SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order) B
A
genus for one species related to Aploneura
and Slavum but with more normal distribution
of rhinaria on the antennae of the alatae, which fold their wings roof-like
in repose. The large characteristic
horn-like galls can be formed on several species of Pistacia. Baizongia
pistaceae (Linnaeus)
Plate 1h Producing large, elongate horn-like galls
15-22 cm long (fig.
126C) on Pistacia spp.,
especially P. palaestina, in the
Mediterranean area, and also in north-west India. Large numbers of alatae, BL 1.8-2.0 mm (fig.
81G), emerge
in September-November and fly to roots of Poaceae (e.g.
Agrostis, Avena, Dactylis, Festuca, Poa, Triticum).
Apterae on grass roots are whitish
or pale yellow, producing wax in threads; BL 1.6‑2.3 mm. They are invariably attended by ants,
especially Lasius flavus, and may
overwinter in ants' nests. The holocycle takes two years (fig. 2). Anholocyclic populations occur on grass
roots more widely in Europe, the Middle East, north Africa, Kenya and the
Indian subcontinent. Roberti (1983)
and Wertheim (1954) gave accounts of this aphid in Italy and Israel
respectively. Wool (1990, 2002), and
Lamb et al. (2012), analysed data
from a 20-year population study on Pistacia
in Israel. Histology of the galls on Pistacia
terebinthus in Spain was studied by Álvarez (2012). Wool & Kurzfeld-Zexer
(2008) studied colonies on cereal roots in culture, rearing them successfully
on oats (Avena sativa and A. sterilis) but not on wheat or
barley. Wool (2011/12) provided a comprehensive review of this
species.
2n=24.
One
species in Central Asia superficially
resembling Liosomaphis but with
better developed antennal tubercles, longer antennae and a short rounded
cauda. Berberidaphis
lydiae (Narzikulov) Apterae are shiny pale yellow, BL 1.7-2.7
mm. On undersides of leaves of Berberis
spp. in mountainous regions of Tajikistan, eastern Kazakhstan (Kadyrbekov
2009a) and northern Pakistan (Naumann-Etienne & Remaudičre 1995).
Oviparae and small apterous males appear in October-November (Narzikulov 1957,
as Liosomaphis lydiae).
Six species are known, all from
east and south-east Asia. All
viviparae are alate, active insects (behaving like Euceraphis), and have a characteristic transverse dark brown to
black bar across the ventral side of the head between the eyes. Most of the species are recorded from
Betulaceae, which are also the hosts of several related genera. One species (prunicola) has apparently become
associated with Prunus. The Indian species are reviewed by
Chakrabarti (1988) and A.K. Ghosh & Quednau (1990), the Japanese species
by Higuchi (1972), and the Chinese species by Qiao et al. (2005b). Betacallis
alnicolens Matsumura
Alatae are pale green with dark tibiae, large (about 3.5 mm), very
active, on leaves of Alnus spp. (hirsuta, japonica, matsumurae)
in Japan, Korea and China, and also collected from Corylus mandshurica in Korea (BMNH collection). 2n=22. Betacallis
luminifera Zhang Alatae yellow; BL 2.6-3.5 mm, on leaves of
Betula luminifera in Zhejiang,
China. Very similar to, and possibly
synonymous with, B. sikkimensis,
but with more secondary rhinaria. Betacallis
odaiensis Takahashi
Alatae are pale lemon-yellow to apricot, with black basal,
sensoriated, part of third antennal segment, black tibiae, two black bars on
the dorsal abdomen, and black siphunculi; BL 2.4-3.1 mm. On leaves of Betula spp. (ermannii, maximowicziana, platyphylla var. japonica)
in Japan. (Indian records of odaiensis refer to other
species.) 2n=22. Betacallis
prunicola R.C. Basu, A.K. Ghosh & Raychaudhuri Alatae light yellowish green with dark
tibiae, dark transverse bars on ABD TERG 3-5 or 4-5 and dark siphunculi; BL
1.9-2.2 mm (A.K. Ghosh & Quednau 1990).
Described from adults and immatures on Prunus cerasus in Meghalaya, India (R.C. Basu et al. 1975), and subsequently
collected from other Prunus spp. so
it now seems likely that this is a valid host association. This species is also recorded from
Arunachal Pradesh, Manipur and West Bengal, and from China and the
Philippines. It can be distinguished
from other Betacallis species by
the long ultimate rostral segment (R IV+V 1.5-2.0 × HT II). Betacallis
querciphaga R.C. Basu, M.R. Ghosh &
Raychaudhuri Alatae light grenish
yellow with dark markings like B.
odaiensis except for the dorsal longitudinal head stripe and differences
in wing vein pigmentation; BL 2.4-3.2 mm.
In India (Manipur, Meghalaya, Sikkim, West Bengal), the Philippines
(Chakrabarti 1988) and Thailand (BMNH collection). It was described from Quercus sp., and alatae and immatures are also recorded from Castanospermum sp.; nevertheless,
specimens from Betula alnoides in
Thailand, from B. luminifera in
China (Qiao et al. 2005b) and from B. utilis in West Bengal (BMNH
collection) indicate that the normal host association is probably with Betula.
Parasitised by Trioxys
(Betuloxys) assamensis in India (Raychaudhuri 1990). Betacallis
sikkimensis R.C. Basu, M.R. Ghosh & Raychaudhuri BL of alata 2.4-3.2 mm, appearance in life
unrecorded but presumably pale, with third antennal segment and tibiae
darkened only at apices (cf. other species).
The siphunculi are black on distal half only and there is often a
consolidated or broken dark central patch on abdominal tergites 4 and 5. Described from Quercus sp. - which is probably only a reserve host - in Sikkim,
India (R.C. Basu et al. 1974);
since collected many times in Uttar Pradesh from Betula alnoides and Betula
sp. (Chakrabarti 1988, and BMNH collection).
Chakrabarti (1988) described the alate male and apterous ovipara from Betula. 2n=20 (Khuda-Bukhsh & Pal 1983b).
A genus
of 6 or 7 species rather evenly distributed through the holarctic on Betula. They are small, rather flat oval aphids
with short 6-segmented antennae, the PT being about as long as the base of
ANT VI. Viviparae have a short,
conical cauda and a bilobed anal plate.
Oviparae have dark dorsal markings and the posterior abdomen is
extended as an "egg-laying organ".
Males are apterous with extensive dark dorsal markings. Fundatrices are unusual in being either
apterous or alate (at least in B.
quadrituberculata). Stroyan (1977)
gave a detailed generic diagnosis, and accounts are available for North
America (Richards 1961), Canada (Richards 1969b), Fennoscandia and Denmark
(Heie 1982), India (Chakrabarti 1988, A.K. Ghosh & Quednau 1990) and
China (Qiao et al. 2005b). There has been much confusion about the
number of species in the brevipilosa/quadrituberculata group, due to
seasonal variation in hair lengths.
Here, B. helvetica Hille Ris
Lambers is regarded as a synonym of B.
brevipilosa, although a study of haemolymph proteins by Karwanska (1987)
suggests that it may be a distinct species.
Parasitised by Aphidius aquilus,
which also attacks other related aphids feeding on Betula (Stary 1973). Betulaphis
brevipilosa Borner (= helvetica Hille Ris Lambers)
(fig. 31B)
Apterae
are pale to grass-green aphids, BL 1.5-2.0 mm, with the tips of the antennae
and the tarsi dark. Alatae almost
always have a dark dorsal abdominal patch.
This species is frequently confused in the literature with B. quadrituberculata. It seems to feed mostly on Betula pendula, usually on the upper
surfaces of the leaves, occurring in north, central and eastern Europe (but
not the UK), Central Asia (Kazakhstan; Kadyrbekov 2017a), and introduced into
North America and New Zealand. However, Buga (1995) reports its occurrence on
several other native and introduced Betula
spp. in Belarus. Accounts are given by
Cottier (1953, as B. quadrituberculata)
and Heie (1982). Ecology of introduced
populations in northern California was studied by Hajek & Dahlsten
(1988). B. viridis Richards 1969 seems to be this species. 2n=20. Betulaphis
hissarica Narzikulov
Appearance in life is unknown.
