Heie 1992). Gynoparae sometimes produce numerous oviparae on Crataegus sp. in autumn (BMNH collection, leg. H.L.G. Stroyan). Spring generations on Crataegus sp(p). are undescribed, but presumably they form similar galls to other members of the crataegi group. Spring migrant alatae collected from galls are not available, and therefore this species is not included in the key under Crataegus.  It is closely related to C. angelicae and C. lauberti (Stroyan 1963), and specimens could key to either of those species. (Alatae collected in July on Laserpitium in Sweden have 0-10 secondary rhinaria on ANT V, and longest hairs on ABD TERG 2-3 are 22-36 µm long and rather abruptly pointed.)

Dysaphis lauberti (Börner)  Forming red blistered-leaf galls on Crataegus leaves in spring, with second generation alatae migrating to form ant-attended summer colonies on lower leaf bases and root collar of Heracleum sphondylium. There are also records from other Umbelliferae/Apiaceae; Conium, Foeniculum, Pastinaca and Smyrnium. Apterae on secondary hosts are pinkish-grey to greenish grey, wax-powdered; BL 1.7-2.6 mm. Alatae have secondary rhinaria distributed III 46-93, IV 11-40, V 0(-5). Return migration of gynoparae and males occurs in September.  In Europe (UK, Netherlands, Germany, Spain, Belarus). Similar aphids collected on Heracleum lehmannianum in Tajikistan show some differences from European populations (Stroyan 1985). Some samples from Pastinaca in UK have shorter, blunter antennal hairs and could be hybrids between lauberti and crataegi (Stroyan 1963). (Note: spring migrant alatae of the closely related species laserpitii (q.v.) may key to lauberti.)

Dysaphis leefmansi (Hille Ris Lambers)    Apterae are pale green to olive with a brownish dorsal sclerotic pattern; BL 1.8-2.2 mm. On roots of Valeriana officinalis in ants’ nests in Europe. Monoecious holocyclic with alate males. Müller (1961a) described German populations as a distinct species (krumbholzi), but Shaposhnikov & Stekolshchikov (1994) compared populations from Netherlands, Germany and Russia (St Petersburg), and gave the German and Russian populations subspecies status (D. leefmansi ssp. krumbholzi Müller and D. leefmansi ssp. valentinae Shaposhnikov respectively). Apterae of the three forms are less easily distinguished than other morphs (fundatrices, alate viviparae, oviparae and males; see keys in Shaposhnikov & Stekolshchikov 1994).

Dysaphis libanotidis Shaposhnikov    Apterae are dark grey with slight greenish tinge, slightly pulverulent; BL is not given. On Libanotis montana (= Seseli libanotis) in Russia (north Caucasus). D. seselii may be this species.

Dysaphis ligulariae Narzikulov    Apterae are greenish grey, wax-covered; BL c. 2 mm. On Ligularia spp. in Tajikistan (Narzikulov & Daniyarova 1990) and Kazakhstan (Kadyrbekov 2013e).

Dysaphis malidauci Shaposhnikov   Rolling and reddening the lateral margins of leaves of Malus in spring, migrating in the second generation to Daucus (Shaposhnikov 1987a,b).  Immatures in the leaf-rolls are green.  In Kazakhstan, and also recorded from Pakistan (Balichistan; Naumann-Etienne & Remaudière 1995).  Described as a subspecies of foeniculus Theobald, which occurs throughout the world as anholocyclic populations on Umbelliferae/Apiaceae (especially Daucus, Foeniculum) and Polygonaceae (Rumex), but which is not known to have a sexual phase. As it is untenable for anholocyclic taxa to have subspecies that are holocyclic, D. malidauci is here regarded as a separate species, although the two forms are practically indistinguishable (for partial discriminants see Shaposhnikov 1987a).  2n=12.

Dysaphis (Pomaphis) maritima (Hille Ris Lambers)     Apterae are brownish or pinkish yellow or pinkish red, with dark tips to appendages; BL 1.8-2.1 mm. Alatae have secondary rhinaria distributed III 25-49, IV 5-12, V 0-4. On aerial parts of Plantago spp. in coastal areas, not usually ant-attended. In northern Europe including Iceland. Monoecious holocyclic with alate males. 2n=12.

