SYSTEMATIC TREATMENT OF APHID GENERA (in
alphabetical order) D
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 (see influentialpoints.com/Gallery).
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. Benheim et al. (2012)
reviewed alternative pest management options. 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;
Savoě et al. 2020), and of
differences in virulence between clonal lineages (Herbert et al. 2010; Powell et al. 2013). The sequencing of the
grape phylloxera genome has now been completed (http://bipaa.genouest.org/is/aphidbase/). For further general information see Blackman & Eastop (2000, as Viteus vitifoliae). 2n=10.
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. R. Zhang et al. (2019) studied the genetic mechanisms underlying the
determination of the size of the dorsal processes. 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).
A genus for one very long-bodied species on Phragmites, somewhat resembling Brachycolus but with Phorodon-like processes on the
antennal tubercles.
Davatchiaphis persica Remaudičre Plate 14e Aptere are pale green; BL 2.1-2.4 mm. On
upper sides of leaves of Phragmites
australis (var. stenophyllus)
in Iran (Remaudičre 1964). Monoecious holocyclic with oviparae and apterous
males in November.
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.
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.
A genus for one South American species very similar to Wahlgreniella, but the primary
rhinaria have a thick unciliated rim (Nieto Nafría et al. 2017a). Delfinoia peruviana (Essig) (= Wahlgreniella australis Delfino) Colour of apterae in life is unrecorded,
presumably pale green, with dusky/dark siphunculi; BL 2.2-3.6 mm. Alatae have
4-19 secondary rhinaria on ANT III. On a vine of the genus Cayaponia in
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
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, west Siberia and Kazakhstan. 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, Greece, 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).
Five species related to Thoracaphis,
four of which have apterous viviparae living on Fagaceae with a heavily
sclerotised and sculptured dorsal cuticle, and all except one of these
species have long pointed marginal prosomal hairs (the exception being D. coccidiformis). A fifth species (D. autumna) lives without host
alternation on Distylium, the
presumed ancestral primary host of the genus. Siphunculi are lacking and the
tarsi are often without claws.
Descriptions and available specimens are inadequate to provide
reliable discriminants for all the species. Sorin (2006) reviewed the species
in Japan. Dermaphis 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. This species was until
recently placed in a separate genus (Dinipponaphis),
but was transferred to Dermaphis on
DNA evidence (Aoki et al. 2017). Dermaphis coccidiformis Aoki & Kurosu Apterae are almost black fringed with a
white wax secretion; BL 0.56-0.93 mm. Body oval, usually with asymmetrical
sclerotisation. On evergreen Quercus spp. (glauca, myrsinifolia, salicina) in Japan, feeding at bases
of winter buds or leaf petioles or near bases of new shoots. Alate sexuparae
are produced in February, developing from nymphs feeding through the winter
on the upper surfaces of leaves, and
presumably migrating to an as yet unidentified primary host (Aoki et al. 2017). 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).
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).
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 Remaudičre &
Remaudičre (1997), but is described as bright red in life with long wax filaments (Zhang
et al. 1992a).]
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.
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).
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, Iran,
Kazakhstan, 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). Peńa-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), South America (Chile 1988,
Argentina 1992; Clua et al. 2004),
and now reported from Australia (South Australia, Victoria, New South Wales;
FAO 2016, Yazdani et al. 2018). 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. Parasitoids (Aphelinus
spp.) introduced for biological control in western USA appear to have become
established (Burd et al. 2001).
Earlier Russian language literature was reviewed by Kovalev et al. (1991). 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 both North and South
American populations, and might be occurring undetected in other places, e.g.
South Africa, the pervasive use of the
term ”biotype” in the economic literature about this species should be
reconsidered, as annual genetic recombination has the potential to generate an
unlimited number of “biotypes” (Puterka et
al. 2014, 2015). This will inevitably influence the interpretation of
data, for example, on the incidence of resistance to this aphid in wheat
cultivars (e.g. Jankielsohn 2016). The complete mitochondrial genome has been
sequenced (Bo Zhang et al. 2013; de
Jager et al. 2014; Nicholson et al. 2015). Steyn et al. (2020) undertook isolation and
sequence analysis of the unusually large X chromosome. 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). Puterka (2017) crossed D. tritici with D. noxia
and obtained a hybrid line with reduced fitness but greater virulence to
wheat and barley than either parent. See Parker (1916) for an early account
of the biology.
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) and Qiao et al.
(2018) have given accounts 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 cathayana, ciliata, sieboldii, suaveolens, yunnanensis).
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.
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 (see influentialpoints.com/Gallery). 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), Hungary (Ripka 2010) and Serbia.
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.
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 &
Remaudičre (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 Remaudičre &
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.
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, and
also recorded from Iran (Abaii 1984).
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. Peńa).
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 (Remaudičre 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 oregonensis 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 & Remaudičre
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,
south-west and 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.
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).