BL of aptera about 1.7-1.9 mm.
Chakrabati (1988) redescribed both apterous and alate viviparae, and
also apterous oviparae and males.
Recorded from various Betula
species (hissarica, alnoides, turkestanica, utilis) in Kazakhstan, Tajikistan, Pakistan and
India (Himachal Pradesh, Uttar Pradesh). Betulaphis
japonica Takahashi
Appearance in life and biology unrecorded, and alatae are
unknown. BL of aptera 1.1-1.6 mm. Found on various Betula species in
Japan, Korea, and Mongolia. For
further details see Higuchi (1972). Similar to the short-haired form of B. quadrituberculata, but with a
relatively shorter R IV+V. Betulaphis
longicornis Quednau & Chakrabarti Apterae yellow: BL 2.3-2.9 mm. The alatae lack a dorsal abdominal dark
patch, and have numerous (26-35) closely-spaced elongate transverse secondary
rhinaria an antennal segment III.
Apterous and alate viviparae are described by Quednau &
Chakrabarti (1980), and apterous oviparae and males by Chakrabarti (1988). Recorded from Betula alnoides and B.
utilis in India (Assam, Bhutan, Himachal Pradesh, Uttar Pradesh, West
Bengal) and Pakistan. Betulaphis
pelei Hille Ris Lambers (fig. 31C)
Apterae
are yellow with dark tarsi and tips of antennae, BL 1.4-1.7 mm. Alatae sometimes have a dark dorsal
abdominal patch. On Betula nana with a boreo-alpine
distribution in North Europe, Iceland and Greenland (Heie 1982). Aphids occurring on Betula glandulosa, described from Baffin Island as Betulaphis arctosetis Richards 1961,
are probably also this species. It has also been reported to occur on B. albosinensis in western China (Qiao
et al. 2005b). 2n=20 Betulaphis
quadrituberculata Kaltenbach (fig. 31A,D)
Apterae
have BL 1.5-2.0 mm and are pale yellowish green or pale yellow to almost
white, sometimes with patches of darker pigment (especially in autumn),
normally feeding on the undersides of leaves (see
influentialpoints.com/Gallery). Alatae sometimes have a dark dorsal
abdominal patch. Recorded from various
Betula species, especially B. pubescens and also occasionally
found on Alnus incana. Widely distributed in Europe (including
Iceland) and across Asia to Mongolia and China, and introduced into North
America. Apterous males and oviparae
may be found from September to November. There is much variation in length of
the spinal and pleural hairs on the anterior abdominal tergites which causes
short-haired specimens of this species, found particularly in the summer
months, to be confused easily with B.
pelei and B. brevipilosa. B.
aureus Richards 1961, described from Betula
sp. in Canada, seems to be this short-haired form of B. quadrituberculata.
Varty (1964) has studied its biology in New Brunswick, Canada. See
also Heie (1982) and Stroyan (1977). 2n=20.
Five species in the Middle East
and Central Asia closely related to Therioaphis
but specific to Astragalus (and
the related genus Astracantha) with
a conical dorsal process on the knob of the cauda. Quednau (2003) restored
the group to full generic status, and provided illustrations of all available
morphs. Bicaudella astragalensis Rusanova (Fig.16a) Apterae are pale brownish or dirty yellow,
BL 1.1-1.5 mm. On Astragalus spp.
in Afghanistan, Azerbaijan and Iran. Oviparae and alate males occur in
October (Remaudičre 1989b). Bicaudella denaensis (Remaudičre) Apterae are squat-bodied, yellowish with
dark dorsal markings and black siphunculi; BL 1.1-1.5 mm. On Astragalus sp. in Iran. Alatae are
unknown. Oviparae and alate males
occur in September (Remaudičre 1989b). Bicaudella farsiana (Remaudičre) (Fig.16b,d) Apterae are yellow, rather pale, BL
1.6-1.9 mm. On an Astragalus sp. of
the microthrix group in Iran.
Oviparae occur in October (Remaudičre 1989b). Bicaudella
laurestanica (Remaudičre) (Fig.16c,e) Apterae are yellowish with dusky head and
thorax and black dorsal abdominal spots; BL 1.3-1.6 mm. On an Astragalus sp. in Iran. Alatae are
unknown. Oviparae and alate males occur in September-October (Remaudičre
1989b). Bicaudella rostrata (Remaudičre) Apterae are yellowish, rather pale; BL c.1.2-1.4 mm. On Astracantha gossypina in Iran. Alatae and sexual morphs are unknown (Remaudičre 1989b).
A genus
for two nearctic species on Cirsium
related to Artemisaphis and Obtusicauda but with a distinctive
cauda and anal plate. Miller & Jensen (2006) reviewed the genus, removing
one species from synonymy. They are active aphids, and seem invariably to be ant-attended.
Bipersona
ochrocentri (Cockerell)
Plate 28h Apterae have a pale pink head and
thorax, brownish purple abdomen, with last two segments pale green, and black
antennae, black legs except for yellow basal parts of femora, black
siphunculi and black cauda; BL 2.5-3.5 mm. In ant-attended colonies on leaves
and stems of Cirsium spp. (ochrocentrus, arvense) in Arizona and New Mexico, USA. Miller & Jensen
(2006) provided a redescription of apterae and alatae. Sexual morphs have not
been described. Bipersona
torticauda Gillette (Fig.22e) Apterae are brick red to ochre red with
black antennae, black legs except basal parts of femora, black siphunculi,
and anal plate and cauda reddish to dusky; BL 2.5-3.8 mm. In ant-attended
colonies on leaves and stems of Cirsium
spp. (there are records from Carduus and
Cnicus, but these are probably
referable to this genus). In western USA and Canada. Monoecious holocyclic,
with oviparae and apterous males in September-October. Miller & Jensen
(2006) provided a redescription of all available morphs.
A
genus for one species closely resembling Acaudella
in many features such as shape of siphunculi and lack of an evident cauda,
but with a much longer processus terminalis. Blackmania
eastopi Kanturski & Wieczorek Apterae are red-brown in life (V.F.
Eastop, field notes); BL of aptera c.1.6 mm. Alata has 21-31 secondary
rhinaria on ANT III. On Polygonum
equisetiforme in Israel and Cyprus (Kanturski & Wieczorek 2015b). The
life cycle is unknown.
Two species
on Senecio in South America with
dark cylindrical siphunculi having a little poorly-defined reticulation at
apices, and 5 hairs on first tarsal segments. A third species from Cuphea in Brazil mentioned under this
genus name by Blackman & Eastop (2006; p. 1106) has subsequently been
identified as Impatientinum paranaense.
Originally described as Blanchardia,
the name was changed to Blanchardaphis by
Ortego et al. (1998). Blanchardaphis
capitophoroides (Blanchard) Apterae are dark reddish; BL c. 3 mm.
Apterae have 18-20 secondary rhinaria on ANT III, alatae are undescribed. On Senecio sp. in Argentina. Ortego et al. (1997, as Blanchardia) compared measurements with those of B.
poikila. Blanchardaphis
poikila (Ortego, Nieto Nafría & Mier Durante) Apterae are dark pink with a narrow
longitudinal greenish mark on abdominal tergites 2-5; BL 2.2-3.0 mm. Apterae
have 5-14 secondary rhinaria on ANT III, alatae have 16-19. In dense colonies
on flowering stems of Senecio
subumbellatus in Argentina, and recently also reported from Chile (Nieto
Nafría et al. 2018). Monoecious
holocyclic, with alate males (Ortego et
al. 1997, as Blanchardia).
A
genus of 4-5 species associated with Alnus
(including Duschekia). The apterae are dorso-ventrally flattened,
rather elongate oval, and apply themselves closely to the leaf surface. They have paired anteriorly-directed
rectangular projections on the head, lateral abdominal hairs arising from
asymmetric tubercles that give the abdominal margin a stepped appearance, and
typically 5-segmented antennae. The
alatae are more normal-looking Calaphidini.