Dysaphis (Cotoneasteria) microsiphon (Nevsky)   Plate 11d   Apterae are greenish yellow, probably somewhat waxy; BL c. 2.1-2.3 mm. In spring slightly rolling leaves of Cotoneaster spp., then migrating to roots of herbaceous plants in several different families including Labiatae/Lamiaceae (Clinopodium, Mentha), Rosaceae (Fragaria, Geranium) and Compositae/Asteraceae (Cnicus, Artemisia, Taraxacum). The root-feeding apterae are described as black in colour by Chakrabarti & Das (2014). Recorded from Belgium (Nieto Nafría et al. 1999), Greece, Morocco, Ukraine, Turkey, Iran, Georgia, Turkmenistan, Tajikistan, Uzbekistan, Kazakhstan, Pakistan, India and Bhutan (Chakrabarti & Das 2014).  Chakrabarti & Medda (1993) compared the morphology of populations on different hosts.  The extent of the dorsal sclerotic pattern in apterae on secondary hosts varies greatly between populations. 2n=12.

Dysaphis mirabilis (Nevsky)    Apterae are yellowish with dark brown cross-band on last abdominal tergite; BL c. 2.7 mm. Forming large colonies in curled leaves of  Ferula spp. in Central Asia. A member of the difficult D. foeniculus species complex; Shaposhnikov (1987b) suggested synonymy with D. ferulae.

Dysaphis mordvilkoi Shaposhnikov    Rolling and reddening the lateral margins of leaves of Malus spp. in spring in Siberia (Amur, Primorsk). Heteroecious holocyclic, migrating to basal parts of Libanotis seseloides; apterae on Libanotis are pinkish grey dusted with greyish-white wax; BL 1.3-2.3 mm (Shaposhnikov & Stekolshchikov 1989). Specimens from Cnidium monnieri in China (Beijing; BMNH collection, leg. V.F. Eastop) may also be this species.

Dysaphis munirae Shaposhnikov    Colour of apterae in life is unknown; BL 2.0-2.4 mm. On Elaeosticta ( = Scaligeria) bucharica in Takjikistan and Kazakhstan (Kadyrbekov 2004b). Heteroecious holocyclic, with a sexual phase on Crataegus spp. (korolkovii, pontica, songorica), presumably forming crataegi-like red curled-leaf galls in spring. Unlike many Dysaphis, the first tarsal taxonomy is not 3,3,2 but 3,3,3 (i.e. HT I has a sense peg).

Dysaphis narzikulovi Shaposhnikov    Apterae are pale brown, with wax; BL c.2.4-2.5 mm. On basal parts of Arctium leiospermum in Tajikistan (Narzikulov & Daniyarova 1990).

Dysaphis neostroyani Ilharco    Apterae are greenish grey with wax powder; BL 1.5-2.4 mm. On stems and leaves of Ophrys fusca in Portugal, and found on Orchis sp. in southern France and Ukraine. Stroyan (1985) summarised its distinguishing characters. Stekolshchikov (2008) confirmed that specimens collected by Bozkho (1976a) were this species, redescribed apterae and alatae and compared its morphology with that of  D. alii and D. eremuri.

Dysaphis newskyi (Börner)    Apterae are pinkish to lilac grey, wax-powdered; BL 1.5-2.7 mm. Apterae are all alatiform and have secondary rhinaria distributed III 4-55, IV 0-20, V 0-3; alatae have them distributed III 42-69, IV 11-24, V 0-5 (in one alata of ssp. aizenbergi from Finland III 33-35, IV 8-10, V 2). In ant-attended colonies in basal leaf sheaths and on root collar of Heracleum spp. in Europe (UK, Finland, Austria) and Kazakhstan (Kadyrbekov 2012a). Monoecious holocyclic with apterous males. Vicariant forms found on Angelica spp. in Sweden, Finland and Kazakhstan (Kadyrbekov 2004b), on Heracleum sibiricum in Finland and northern Russia, and on Heracleum dissectum in Ukraine and Kazakhstan (Kadyrbekov 2009b) are currently regarded as subspecies; D. newskyi ssp. ossiannilssoni Stroyan, D. newskyi ssp. aizenbergi Shaposhnikov and D. newskyi ssp. mamontovae Shaposhnikov respectively. 2n=12 for ssp. aizenbergi.