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, Iran, 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 & Remaudičre (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 Allium lineare and Allium sp. in
Kazakhstan (Kadyrbekov 2017a). Stekolshchikov (2008) redescribed the apterae
(possibly they are fundatrices) of this species. 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), and also recorded from Iran (Alikhani et al. 2010). 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 is also reported to occur
in Kazakhstan (Kadyrbekov 2017a). It
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 (see influentialpoints.com/Gallery);
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 (see influentialpoints.com/Gallery)
of this form occur on Crataegus in
spring. The fundatrix is
greenish or bluish grey, wax-powdered, 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). Aphids colonising Cenolophium fischeri (= denudatum) in Kazakhstan
have also been identified as this subspecies (Kadyrbekov 2017a). 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 & Remaudičre 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, Caucasus and Iran, 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
Iran (Rezwani et al. 1994), 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
(Ĺland; 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) and Kazakhstan (Kadyrbekov 2017a). 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) and Kazakhstan (Kadyrbekov 2017a). 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 to crimson
curled-leaf galls on Crataegus spp.
(see
influentialpoints.com/Gallery)
in spring. The fundatrix is greenish-grey, thickly
wax-powdered, the immature alatae are pinkish. The 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 (see
also influentialpoints.com/Gallery);
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 (although
in Sicily damaging infestations of what is apparently the nominal subspecies
occur on the fruits of Cucurbita;
Massimino Cocuzza & Barbagallo 2017); D.
crataegi ssp. kunzei (Börner) goes
to Pastinaca; and D. crataegi ssp. aethusae (Börner) to Aethusa
cynapium and Torilis japonica.
In Italy (Sicily and Sardinia), populations feeding on Ferula, and probably anholocyclic, have also been given
subspecies status, as D. crataegi
ssp. siciliensis (Barbagallo &
Stroyan 1982). 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, as they are in southern Europe.
2n=12 (on Daucus). Dysaphis crithmi (Buckton) Apterae are grey to greenish grey,
wax-powdered (see influentialpoints.com/Gallery);
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 see influentialpoints.com/Gallery;
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 Iran, Kazakhstan (Kadyrbekov 2017a) and 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 and M. sieversii 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 & Remaudičre 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, Italy, European Russia, Altai Republic and Kazakhstan.
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 kadyrovi Depa & Kanturski Apterae are greyish green to dark green,
without wax; BL 1.2-2.1 mm. On Anacantha
darwasica (Asteraceae/Compositae) in Tajikistan, feeding on upper sides
of leaves and shoots near inflorescences (Depa et al. 2017b). Other morphs and life cycle are unknown. Unusually
for a Dysaphis, this species lacks
both marginal and spinal tubercles. Dysaphis koryakini Stekolshchikov & Buga Apterae are pale greyish-green to greenish
yellow, lightly wax-powdered, with pinkish head and thorax, dark siphunculi
and pale cauda; BL 1.4-2.4 mm. Colonies are formed on the root collar and
basal leaf-sheaths of Cenolophium
denudatum, tented over by ants. In north-west Russia (Murmansk region;
Stekolshchikov & Buga 2018). Monoecious holocyclic; oviparae and apterous
males were collected in late July-August. From the description this species
seems to be very close to D. apiifolia (s.
lat.), which has also been recorded from Cenolophium.
[The name is an amendation of an originally incorrect spelling as karyakini (Stekolshchikov & Buga
2019a).] 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, Iran,
Central Asia, and Siberia eastward to Irkutsk; also north Africa (Egypt,
Eritrea, Tunisia), and introduced to Australia (about 1992;
Brumley 2020). 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 spp. 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 (see influentialpoints.com/Gallery);
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), and alo recorded from Iran (Rezwani
1990) and Kazakhstan (Kadyrbekov 2017a). 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, where Aegopodium and Angelica are also recorded as
secondary hosts (Kadyrbekov 2017a), and also recorded from Kyrgyzstan and
Pakistan (Balichistan; Naumann-Etienne & Remaudičre 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 Tajikistan and Kazakhstan (Kadyrbekov 2004b).
Heteroecious holocyclic, with a sexual phase on Crataegus spp. (altaica,
korolkowii, pontica,
songorica, turkestanica),
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 (see influentialpoints.com/Gallery);
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 & Remaudičre 1995),
and were also found on S. graeca in
Turkey (G. Remaudičre, 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 &
Remaudičre 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 & Remaudičre (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 and P. media in Central Asia (Nevsky
1929a, Kadyrbekov & Aoitzhanova 2005, Kadyrbekov 2017a), 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 Iran, in Kazakhstan, 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
Turkey, 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, Iran 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, the Middle East 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 & Remaudičre 1995). Dysaphis tulipae (Boyer de Fonscolombe) Plate 11b Apterae are whitish to pale yellow,
wax-powdered (see influentialpoints.com/Gallery);
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 and P. morisonii 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 and L. latifolium
in Kazakhstan by Kadyrbekov (2005d, 2017a). Dysaphis (Pomaphis) viennoti Remaudičre
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). |