One species (occidentalis)
differs in that the apterae often have 6-segmented antennae, the frontal
projections are only weakly developed and the abdominal margin is more
indented than stepped. Hille Ris
Lambers & Hottes (1962) revised the genus. Boernerina
alni
Takahashi (fig. 29A,B) Apterae are pale
yellowish-white, dorso-ventrally flattened, elongate oval, BL 1.7-2.6
mm. Alatae are whitish and waxy. Sorin (1970a) described the apterous male
and ovipara, collected in mid-August in Japan, on Alnus spp. Pashchenko
(1983b) described a subspecies (insularia)
in east Siberia, which is very likely to be the aphid from the Irkutsk region
described by Rupais (1989b) as B.
sibirica. This subspecies is also now reported from the Nenets region of
north-west Russia (Stekolshchikov 2017), and specimens with characters of
both insularia and alni s.str. have been collected in
west Siberia (Stekolshchikov & Khureva 2020). Boernerina
depressa Bramstedt
Apterae are whitish, flattened and elongate oval, BL 1.5-2.5 mm, on
undersides of leaves of Alnus viridis
in the Alps and Carpathians, and also reported from A. maximowiczii in the Kuril islands (see Holman 2009). An alata
recorded from west Siberia by Ivanoskaya (1977) has been re-identified as B. alni (Stekolshchikov & Khureva
2020). Oviparae and very small apterous males are produced in July-August
(Hille Ris Lambers & Hottes 1962). Boernerina
occidentalis Hille Ris Lambers & Hottes Apterae are pale, flattened, oval, BL
1.7-2.4 mm, with broad sclerotic bars on abdominal tergites, the anterior
ones being sometimes browhish pigmented.
Alatae are variably pigmented, darker specimens have blackish-brown
sclerotic bars on abdominal tergites.
On Alnus sinuata in Alaska,
also collected in Oregon (BMNH collection; leg. D. Hille Ris Lambers), and
now also recorded from A. alnobetula ssp. fruticosa (=A. fruticosa) in
north-east Siberia (Chukotka; Stekolshchikov & Khruleva 2015). Oviparae and small, slender apterous males
are produced in July in Alaska (Hille Ris Lambers & Hottes 1962). Boernerina
variabilis Richards
Apterae are pale to bright yellow with black on distal parts of
antennae and legs, flattened, elongate oval, BL 1.9-2.6 mm. Alatae vary in pigmentation; when well-pigmented
they have broad dark bars dorsally and ventrally on the abdomen. On undersides of leaves of Alnus spp. (sinuata, rugosa, alnobetula) in east and west Canada
(British Columbia, Northwest Territory, Ontario, Quebec, Labrador). Oviparae and small, apterous males occur in
August in Northwest Territory (BMNH collection, R. O'Doherty), but oviparae
were collected in October in northern Ontario (BMNH collection, leg. J.
S˙pkens). 2n=16*.
This
genus of c.50 mainly palaearctic species is characterised by the large round
spiracular apertures, short often semicircular cauda, and siphunculi with a
subapical annular incision. Brachycaudus is primitively
heteroecious with Prunus as primary
hosts, and about 14 species still host-alternate. There
are species groups associated with secondary hosts in Ranunculaceae (subgenus
Brachycaudina), Caryophyllaceae
(subgenus Acaudus), Boraginaceae
and Compositae (subgenus Prunaphis),
and plants formerly in Scrophulariaceae but now placed in Orobanchaceae
(subgenus Scrophulaphis). For
an introduction to the genus and more detailed accounts of seven of the Prunus-feeding species see Blackman
& Eastop 2000, p.246. Heie (1992)
reviewed the species in Fennoscandia and Denmark, and Blackman (2010)
reviewed the British species (but there are errors in the subgeneric
classification in the latter work). Burger
(1975) reviewed European species then placed in Acaudus. Andreev (2004) revised the subgeneric
classification (but his synonymy of subgenus Thuleaphis with subgenus Mordvilkomemor
is not accepted here – see Blackman & Eastop 2011). Cocuzza et al. (2007) made a morphological and
molecular study of subgenus Appelia.
Coeur d’acier et
al.
(2008) and Jousselin et al. (2009, 2010) studied the molecular
phylogenetics of Brachycaudus species and their bacterial symbiont,
which largely confirm the subgeneric groupings used here. Jousselin et al. (2010) analysed their data
further to reconstruct ancestral character states, and reached the rather
surprising conclusion that several species may have reacquired the primary
host after periods of monoecy on secondary hosts. This interesting finding
requires further investigation. Brachycaudus
(Thuleaphis) acaudata (Hille Ris
Lambers) Colour of apterae in life unknown, probably
greenish with extensive blackish dorsal sclerotisation; BL 1.5-1.7 mm. Alatae
have 10-14 secondary rhinartia on ANT III. The host of type specimens from
Greenland is unknown, but it has subsequently been collected from stems of Persicaria viviparum in arctic Canada
(Baffin Island; Richards 1963a). Records of
B. acaudata from Rhodiola (= Sedum) rosea in Iceland
and Wales are referable to B. sedi.
Monoecious holocyclic with apterous males (original description). Brachycaudus
(Brachycaudina) aconiti (Mordvilko) Apterae are shining black, BL 2.0-2.6 mm.
Apterae have 0-8 secondary rhinaria on ANT III, and alatae have 7-13. On Aconitum spp.
and Delphinium spp. colonising
flower stems and leaf petioles, and rarely on undersides of leaves. In
central, south and south-east Europe, Central Asia and Siberia. Monoecious
holocyclic, with apterous males (Nosyrev 1968). Closely related to B. napelli in western Europe;
differing rhinarial numbers on ANT III are often given as a discriminant
between the two, but may not always be
reliable due to alatiformity. Brachycaudus (Appelia) almatinus
Nevsky Apterae according to the
original description are pale brown with black lateral spots, black
siphunculi, femora, tibial apices and tarsi, and dusky antennae. BL about 1.8 mm. The cauda is very short and broad. Alata have a pale brown abdomen, presumably
without a dark dorsal patch. Found in
Kazakhstan on several Prunus spp.,
rolling leaves longitudinally (Nevsky 1951, Kadyrbekov 2014d). Biology is unknown. Brachycaudus
(Thuleaphis) amygdalinus (Schouteden) Host-alternating between Prunus spp., especially P.dulcis or P. persica, and Polygonaceae.
Apterae are green with variably developed dark dorsal markings,
squat-bodied with rather short pale legs and antennae; BL 1.6-2.1 mm. Alatae have a dark central abdominal patch
and secondary rhinaria distributed III 12-21, IV 0-1 (spring migrants) or III
9-16, IV 0 (gynoparae). Colonies in
spring cause rolling of young almond or peach leaves and stunting of new
growth. The leaves are rolled somewhat
obliquely with respect to the mid-rib. rather than perpendicularly to the
mid-rib as in B. helichrysi.
Migration occurs to Polygonum (particularly
small-leaved) species, Persicaria maculosa and Fagopyrum cymosum. On Polygonum
aviculare they feed hidden under the thin bracts surrounding the nodes
(Talhouk 1977). It is also frequently anholocyclic on Polygonaceae, even
where primary hosts are available. In
Europe, the Middle East, North Africa (Tunisia: Boukhris-Bouhachem et
al.