Dysaphis oreoselini Szelegiewicz    Apterae are pale greenish to whitish yellow, without wax; BL 1.5-2.0 mm. In Poland, especially in southern steppes, living on roots and basal parts of  Peucedanum oreoselinum, not attended by ants. Monoecious holocyclic, with apterous males (original description). 

Dysaphis papillata (Nevsky)    Apterae are cream-yellow, with black dorsal spots, and cross-bands on posterior abdomen; BL c. 2.3 mm.   On leaves of Prangos spp. in Central Asia. A member of the difficult D. foeniculus species complex; Shaposhnikov (1987b) suggested synonymy with D. ferulae.

Dysaphis (Pomaphis) parasorbi (Börner)  Forming leaf nests on Amelanchier ovalis in Austria (Börner 1952).  Life cycle unknown.  Very similar to D. sorbi, except for the very short antennal hairs (those on ANT III less than 0.2 × diameter of segment).  See Stroyan (1963).

Dysaphis (Pomaphis) pavlovskyana Narzikulov  (fig. 109D)  Spring generations distort and discolour the leaves of Sorbus spp. (persica, turkestanica) in Tajikistan (Narzikulov 1957), and subsequently recorded from other Sorbus spp. in Turkey, countries in central Asia, Pakistan, India, and across Siberia.  Migration occurs in late May-June to Plantago sp(p).; a report of Hesperis matronalis as a secondary host in western Siberia (Ivanoskaya 1977) seems to be erroneous (Stekolshchikov & Shaposhnikov 1998). Apterae on Plantago are dark brown with pale head, and appendages pale except at apices; BL 1.4-2.2 mm.  A full description of the life cycle and all morphs except oviparae was provided by Stekolshchikov & Shaposhnikov (1998), including a study of inter-population variation. Similar aphids (in densely wax-covered colonies) have been collected on S. lanata in Pakistan (Naumann-Etienne & Remaudière 1995), and were also found on S. graeca in Turkey (G. Remaudière, cited by Stroyan 1985); data provided by Stekolshchikov (2014) indicate clinal variation wihin one species.  Khuda Bukhsh & Pal (1983) recorded 2n=12 for D. pavlovskyana collected in India on Sorbus (= Pyrus) lanata.

[D. pavlovskyana ssp. indica Chakrabarti & Medda has now been given full species status by Stekolshchikov (2014).]

Dysaphis peucedani Szelegiewicz    Apterae are greenish-white, wax-powdered, with blackish antennae, legs and siphunculi; BL 1.8-2.6 mm. On roots of Peucedanum oreoselinum and P. alsaticum, attended by ants. In Hungary and Romania (Holman & Pintera 1981, as Dysaphis sp. ?uralensis). This species seems very similar to D. apiifolia.

Dysaphis physocaulis Shaposhnikov    Rolling and reddening the leaves of Malus domestica in Crimea, migrating in early May to Myrrhoides (= Physocaulis) nodosus. Apterae on the secondary host are dark grey-green coated with white wax; BL 1.8-2.4 mm.  Alatae (gynoparae) have secondary rhinaria distributed  III 48-55, IV 18-24, V 0-2.  Return migration to apple occurs in June after only 1-2 generations on the secondary host (Shaposhnikov 1990). Alatae trapped in Pakistan may also be this species (Naumann-Etienne & Remaudière 1995). It is a member of the D. anthrisci species complex.