2007, Algeria:
Laamari et al. 2010), Central Asia,
Pakistan and South Africa. Darwish
(1983) described the morphology of the spring generations. See also Blackman & Eastop 2000, p.246. 2n=12. Brachycaudus
(Nevskyaphis) ballotae (Passerini) Colour of apterae in life unknown,
probably shiny dark green to black; BL c.1.7-1.9 mm. On shoots of Ballota
nigra, attended by ants (Szelegiewicz 1981a). In continental Europe
(Germany, Czech Republic, Hungary, north Italy and Poland), and also recorded
from Iran (Hodjat 1993). Monoecious holocyclic. Molecular data (Jousselin et al. 2010) indicate that B. lamii is probably a synonym. Brachycaudus
(Nevskyaphis) bicolor (Nevsky) (Fig.9b) Apterae are shining yellowish tinged with
pink, to pale green, with a large shiny black dorsal abdominal patch (see influential
points/gallery); BL c.2.1-2.4 mm. Alatae have secondary rhinaria
distributed III 28-52, IV 0-8, V 0. Ant-attended colonies occur on root
collars, and at bases of leaves near ground level, of Boraginaceae (e.g. Anchusa, Cerinthe, Cynoglossum,
Echium, Lindelofia, Myosotis)
and also recorded from a species of Codonocephalum
= Inula (Compositae). In UK
(Stroyan 1955, Baker 2009), north-west France (Cocuzza et al. 2010), Madeira, Portugal, Spain, Italy, Greece, Egypt,
Iran (Rezwani et al. 1994),
Caucasus, west Siberia, Tajikistan, Uzbekistan, Kazakhstan, Afghanistan, and
alatae have been trapped in Pakistan. Sexual morphs are unrecorded, and it is
apparently anholocyclic in Italy (Barbagallo & Patti 1998), but spring
colonies have been observed on Amygdalus scoparius in Iran, probably
the primary host (Naumann-Etienne & Remaudičre 1995). The disjunct distribution and paler siphunculi
of British populations suggest that
they may be a distinct taxon, possibly specific to Cynoglossum (Blackman 2010). 2n=12. Brachycaudus
(Thuleaphis) brevirostratus Pashchenko Apterae are dirty green to pale brown;
BL c.1.3 mm. On Polygonum spp. in
east Siberia. Possibly this is a synonym of B. rumexicolens. Brachycaudus
(Prunaphis) cardui (L.) Plate 10a Apterae on herbaceous plants are shiny
black dorsally, light green to yellowish or reddish ventrally (immatures are
green or reddish) (see influentialpoints.com/Gallery);
BL 1.9-2.3 mm. Alatae have extensive dark dorsal pigmentation, and secondary rhinaria distributed III 12-39, IV
0(-4). B. cardui lives in dense
ant-attended colonies on stems and leaves of many species of Compositae (e.g.
Arctium, Carduus, Cirsium, Cynara, Chrysanthemum, Tanacetum,
Matricaria) and Boraginaceae (e.g Borago, Cynoglossum, Echium, Symphytum), and also frequently on
other plants, e.g. Capsella. In
Europe, Asia, north Africa and North America. Heteroecious holocyclic in
continental climates with a sexual phase on Prunus, esp. P. domestica or
P. spinosa (or sometimes P. avium or P. armeniaca). Apterae in rolled leaves of
plum in spring, are usually green with the dorsal cuticle sclerotic and often
brownish, but not shiny black like the summer generations on secondary
hosts. Spring migrant alatae have
secondary rhinaria distributed III 20-30, IV 0(-1). According to Börner
(1952), the aphids which use P. spinosa
as primary host and migrate to basal parts of stems of Compositae are a
distinct species, B. lateralis
(q.v.), but further work seems necessary to confirm this. Central
Asian populations with longer hairs are regarded as a subspecies, B. cardui ssp. turanica Mordvilko (differentiated by Andreev 1999). See
also Blackman & Eastop 2000, p.247.
2n=10. Brachycaudus
cerasicola Mordvilko ex Nevsky (= phlomicola
Nevsky) Apterae are shiny yellowish brown
to dark brown or black due to extensive dorsal sclerotic shield; BL 1.7-2.1
mm. Immatures are dirty brown. Alatae have
secondary rhinaria distributed III 21-50, IV 2-23, V 0(-5). Spring
colonies roll and twist the leaves of Prunus
spp. and turn them red. There is apparently a facultative host alternation to
Lamiaceae (Dracocephalum, Eremostachys, Nepeta,
Perovskia, Phlomis, Stachys) and
Orobanchaceae (Pedicularis) (Shaposhnikov
1964b, as phlomicola; Narzikulov & Daniyarova 1990). Kadyrbekov (2014d) records it also from
plant genera in several other families, including Rhodiola (Crassulaceae) and Gentiana/Gentianella
(Gentianaceae), where there is possible confusion respectively with B. sedi and B. gentianae. In Central Asia, and there is a redescription from
Iran by Sedighi et al. (2018). This
species was erroneously synonymised with B. pilosus by Blackman &
Eastop (1994). Brachycaudus
(Appelia) cerinthis Bozhko Apterae are shining blackish, BL 1.6-2.1
mm. Alatae have secondary rhinaria distributed III 8-12, IV 0-2, V 0. On
stems, flower stalks and bracts, and deforming flower buds, of Cerinthe spp. In France, Italy, Czech Republic, Poland,
Hungary and Ukraine. Monoecious holocyclic on Cerinthe (Cocuzza et al.
2007). Brachycaudus
crassitibiae Nevsky Apterae are dark brown with paler
appendages; BL c.1.3-1.4 mm. Alatae have 4-6 secondary rhinaria on ANT III.
On Turgenia (~Caucalis) latifolia in
south Kazakhstan. Brachycaudus (Acaudus)
divaricatae Shaposhnikov Host-alternating between Prunus cerasifera (sometimes P. domestica or P. spinosa) and Caryophyllaceae (usually Silene latifolia s. lat.) in east Europe and the Middle
East. Apterae are shining dark brown
to black, BL 2.0-2.2 mm, rolling leaves of Prunus in spring. Spring migrant alatae have secondary rhinaria
distributed III 16-31, IV 0, but later generations have III 30-50, IV 3-4, V
0. The migration to Silene is
apparently facultative (Shaposhnikov 1962). In recent years this species has
spread north-westward to the eastern Baltic coast, where it has an
abbreviated life cycle, staying on P. cerasifera and producing sexual
morphs and eggs from late June (Rakauskas &
Juronis 2006, Rakauskas & Turčinavičiene 2006). It
has also now reached the Czech Republic, where its morphology and DNA have
been compared with eastern Baltic populations (Bašilova et al. 2012).
Havelka
et al. (2017) have compared its DNA
with that of the closely related B.
lychnidis. The male genitalia were described and illustrated by Wieczorek
et al. (2011). Native predators
attacking B. divaricatae in
Lithuania were studied by Danilov et al.
(2016). Brachycaudus
divaricatellus Shaposhnikov Apterae are shiny yellowish-rose, elongate
oval, BL c.2.0 mm. Alatae have
secondary rhinaria distributed III 30-50, IV 15-22, V 0. On
undersides of leaves of Prunus
cerasifera, causing slight curling of tips and discoloration. In south-west Turkmenistan (Shaposhnikov
1964a). Life cycle is unknown. Shaposhnikov (1964b) described the ovipara. Brachycaudus (Thuleaphis) eurotiae (Mamontova-Solukha) Apterae are shiny yellow, without wax; BL
1.1-1.6 mm. On Krascheninnikovia spp.,
living in galls formed by curling and thickening of the upper sides of the
leaves. Alatae have 5-7 secondary rhinaria on ANT III only, and dark dorsal markings. In Ukraine (Mamontova-Solukha, 1968, as Cryptosiphon eurotiae), Kazakhstan and
western China (Kadyrbekov et al.,
2002). Kadyrbekov (2002) erected a new genus (Scythaphis) for this species which has an unusually setose
abdominal tergite 8 and lacks marginal tubercles (and has a somewhat
distinctive biology), but
Stekolshchikov (2007) transferred it to Brachycaudus (Thuleaphis).
Brachycaudus
gentianae Daniyarova Apterae are shining brown; BL c. 1.8
mm. On Gentiana olgae in Tajikistan, and on Gentiana tianshanica in Kazakhstan (Kadyrbekov 2017a). Possibly
this is a synonym of B. cerasicola,
which is also recorded from Gentiana/Gentianella by Kadyrbekov (2014d). Brachycaudus
helichrysi (Kaltenbach) Plate 9i Apterae are very variably coloured (but
always pale); pale green, pale yellow, whitish or pinkish (see influentialpoints.com/Gallery);
BL 0.9-2.0 mm. Alatae have secondary rhinaria distributed III 13-46, IV 0-18.