Dysaphis pimpinellae Shaposhnikov    Apterae are brown, green, yellowish or pink-green, pruinose ventrally; BL 1.8-2.2 mm. At bases of lower leaves of Pimpinella sp., in ant shelters, and on roots of P. aromatica (Shaposhnikov 1986). In southern Caucasus (Armenia, Georgia). The life cycle is unknown.

Dysaphis (Pomaphis) plantaginea (Passerini)  Plate 11c   Forming yellowish crumpled-leaf galls on Malus spp., and sometimes on Pyrus in warmer regions.  Apterae on the primary host are dull reddish to pinkish grey with a pale grey wax bloom (see influentialpoints.com/Gallery); BL 2.1-2.6 mm. Spring migrant alatae appear in the third generation and migrate to form colonies along veins on undersides of leaves of Plantago spp. These colonies are not attended by ants. Apterae on plantains are pinkish or reddish yellow, with dark-tipped siphunculi; BL 1.4-1.7 mm. Alatae produced on the secondary host have secondary rhinaria distributed III 54-92, IV 16-40, V 1-11. In Europe, Africa, south-west and central Asia, India, Pakistan, Nepal, and also in North and South America. In east Asia (Japan, Taiwan, Korea, Thailand) there are populations on Plantago that are morphologically indistinguishable from plantaginea, but they have a different feeding site, at the bases of leaf petioles, and there are no records from apple, suggesting that a distinct species, for which the name D. plantagifoliae (Shinji) is available, occurs in that region. However, D. rotalis Zhang, Chen, Zhong & Li was described from curled leaves of Malus baccata in China (G. Zhang et al. 1999), and was synonymised with D. plantaginea by Stekolshchikov (2007).

Work on this well-studied pest of apple in Europe includes population dynamics (Graf et al. 1985), genetic variation within and between populations in space and time (Guillemaud et al. 2011), interactions with hypersensitive apple plants (Lyth 1985), and timing of control measures (Hull & Starner 1983). Its ecology on Plantago in western USA was studied by Cockfield et al. (2011). It has now been demonstrated that males of D. plantaginea are specifically attracted to female sex pheromones when in conjunction with a particular combination of apple tree odours released by the feeding oviparae (van Tol et al. 2009). See also Blackman & Eastop 2000, p.267.  2n=12.

[A population on apple in Pakistan (Baluchistan) differs from European D. plantaginea in having pale basal parts to siphunculi and shorter antennal and dorsal hairs (Stroyan 1957, Chakrabarti & Medda 1993) and was suggested by Naumann-Etienne & Remaudière (1995) to be the primary host form of D. gallica, a species only known from Europe on secondary hosts. However, D. ariae has also been suggested as the primary host form of D. gallica, so further work is needed to resolve this question.]

Dysaphis (Pomaphis) plantaginis (Pašek)    Apterae are whitish or yellowish, with dark-tipped appendages; BL c.2 mm. On roots and leaf bases of Plantago spp. in Slovakia, and also reported from Poland (Szelegiewicz 1966b).  Alatae are unknown.

Dysaphis pseudomolli Narzikulov    Apterae are dark bluish green dusted with grey wax; BL c.1.4-1.8 mm. On basal parts of Cousinia pseudomollis in Tajikistan.

Dysaphis pulverina (Nevsky)    Apterae are pale green to greenish yellow, densely covered with bluish mealy wax; BL 1.8-2.4 mm. Alatae have secondary rhinaria distributed III 19-36, IV 0-9. On upper and lower sides of leaves of Plantago major in Central Asia (Nevsky 1929a, Kadyrbekov & Aoitzhanova 2005), and also recorded from Iran on Veronica persica and Ceratocephalus falcatus as well as Plantago (Stroyan 1972a, as ssp. iranica, synonymised with pulverina by Shaposhnikov 1987a).  Very close to D. foeniculus, and treated as a subspecies of foeniculus by Shaposhnikov (1987a).

Dysaphis (Pomaphis) pyraria Narzikulov   Spring generations distort the leaves of Pyrus korshinskyi in Tajikistan.  The life cycle is unknown; only the apterous fundatrigenia is described (Narzikulov 1961, Stroyan 1963, Shaposhnikov 1988).  It is closely related to D. pyri. A record from Turkey (Şenol et al. 2014c) requires additional confirmation.

Dysaphis (Pomaphis) pyri (Boyer de Fonscolombe)  (Fig.29d)   Spring generations distort and yellow the leaves of Pyrus communis; it is also recorded from P. amygdaliformis, P. nivalis, P. regelii and P. ussuriensis, Malus sargentii and Cydonia vulgaris.  Records from Sorbus appear to be unsubstantiated. Apterae are wax-covered, brownish-red to dark brown with black siphunculi (see influential points/gallery).  Migration to Galium spp. occurs after 2-3 generations on pear, which in some years can result in heavy attacks. Apterae on Galium are pink to reddish brown, powdered with grey wax; BL 1.0-1.4 mm. They feed at or below ground level on roots and prostrate stems, and sometimes colonise other Rubiaceae (Asperula, Rubia). Alatae (gynoparae) produced on Galium have secondary rhinaria distributed III 15-60, IV 6-22, V 0-8.  In Europe, North Africa, south-west and central Asia, northern India, Nepal, Pakistan, and introduced to Colorado, USA (Smith & Parron 1978).  Kolesova (1974) gave a detailed account of its life history in Crimea. Santas (1987) recorded predators of colonies on P. amygdaliformis in Greece.  To judge from a paratype aptera in the BMNH collection, D. multisetosa Basu, described from P. communis in West Bengal, seems to be a synonym of pyri. Bašilova & Rakauskas (2012a) compared nuclear and mitochondrial DNA sequences with those of D. reaumuri.  See also Blackman & Eastop 2000, p.267 and Stroyan (1957). 2n=12.

Dysaphis radicivorans (Nevsky)    Apterae are yellowish green, with brown head, prothorax and dorsal cross-bands on posterior abdominal tergites ; BL 1.8-2.2 mm. On roots of Compositae/Asteraceae (Cirsium, Cousinia, Sonchus) in Central Asia (Shaposhnikov 1956). Closely related to Dysaphis lappae ssp. cirsii in Europe, but longer-haired (judging from the redescription of radicivorans by Narzikulov & Daniyarova (1990), which differs in several respects from Nevsky’s original account).

Dysaphis radicola (Mordvilko)  (Fig.2a)   Spring generations roll and redden the lateral margins of leaves of Malus (orientalis, domestica), in the same way as others species of the anthrisci group.  However, the small primary gall near the apex of the leaf produced by the fundatrix comprises a longitudinal fold near the mid-rib, rather than the transverse fold of made by fundatrices of anthrisci, etc. (Shaposhnikov 1964).  Alatae are produced in the second generation and migrate to form colonies on the roots of Rumex spp.  Apterae on Rumex are greenish grey to leaden grey, wax-powdered (see influentialpoints.com/Gallery); BL 1.5-2.6 mm. Alatae produced on Rumex have secondary rhinaria distributed III 26-53, IV 3-18, V 0-1. It occurs throughout Europe, in the Caucasus, in Japan and Australia, and alatae apparently of this species have been trapped in USA (BMNH collection).  Stekolshchikov (2006) gave a redescription of all morphs. Shaposhnikov (1964) described a shorter-haired form with similar host relations in southern Russia as a subspecies, D. radicola meridialis, and this was given separate species status by Eastop & Hille Ris Lambers (1976). D. radicola-group populations in Greece, Spain and Portugal are similarly short-haired. Stekolshchikov (2006) showed that there was a latitudinal cline in hair length of D. radicola, and therefore synonymised meridialis with radicola.  Anholocycly on Rumex roots occurs commonly where this species has never been found on Malus; e.g., in Britain (Stroyan 1963), Japan (Sugimoto 2003) and Australia (Australian Capital Territory; BMNH collection, leg. V.F. Eastop). An anholocylic population in Georgia was described as a subspecies, “D. radicola ssp. anholocyclica”, by Stekolshchikov & Lobanov (2002), and Stekolshchikov (2006) considered that anholocylic populations in other parts of the world might belong to this subspecies. However, anholocycly per se is not a valid reason for subspecies designation, the morphological distinction between the two forms is not clear, and more evidence from experimental studies and DNA analysis is needed to clarify genetic relationships. 2n=12.