On stems and in flowerheads of numerous plant species, esp.
Compositae/Asteraceae (e.g. Achillea,
Ageratum, Aster, Bidens, Chrysanthemum, Cineraria, Erigeron, Gnaphalium, Helianthus, Matricaria,
Senecio) and Boraginaceae (e.g. Anchusa, Cynoglossum, Myosotis, Symphytum), but also many others
including Rumex, Saxifraga, Trifolium, Veronica.
World-wide, and a major pest (for further information see Eastop & Blackman
2000, p.247). Heteroecious holocyclic with sexual phase on Prunus spp. (especially P. domestica, P. insititia, P. spinosa) in colder climates. Apterae
in spring populations on Prunus are
green, brownish or yellowish, shiny and slightly waxy, causing leaves to roll
up tightly perpendicular to the mid-rib.
BL is 1.4-2.0 mm on Prunus,
down to 0.9 mm on secondary hosts.
Darwish (1984) described the morphology of spring generations. Anholocyclic
populations occur in warmer regions and in glasshouses, and
are sometimes found on new growth of various trees. Molecular studies have now revealed that
populations identified as B. helichrysi
throughout the world comprise two forms that are sufficiently distinct to be
regarded as sibling species, genetically isolated from each other (Pifaretti et al. 2012, 2013a,b and
possibly Rebijith et al. 2013).
Host alternation to Prunus domestica
and related species only occurs in one of these forms (H1), in regions with a
continental climate. The second form (H2) is mainly anholocyclic,
and is mostly absent from the coldest regions, but has now been shown to have
a sexual phase on Prunus persica in
northern India (Pifaretti 2013b). Mixed populations of the two forms often
occur on herbaceous hosts, and no clear differences in host preference on
herbaceous plants between these two forms have so far been detected. Further
studies have revealed a third form (H3) restricted to East Asia, associated
with apricots and Prunus species
endemic to that region (Popkin et al.
2017). 2n=12. Brachycaudus
(Prunaphis) iranicus Davatchi
& Remaudičre (Fig.9a) Apterae are greenish yellow to pale green
with shiny brown dorsal markings (a transverse bar on the mesonotum, a central
abdominal patch and presiphuncular sclerites) and brown siphunculi; BL
1.6-2.0 mm. On undersides of leaves of
Anchusa azurea (= italica) and A. strigosa, feeding close to the main veins, in the Middle East
(Iran, Turkey, Israel, Lebanon and Syria). The alata (hitherto undescribed)
has dark antennae, 13-16 secondary
rhinaria on ANT III, and a central abdominal sclerite on abdominal tergites
5-6, fused each side with presiphuncular sclerites, and with a large central
lacuna. Monoecious holocyclic with alate
males (Davatchi & Remaudičre 1953). Brachycaudus
(Prunaphis) jacobi Stroyan Apterae are black and shiny dorsally,
olive green ventrally; BL 1.4-2.0 mm. Alatae have secondary rhinaria
distributed III 6-16, IV 0-2. At base of stem or on roots of Myosotis and Pulmonaria spp., ant-attended. In Europe (UK, Netherlands,
Germany, Italy). Monoecious holocyclic with apterous males (Müller 1975a).
2n=12. Brachycaudus
(Acaudus) klugkisti (Börner) Apterae are shining black dorsally, red-brown
ventrally; BL 1.5-2.5 mm. Alatae have secondary rhinaria distributed III
11-31 (in a row), IV 0-8. On upper parts of Silene (incl. Melandrium) spp. In
Europe (UK, Scandinavia, Finland, Germany, Poland, France, Italy). Monoecious
holocyclic with apterous males (Müller 1987). 2n=10. Brachycaudus (Nevskyaphis) lamii Koch
Apterae have a shining black dorsum; BL 1.8-2.2 mm. On Lamium spp., especially L. album, in Europe (Germany, Poland, Switzerland, Italy, Slovakia).
Monoecious holocyclic, with oviparae in October (BMNH collection, leg. D.
Hille Ris Lambers). Molecular data (Jousselin et al. 2010) indicate that this
is probably a synonym of B. ballotae.
Brachycaudus
(Prunaphis)
lateralis (Walker)
Apterae with a shining black dorsum, ventrally green or reddish; BL
1.6-2.6 mm. On stems and leaves, usually close to ground, of plants in
numerous genera of Compositae including Arctium,
Anthemis, Carduus, Cirsium, Chrysanthemum, Galinsoga, Leucanthemum, Matricaria and Senecio. Also found on Capsella.
In Europe, eastward to Ukraine. Mainly
anholocyclic, although a sexual phase may sometimes occur on Prunus (Müller & Horatschek 1979).
It is closely related to B. cardui,
often treated as a subspecies, and DNA studies (Jousselin et al. 2009, 2010) indicate that it
may be a variant form of that species. Brachycaudus
(Scrophulaphis)
linariae Stroyan
Apterae are deep blackish green, shiny, immatures being paler green
(see influential
points/gallery); BL 1.4-1.9 mm. Alatae have secondary rhinaria
distributed III 11-37, IV 5-10, V 0-2. On basal parts of Linaria spp., attended by ants. In UK, Denmark, Sweden, Finland,
France, Germany, Italy and most of eastern Europe (Andreev & Mamontova
1998). Monoecious holocyclic; Patti & Barbagallo (1997) described
oviparae and apterous males collected from L. purpurea in Sicily in October. Brachycaudus (Nevskyaphis)
lucifugus Müller
Apterae are yellowish green with shiny dark brown to black dorsum; BL
1.3-2.2 mm. Alatae have secondary rhinaria
distributed III 15-30, IV
0-9, V 0-1(-5). On roots and at leaf bases of Plantago lanceolata, in Europe (UK, Germany, Hungary, Italy).
Ant-attended. Monoecious holocyclic with apterous males (original description
and Stroyan 1964a). Brachycaudus (Acaudus) lychnicola Hille Ris Lambers
Apterae are reddish brown with shining black dorsal shield; BL 1.9-2.3
mm. Alatae have secondary rhinaria
distributed III (15-)
23-36, IV 0-4. On roots, stem-bases and lower leaves of Lychnis
flos-cuculi and Silene (incl. Melandrium)
spp., attended by ants. In north-west Europe (Netherlands, Sweden). Monoecious holocyclic with apterous males
(original description). Molecular data (Jousselin et al. 2010) indicate that, if the specimens from Czech Republic
used in their analysis are correctly identified as this species, then it is
probably a synonym of B. lychnidis. Brachycaudus (Acaudus) lychnidis (L.) Apterae
are reddish brown with shining brown-black dorsal shield (see influentialpoints.com/Gallery); BL 1.8-2.9 mm. Alatae
have secondary rhinaria distributed III 15-34, IV 0-4. On stems, leaves and flowers of Lychnis and Silene (incl. Melandrium)
spp. Europe, eastward to west Siberia, Turkey,
Iran, Caucasus and Kazakhstan. Monoecious
holocyclic with apterous males (Heie 1992). 2n=12. Brachycaudus
(Nevskyaphis) malvae Shaposhnikov Apterae are shining blackish green (see influential
points/gallery); BL 1.8-2.3 mm. Alatae have secondary rhinaria
distributed III 17-31, IV 0-3, V 0. On Malva
spp., in ant-attended colonies at base of stem and on lower leaves. In
England, Spain, southern Russia, Ukraine, and also China, if Brachycaudus atuberculatus Zhang 1981
is a synonym (see Andreev 2004). 2n=12. Brachycaudus
(Scrophulaphis) mimeuri Remaudičre Apterae are shining pale to dark brown;
BL 0.7-1.4 mm. Alatae have secondary rhinaria distributed III 12-33, IV 1-10,
V 0-1. On roots of Euphrasia and Odontites spp. in southern and eastern
Europe; for distribution see Jörg & Lampel 1988, but also in Denmark
(Heie 1992) and Sicily (Barbagallo & Stroyan 1982), and possibly also in
South Australia on Parentucellia
latifolia (Eastop 1966, as ?B .