Dysaphis ramani Das & Raychaudhuri   Only apterous viviparae are described, as dark brown, wax-covered aphids heavily infesting and curling leaves of Crataegus sp. in April in Nepal (Das & Raychaudhuri 1983).  Presence of apterae without alatae indicates that colonies persist on the primary host beyond the second generation, which is unusual for a Crataegus-feeding species of Dysaphis.  The life cycle is unknown.

Dysaphis ranunculi (Kaltenbach)  (Fig.45c)   Curled-leaf galls on Crataegus are pale yellowish-green, often suffused with rosy pink, but never with the sharp demarcation line between red of gall and green of leaf lamina found in crataegi.  The fundatrix is deep blue-grey, with a grape-like bloom, immature alatae brownish to grey.  Migration occurs in the second generation to basal parts of Ranunculus spp. Apterae on Ranunculus are mottled grey-green, brownish around siphuncular bases, wax-dusted, with variable darker sclerotisation; BL 1.7-2.3 mm. They are mostly alatiform, with secondary rhinaria distributed III 0-26, IV 0-13, V 0-5. Alatae produced on Ranunculus have secondary rhinaria distributed III 40-71, IV 13-30, V 0-11.  The return migration to Crataegus occurs in September (Stroyan 1963).  In Europe and central Asia.

Dysaphis rara Shaposhnikov    Described from oviparae, with BL c. 1.8 mm. On Capsella bursa-pastoris  in Ukraine, and possibly apterous viviparae collected in Czech Republic (Stroyan 1985, Holman 1991c) are this species (although other short-haired Dysaphis may also be using Capsella as a reserve host). Presumably monoecious holocyclic on Capsella; oviparae occur in late September (original description).

Dysaphis (Pomaphis) reaumuri (Mordvilko)   (Fig.29e)   Colonies roll and twist both leaf laminae and petioles of Pyrus spp. (see Patti & Barbagallo 1997). Apterae on pear are light green covered with wax in early spring, later dark green, with pale siphunculi; BL 1.3-2.0 mm.  Migration occurs to stem bases and roots of Galium spp., but colonies also persist on pear and even produce oviparae in autumn (Kolesova 1974). Apterae on Galium are greyish green, with rust-red spots around siphunculi, lightly wax-dusted; BL 1.0-1.5 mm. Alatae produced on Galium have secondary rhinaria distributed III 29-42, IV 4-12, V 0-4. In southern Europe and central Asia.  Kolesova (1974) gave a detailed account of its life history in Crimea. Patti & Barbagallo (1997) provided detailed redescriptions of apterae and alatae from populations on Pyrus in Italy. Bašilova & Rakauskas (2012a) compared nuclear and mitochondrial DNA sequences with those of D. pyri. See also Blackman & Eastop 2000, p.268. 2n=12. 

Dysaphis rumecicola (Hori)    Apterae are milky white, pale green or dark purplish brown, wax-dusted; BL 2.2-2.6 mm. On Rheum and Rumex spp., at bases of stems sheltered by ants.  In Japan (Miyazaki 1971), Korea (Lee et al. 2002c) and India (Himachal Pradesh, Uttar Pradesh; Chakrabarti & Medda 1993). [Closely related to D. emicis (q.v.), which should perhaps be treated as a synonym, since Stekolshchikov (2006) demonstrated that there is an east-west cline in hair length and shape in populations of the emicis/rumecicola group. We retain both names here pending further investigation, in order to avoid hiding potentially useful information.]

Dysaphis selinumi Chakrabarti & Medda    Appearance of apterae in life is unrecorded; BL c.2 mm. On Selinum tenuifolium (= wallichianum) in Uttaranchal and Uttar Pradesh, India. Similar to the Japanese species D. taisetsusana.