persicaecola). Not attended by ants (Hille Ris Lambers 1967). Presumably
anholocyclic, and closely related to B.
persicae (q.v.). Brachycaudus
(Prunaphis) mordvilkoi Hille Ris
Lambers Apterae are green with shiny
black dorsal shield; BL 1.7-2.3 mm. Alatae have secondary rhinaria
distributed III 12-21, IV 3-6, V 0-1. On certain Boraginaceae (Anchusa, Echium, Solenanthus,
Symphytum), with one record from Compositae (Hieracium pilosella). Widely
distributed in Europe (but not recorded from British Isles, Scandinavia or
Iberian peninsula) and in Middle East (Turkey; Şenol et al. 2017). Monoecious holocyclic
with apterous males (Müller 1975a). Brachycaudus (Brachycaudina) napelli (Schrank)
(Fig.6b,c) Apterae are shining black with end of
abdomen red, immatures are brown; BL 2.0-2.9 mm. Alatiform apterae occur with
3-25 secondary rhinaria on ANT III, alatae have 17-37 on III and 0-4 on IV.
On Aconitum spp., esp. napellus, forming colonies on
stems, flowers and fruits, and sometimes on leaves. Only occasionally on Delphinium spp. In west and north-west Europe (not UK), south to Spain, and
also recorded from Austria and Czech Republic (Cocuzza et al. 2010). Closely related
to B. aconiti (q.v.). Monoecious holocyclic with apterous males.
Brachycaudus
(Acaudus) pallidus Andreev Apterae are shining dark brown; BL
1.5-2.2 mm. On roots of Silene nutans in
Moldova. Brachycaudus (Scrophulaphis) persicae group Apterae have a shiny dark brown or black
dorsum, and black siphunculi (see influential
points/gallery); BL 1.5-2.2 mm. Alate spring migrants have secondary rhinaria
distributed III 23-51, IV 9-21, V 1-6. B.
persicae
(Passerini) s.str. is most commonly found in large spring colonies on young
stems of Prunus persica or P. armeniaca, often persisting into
the summer on root suckers. Alatae from Prunus
have been transferred successfully to Euphrasia
sp. and Rhinanthus glaber and
subsequently to Melampyrum pratense (Burger
1975), so it seems probable that there is a facultative host alternation from
Prunus to Orobanchaceae
(Euphrasia, Melampyrum, Rhinanthus),
at least in Europe, but anholocyclic overwintering is also common on Prunus roots. On
Orobanchaceae it lives on the above-ground parts (Heie 1992). Aphids
commonly regarded as this species are also widely distributed on peach
outside Europe, in the Middle East, Central Asia, southern Africa, Australia,
New Zealand, North and South America. Similar aphids also occur commonly in
Europe on P. domestica and P. spinosa, and occasionally on Pyrus communis and Chaenomeles japonica. These all form a complex of which the host
relationships and life cycles need further clarification. Müller & Steiner (1988a) applied the
name B. semisubterraneus Börner to
anholocyclic populations on plum in Germany, and suggested that the form
distributed world-wide on peach was also an anholocyclic species, to which
they applied the name B. persicaecola
(Boisduval). However, aphids of the B. persicae group are recorded from
Orobanchaceae in both Australia and Chile (BMNH collection), indicating that
some host alternation occurs in southern continents, and Burger (1975) showed
that not all populations on plum and peach are anholocyclic, so the
application of the names semisubterraneus
and persicaecola is still unclear. There are also records from Odontites, but these may be referable to B. mimeuri (q.v.), a closely-related anholocyclic form on roots
of Orobanchaceae that has also become widely distributed. 2n=10. Brachycaudus
(Mordvilkomemor) pilosus (Mordvilko ex Nevsky) Apterae are black with pale appendages and
cauda; BL 1.1-1.7 mm. Alatae have secondary rhinaria distributed III
16-23, IV 6-8, V 0-1. On various Prunus spp.,
especially P. ulmifolia (= P.
triloba), damaged leaves of which
develop gall-like thickenings and red spots, and P. bucharica, the
leaves of which are only slightly curled and reddened (Narzikulov 1965c). Possibly
there is no host alternation; populations on P. prostrata in Pakistan
included apterous males and oviparae in May together with fundatrices, and
apterous and alate viviparae, and additional sexual morphs were collected in
August (Naumann-Etienne & Remaudičre 1995). In mountainous regions of
central Asia (Kazakhstan, Kirgizia, Tajikistan, Turkmenistan), Iran, India
and Pakistan (BMNH collection). Brachycaudus (Nevskyaphis) plantaginis Holman & Szelegiewicz Apterae are yellowish brown with shiny dark
dorsum; BL c.1.9-2.1 mm. Alatae are undescribed. Ant-attended, on root collar
of Plantago
depressa and P. major in Mongolia and east
Siberia (Pashchenko 1988a). Brachycaudus (Acaudus)
populi (del
Guercio) Apterae are red-brown with
shiny black dorsal shield; BL 1.6-2.9 mm. Alatae have secondary rhinaria
distributed III 19-36, IV 0-3. On upper parts of Silene
spp., widely distributed in Europe, eastward to Crimea, Russia (Asltai)
and Kazakhstan. Monoecious holocyclic with apterous and alate males. Closely
related to B. lychnidis, and
confused with that species prior to Burger (1975). 2n=12. Brachycaudus (Appelia) prunicola
(Kaltenbach) Apterae are shiny black,
dark green or deep yellow-green with black dorsal cross-bands, immatures
grey-green (see influential
points/gallery); BL 1.4-2.4 mm. Alatae
have secondary rhinaria distributed III 23-43, IV 7-16, V 0-3. In
spring on new growth of Prunus (in
Europe usually P. spinosa,
sometimes P. domestica or P. insititia), causing severe leaf
curl and discoloration.
Host-alternating between Prunus
and Tragopogon, but part of the
population - possibly a subspecies - may live without alternation on P. spinosa. In continental Europe, Siberia
(Stekolshchikov et al. 2008) east to Irkutsk (Stekolshchikov &
Shaposhnikov 1998), Iran (Rezwani & Radjabi 1986), Kazakhstan (Kadyrbekov
2005c) and Pakistan (Baluchistan; Naumann-Etienne & Remaudičre 1995).
[Note: the sample (no. 992) identified as B.
prunicola from Brittany used in DNA analysis by Jousselin et al. (2010), and branching
separately from the rest of the prunicola/schwartzi/tragopogonis group in their phylogeny, was B. prunifex (q.v).] 2n=12. Brachycaudus
(Appelia) prunifex (Theobald) Apterae are shiny dark green to
black (see influential
points/gallery); BL 1.5-2.4 mm. Alatae have secondary rhinaria distributed III
22-41, IV 2-13, V 0(-2). Colonies occur all-year-round on Prunus spinosa, in spring causing severe leaf curl and discoloration to
new growth. There is also a record from P.
cerasifera var. atropurpurea. Suggestions
of a facultative host alternation to Tragopogon
do not appear to be substantiated. Alate males have been collected in
September (BMNH collection), but oviparae are not yet recorded. In England,
Wales, Ireland, and northern France. This species has been synonymised
previously with B. prunicola (Kaltenbach),
which occurs on the same host throughout the rest of Europe and in central
Asia, but it can be distinguished from that species (and from B. schwartzi) by the longer hairs on
the antennae and anterior abdominal tergites (Blackman 2010). 2n=12. Brachycaudus
(Acaudus) rinariatus Andreev Apterae have a shining black dorsal
shield; BL 1.7-2.2 mm. Alatae have secondary rhinaria distributed III 36-48,
IV 16-19, V 1-6. On Linaria vulgaris in
eastern Europe (Russia, Moldova, Ukraine), and on L.× incompleta in Kazakhstan (Kadyrbekov, 2017a, as B. himalayensis). Brachycaudus
(Brachycaudina) rociadae (Cockerell) (Fig.6a) Apterae are shining reddish brown to
black, fading to hazel brown anteriorly and posteriorly; BL c. 2.0 mm.
Immatures are bright coral red. Alatae have 2-13 secondary rhinaria on ANT
III only. On stems and leaves of Delphinium
spp.; feeding on leaves causes them to curl tightly (Palmer 1952) . Widely distributed in North America.
Monoecious holocyclic, with oviparae and apterous males in mid-July (Ontario;
BMNH collection, leg. S.F. MacDonald) or early October (Colorado and Manitoba).
Brachycaudus
(Thuleaphis) rumexicolens (Patch) Apterae are reddish with variably
developed dark dorsal markings; BL 1.3-2.1 mm. Alatae have more-or-less fused
cross-bands on posterior abdominal tergites, 6-22 secondary rhinaria on ANT
III, and 0(-1) on IV. In inflorescences of
Rumex spp., especially acetosella, and sometimes on other Polygonaceae (Fagopyrum, Persicaria,
Polygonum. Rheum). In Europe,
north Africa (Tunisia: Boukhris-Bouhachem et al. 2007), west and Central Asia, north-east Siberia
(Chukotka: Stekolshchikov & Khruleva 2015), India, Australia, North and
South America. Monoecious holocyclic, with apterous males (Tuatay &
Remaudičre 1964). B. brevirostratus in
east Siberia might be this species. 2n=12. Brachycaudus
salicinae Börner Apterae are shining grey-black; BL 1.4-1.9
mm. Alatae have secondary rhinaria distributed III 16-38, IV 3-13, V 0. In
rolled leaves of Inula spp.,
especially salicina, and also Pulicaria dysenterica (Holman 2009);
in Europe (Germany, France, Poland, Czech Republic, Slovakia), Iran (Rezwani
& Parvizi 1990) and Central Asia (Kazakhstan; Kadyrbekov &
Aoitzhanova 2005). Monoecious holocyclic (Börner 1952). Brachycaudus (Appelia) schwartzi
(Börner) Apterae of BL 1.4-2.1 mm, shiny
yellow-brown to dark brown with extensive dorsal black sclerotization;
immatures are yellow-brown. Alatae have
secondary rhinaria distributed III 23-36, IV 6-17, V 0(-1). Living without host alternation on Prunus persica; spring colonies cause
severe curling and distortion of peach leaves. In Europe, Iran, Kazakhstan,
North Africa (Tunisia, Egypt), India, South America and California. Records from other Prunus spp. with the possible exception of P. serotina, may be referable to other species. Records of B. tragopogonis from peach in the literature are referable to
this aphid. Darwish (1983) gave
descriptions of all developmental stages. A morphometric and molecular study
by Cocuzza et al. (2007) indicated
that this species should perhaps be treated as a subspecies of B. prunicola. (See also Blackman &
Eastop 2000, p.249). 2n=12. Brachycaudus
(Thuleaphis) sedi (Jacob) Apterae are yellowish green, with dark
cross-bands on posterior abdomen; B: 1.0-2.1 mm. Alatae have secondary
rhinaria distributed III 11-19, IV 0-5, V 0(-1), VI base 0(-1). On stem
apices and flowers of Rhodiola (= Sedum) rosea, causing distortion. There is
also a record from Sedum anacampseros
(Cocuzza et al. 2010). In Europe (including Iceland, as Thuleaphis acaudata; Prior & Stroyan 1960) and west Siberia (Ivanoskaya 1975, as
Rhodiolaphis cholsunensis).
Monoecious holocyclic, with oviparae and apterous males appearing in August
(original description). 2n=8*. Brachycaudus
shaposhnikovi Narzikulov Apterae are shining yellowish green with
black dorsal sclerotic markings, siphunculi pale with tips darker; BL 2.1-2.7
mm. Alatae have 15-18 secondary rhinaria on ANT III. Monoecious holocyclic on
Atraphaxis spp. in Tajikistan (Narzikulov & Daniyarova 1990) and Kazakhstan
(Kadyrbekov 2014e). Brachycaudus
spiraeae Börner Apterae are light green to greyish brown
with variable dark dorsal markings; BL 1.2-1.8 mm. Apterae have secondary
rhinaria distributed III 0-11, IV 0-1, and alatae have them distributed III
19-34, IV 4-13, V 0-2. On Spiraea spp.,
in leaves rolled and curled into narrow “pods”. In Europe, west Siberia, Iran
(Rezwani 2010) and Central Asia.
Generally regarded as monoecious holocyclic on Spiraea, but molecular work has
revealed a close relationship with H3 populations of B. helichrysi, possibly involving introgression, and implying
some degree of migration from Spiraea
to herbaceous hosts (Popkin et al.
2017). 2n=12. Brachycaudus
(Appelia) tragopogonis (Kaltenbach) Apterae are shining grey-brown to dark
brown with dark dorsal sclerotic
markings (see influential
points/gallery); BL 1.4 –2.3 mm. Alatae have
secondary rhinaria distributed III 27-49, IV 6-22, V 0-4. On aerial parts of Tragopogon spp. and Scorzonera spp. in Europe, west
Siberia, south-west and Central Asia (Kazakhstan; Kadyrbekov 2005c, 2009a),
and introduced to western USA and South America (Brazil, Chile). Monoecious
holocyclic. In western Europe there is possible confusion with the
morphologically similar B. prunicola,
which has a sexual phase on Prunus spinosus. Hille Ris Lambers (1948) described a
subspecies with long dorsal hairs, B. tragopogonis
ssp. setosus, on T. longirostris in Israel, and
collections from south-west and Central Asia and Pakistan may be of this form
(Mostafawy 1967), which is also reported from Hungary (Basky 2015). 2n=12
(2n=11 and 12 for samples of ssp. setosus
from Israel and Iran respectively). Brachycaudus umbelliferarum Nevsky
Apterae are pale brown or yellowish green with dark brown pleural and
marginal spots, or entirely dark brown; BL c. 2.4 mm. On roots of an
umbelliferous plant, probably Heracleum
sp. south-east Kazakhstan. With marginal tubercles on prothorax,
abdominal tergites 1, 2 and 7, this species is unlikely to be a Brachycaudus, and is regarded by
Andreev (2004) as a nomen dubium. Brachycaudus (Nevskyaphis) virgatus Shaposhnikov Colour of apterae in life is unknown,
probably greenish with darker head and dorsal markings; BL 1.6-1.8 mm (2
specimens). On stems and leaves near
base of host plants, which are Boraginaceae (Anchusa, Symphytum) and the composite Senecio fluviatilis (= nemorensis). Alatae have not been described.
Only known from southern Ukraine.
Brachycaudus
viridanus (Nevsky) Apterae are pale green, BL 1.3-1.7 mm. On
stems of Cousinia sp. in
Uzbekistan. Alatae are also pale (brownish head and thorax and pale green
abdomen), and have about 7 rhinaria on ANT III. This species was synonymised
by with B. cardui (presumably ssp. turanica), and could possibly be a
dwarf form of that species lacking dark sclerotisation.
Four or five species with very
short broad-based conical or barrel-shaped flangeless siphunculi and
triangular cauda associated with Caryophyllaceae. Heie (1992) and Blackman
(2010) reviewed the species in western
Europe. Brachycolus
aestivus Zhang, Qiao & Zhang Colour of apterae in life unrecorded; BL
c.1.5 mm. On Bupleurum chinense in
Shanxi province, China (L. Zhang et al.