Dysaphis seselii Vaskovskaya    Apterae are pale greenish grey, wax-dusted; BL c. 1.2 mm. On Seseli dichotomum in Crimea. The description, although detailed, does not clearly distinguish this species from D. libanotidis, another short-haired Dysaphis described from a related umbellifer.

Dysaphis (Pomaphis) shaposhnikovi Stekolshchikov    Apterae are greenish yellow brown, with dark brown siphunculi; BL c.2.2 mm. In leaf axils and upper sides of leaf bases of Plantago sp. in east Russia (Primorsk). Monoecious holocyclic (Stekolshchikov 1998).

Dysaphis sharmai Stroyan   Appearance in life unknown; on Pyrus pashia in Nepal, presumably distorting leaves.  Also in India (Himachal Pradesh; BMNH collection, leg. D. Hille Ris Lambers).  The life cycle is unknown, but the presence of apterae indicates that the species is either monoecious on Pyrus or migrates to a secondary host in third or later generations (Stroyan 1982b).  Shaposhnikov (1988) discussed the taxonomic affinities of this aphid.

Dysaphis sibirica Shaposhnikov   Galls on Malus are formed by longitudinal folding and swelling of the leaf margin, becoming greenish yellow with red patches or entirely red (Stekolshchikov & Shaposhnikov 1998).  All the second generation is alate, migrating in mid-June to colonise Aegopodium spp. (alpestre, podagraria). Other reported secondary hosts, e.g. Carum carvi and Sphallerocarpus gracilis (Umbelliferae/Apiaceae) and Galeopsis bifida (see Shaposhnikov 1986) were not colonised in cage experiments (Stekolshchikov & Shaposhnikov 1998).  In eastern Siberia.

Dysaphis (Pomaphis) sorbi (Kaltenbach)  Forming crumpled-leaf galls in spring on Sorbus spp., especially aucuparia.  Mid-rib and leaflets curl into tight bunches, with only slight discoloration.  Apterae on Sorbus are dull green or olive-brown or reddish (see influentialpoints.com/Gallery); BL 2.1-3.1 mm. Alatae are not produced in large numbers until the 4th generation on Sorbus in June; they migrate to basal parts of Campanula spp., and perhaps to Jasione montana.  Apterae on secondary hosts are dull yellowish; BL 1.3-1.7 mm. Alatae produced on the secondary host have secondary rhinaria distributed III 45-76, IV 20-26, V 5-9.  Host alternation seems to be facultative; colonies may persist on Sorbus into late summer, and possibly these produce sexuales without host alternation (Stroyan 1957, Shaposhnikov 1963).  Throughout Europe except the Iberian peninsula, and across Asia to east Siberia (Irkutsk; Stekolshchikov & Shaposhnikov 1998). Old records from North America are probably D. plantaginea. Barbagallo & Patti (1994) provided a key to distinguish D. sorbi from two closely-related spp. on Campanulaceae (brevirostris and henrystroyani). 2n=12.

Dysaphis sorbiarum (Narzikulov)   Apterae are pale greenish-yellow, slightly wax-dusted; reported to live on upper surfaces of leaves of Crataegus altaica and C. hissarica in Tajikistan (Narzikulov 1954). Apparently living all year on Crataegus, with alate males, alate females being rare (see also Stroyan 1963).

Dysaphis tadzhikistanica Daniyarova    Apterae are yellowish green to whitish, wax-dusted; BL c.2.3-2.4 mm. On Ferula jaeschkeana in Central Asia. Shaposhnikov (1987a) gave some additional morphological data, comparing it with other members of the D. foeniculus species complex.

Dysaphis taisetsusana (Miyazaki)   Apterae are green, wax-covered; BL c.2.2-2.3 mm. Described from an unidentified umbelliferous plant, living under the leaf sheaths, and subsequently recorded from ?Peucedanum sp. (Stroyan 1985). The cauda is unusual for a Dysaphis in being elongate triangular, distinctly longer than its basal width, and longer than R IV+V. This species is only known from Japan. The Indian species D. selinumi is similar.

Dysaphis taraxaci Daniyarova    Apterae are pale brown, with wax; BL c.1.7 mm. On basal parts of Taraxacum officinale in Tajikistan.

Dysaphis tschildarensis Daniyarova   Spring generations distort the leaves of Pyrus communis in Tajikistan and Kazakhstan.  Host alternation occurs to Ferula spp. Populations on Angelica decurrens are regarded as a subspecies, D. tschildarensis ssp. tuberculata  (Shaposhnikov 1988). Apterae on the secondary hosts are greyish brown, wax-dusted; BL c.2.6-2.7 mm. Alatae trapped in Pakistan may also be this species (Naumann-Etienne & Remaudière 1995).

Dysaphis tulipae (Boyer de Fonscolombe)  Plate 11b   Apterae are whitish to pale yellow, wax-powdered; BL 1.7-2.3 mm. Alatae have secondary rhinaria distributed III 24-55, IV 3-14, V 0. On many monocots including Arum, Chionodoxia, Crocus, Freesia, Gladiolus, Iris, Lilium, Moraea, Musa, Scilla and Strelitzia, colonising shoots and leaves of growing plants, and on underground stems and bulbs. Records from Umbelliferae/Apiaceae, e.g. in Bermuda (Stoetzel & Hilburn 1990) may be due to confusion with the very similar D. foeniculus. Sometimes ant-attended. Almost cosmopolitan (now including South America). Apparently entirely anholocyclic. Stekolshchikov (2008) provided a detailed redescription. 2n = 11 or 12.

Dysaphis ubsanurensis Ivanoskaya    Apterae are brownish; BL c. 2.1 mm. In ant shelters on roots of a Salsola sp., or possibly an Achnatherum (= Stipa) sp. (the actual host is uncertain, but these two plants prevailed at the collection site). In Siberia (Tuva Republic). Stekolshchikov (2008) has redescribed this species. The life cycle is unknown.

Dysaphis unicauli Mukhamediev    Apterae are dark green powdered with greyish wax; BL 1.4-2.0 mm. On Seseli unicaule, feeding on undersides of leaves, which mostly become rolled and twisted. In Central Asia.

Dysaphis uralensis Shaposhnikov    Apterae are pale green, without wax; BL 1.7-2.1 mm. On roots of Peucedanum alsaticum in Russia and Kazakhstan, and alatiform apterae agreeing with the description of this species have also been collected on Pastinaca in Sweden (BMNH collection, leg. V.F. Eastop).

Dysaphis ussuriensis Shaposhnikov & Stekolshchikov    Apterae are greenish grey, lightly dusted with white wax; BL 1.8-2.4 mm. On root collar and basal leaf sheaths of Anthriscus aemula (= nemorosa) in eastern Siberia (Stekolshchikov 1998). Closely related to D. anthrisci (q.v.), but without host alternation.

Dysaphis vandenboschi Stroyan    Colour of apterae in life is unrecorded; BL 1.6-2.4 mm. On Lepidium (= Cardaria) draba in Italy, southern France and Iran. Shaposhnikov (1987b) described a population with more tubercles on Lepidium crassifolium (= L. cartilagineum) in Ukraine as a subspecies, D. vandenboschi ssp. lepidii, and this form was also recorded from.L. perfoliatum in Kazakhstan by Kadyrbekov (2005d).

Dysaphis (Pomaphis) viennoti Remaudière    Apterae are shining black; BL 1.0-1.2 mm. On Seseli sp., active and easily disturbed, ant-attended. Only known from Iran. Monoecious holocyclic with oviparea and alate males in early September (original description).

Dysaphis virgata Shaposhnikov & Moralev   Forming pale green or red curled-leaf galls on Crataegus orientalis in Armenia (Shaposhnikov & Moralev 1978).  Immatures in galls are rose pink with grey pubescence.  The second generation is alate, migrating to an unknown secondary host.  See also under crataegi, and Stroyan (1985).