2008). To judge from the host association this species might belong in Semiaphis. Brachycolus
brachysiphon Richards Colour of apterae in life unobserved; BL
c. 1.8 mm. On Stellaria longipes in
northern Canada. Brachycolus
cerastii (Kaltenbach) (Fig.51b) Apterae are dirty green, wax-powdered; BL
1.3-1.7 mm. Alatae have 3-6 secondary rhinaria on ANT III. On Cerastium spp., esp. C. arvense, causing shoots to be
stunted and deformed into gall-like structures. Apparently widely distributed
in Europe, but rarely collected. Also recorded from Kazakhstan on C. davuricum and C. tianschanicum (Kadyrbekov 2014e, 2017a). Monoecious holocyclic
(but sexual morphs are apparently undescribed). 2n=14. Brachycolus
cucubali (Passerini) Plate 13d Apterae are whitish yellow to pale green,
wax-powdered; BL 1.4-2.2 mm. Alatae have secondary rhinaria distributed III
9-16, IV 0-2. On Silene spp.,
rolling leaves into elongate pseudogalls (Heie 1992, as Hayhurstia). Widely distributed in Europe, in eastern Siberia and
Kazakhstan and in north Africa. In Kazakhstan it is also recorded from Oberna and Cerastium (Kadyrbekov 2017a). Monoecious holocyclic; Casiraghi et al. (2020) studied the life cycle
on Silene vulgaris in northern
Spain, redescribed the aptera and alata, and described oviparae and apterous
males found in the pseudogalls in November to January. Apterous viviparae
also live through the winter in the galls along with the overwintering eggs. Brachycolus
stellariae (Hardy) Apterae are pale green, wax-powdered; BL
1.1-2.0 mm. Alatae have secondary rhinaria distributed III 4-9, V 0-1. On
upper sides of leaves of Stellaria spp., which are rolled into
oblong pseudogalls. Also recorded from Moehringia
trinervia. In Europe. Monoecious holocyclic with oviparae and very
small apterous males produced in September (Blackman 2010).
Two palaearctic species with
different biologies, similar to Brachycolus
but with shorter antennae and terminal processes, and very short conical
siphunculi. Brachycorynella
asparagi (Mordvilko) Apterae are green, covered with grey
mealy wax; BL 1.2-1.8 mm. On Asparagus spp.,
causing stunting of shoots and “rosetting” of leaves, which turn blue-green.
In Europe (not yet in UK, but recently expanding its range), North Africa
(Tunisia; Boukhris-Bouhachem et al. 2007), south-west and Central
Asia (Kazakhstan; Kadyrbekov & Aoitzhanova 2005), China, and introduced
into North America (see also Blackman & Eastop 2000). Monoecious holocyclic with alate males.
2n=10. Brachycorynella
lonicerina (Shaposhnikov) (Fig.38c) Apterae are green, dusted with greyish wax
powder, BL c.1.8 mm. On Lonicera spp.,
feeding on upper sides of leaves, which are folded in half (Shaposhnikov,
1964, as Semiaphis). In Central
Asia, southern Russia, Ukraine, Iran and Pakistan (Naumann-Etienne &
Remaudičre 1995). Heteroecious holocyclic but with with an unknown secondary
host (Remaudičre & Remaudičre 1997, p. 298). Oviparae and alate males
appear on Lonicera in late
September-October (Narzikulov 1965b; Mukhamediev & Akhmedov 1982).
2n=10*.
One species in India with a Myzus-like head but with thick
truncate siphunculi and an unusual cauda. Brachymyzus
jasmini A.N. Basu Apterae have head, thorax and antennae
blackish brown, abdomen greenish, BL 1.6-2.1 mm. On leaves and young shoots
of Jasminum humile in north-east India (West Bengal).
Oviparae were described from Sikkim on Nellia
(Neillia?) sp. (Rosaceae), and an alate male was
found on Pilea microphylla (Mondal et al. 1978). There is possibly host
alternation between Neillia and Jasminum.
One or two species of
grass-feeding aphids in east and south-east Asia, with distinctive very short
black siphunculi and a long dark rather hairy cauda. Brachysiphoniella
apiaca Zhang & Zhang Colour of apterae in life is unrecorded;
BL c.1.6 mm. On Apium graveolens in
China (L. Zhang & G. Zhang 2000a). The host needs confirmation, as this
aphid is very similar to the grass-feeding B. montana. Brachysiphoniella
montana (van der Goot) Plate 14d Apterae are brownish green, densely
covered in mealy wax (see aphids
of Karnataka website); BL 1.1-1.9
mm. On grasses, well-adapted to aquatic species such as Leersia hexandra due to its waxy coat. Also recorded from Cynodon, Eleusine, Microstegium,
Miscanthus, Oryza (as Melanaphis
vandergooti Raychaudhuri & Banerjee), Panicum, Pennisetum and
Phragmites. India, Bangladesh,
Nepal, China, Taiwan, Vietnam, Korea, Japan, Philippines, Malysia, Indonesia,
and Australia (NSW). Presumably mainly anholocyclic, but at least in Korea
there may be host alternation to Pyrus spp.
(Lee et al. 2002c).
About 40 species mainly on xerophytic plants, mostly Amaranthaceae.
Related to Aphis but with a very
short antennal terminal process (usually shorter than in Aphis subg. Protaphis) and
very short, often pale siphunculi. The clypeus is often globose, but the
extent of its enlargement varies considerably, and subgeneric separation of Xerophilaphis on grounds of greater
enlargement of clypeus (Remaudiere & Halbert 1996) does not seem warranted.
Many palaearctic species and one described from South America are coated in
mealy wax in life, but wax production is not reported for North American
species. Kadyrbekov (1999c) reviewed and keyed the species in Central Asia,
Kadyrbekov (2001c) summarised the characters of the genus and transferred
several species to it, and Kadyrbekov (2014c) described several more species
from Kazakhstan. Nieto Nafría et al.
(2005a) reviewed the two species on the Iberian peninsula and provided lists
of synonymies. Despite this the genus is still poorly known; many of the
nominal species are very similar to one another and their separate identity
needs experimental confirmation.
Brachyunguis
agriphylli Bozhko Apterae are green, wax-powdered, with
dark green pleural markings; BL c.1.5-1.6 mm. In large colonies on Agriophyllum squarrosum (= A. arenarium)
in Ukraine. Not clearly distinct from B.
harmalae. Brachyunguis
armelliae Kadyrbekov Apterae are greenish with brown head,
without wax powdering; BL 1.5-1.7 mm. The single known alata has 7-8
secondary rhinaria on ANT III. On inflorescences of Nitraria schoberi, ant-attended, in south-east Kazakhstan
(Kadyrbekov 2014c). Brachyunguis
atraphaxidis (Nevsky) Apterae are dark green, with paler
appendages, body wax-dusted; BL c.1.6-1.7 mm. In small colonies on leaves and
flowers of Atraphaxis spp.,
attended by ants. Recorded from Iran, Georgia, Turkmenistan,Uzbekistan,
Tajikistan Kyrgyzstan and Kazakhstan (Kadyrbekov 2017a). Brachyunguis
bahamondesi Remaudičre
& Halbert Apterae are pale, with
apices of tibiae, tarsi and siphunculi black, body colour in life unrecorded,
BL 1.0-2.2 mm. On Amaranthaceae (Chenopodium,
Atriplex) in Argentina. Brachyunguis
berezhkovi (Ivanoskaya) Apterae are pale green, pruinose, with
pale appendages; BL c.1.4-1.6 mm. In small ant-attended colonies at shoot
terminals of Salsola richteri in
Central Asia (Turkmenistan). Brachyunguis
bicolor Ivanoskaya Apterae are grey with darker patches
laterally, wax-dusted; BL c. 1.3 mm. On Cynanchum
acutum, singly or in small colonies on stems, flowers and flower stalks,
attended by ants. In Ukraine (as B.
cynanchiacuti Bozhko),
Turkmenistan and Kazakhstan. Brachyunguis bishopi Remaudičre &
Halbert Apterae are pale, colour in
life unrecorded, BL 1.2-2.3 mm. On terminal parts of Sarcobatus
vermiculatus in western USA. Oviparae
and apterous males occur in October (original description).
Brachyunguis
blanchardi Remaudičre
& Bahamondes Apterae are dark
grey, wax-dusted, BL 1.6-2.0 mm. On Chenopodium
(=Allenrolfea) vaginata in Argentina. The clypeus is
greatly enlarged in this species, indicating specialisation on Amaranthaceae. Brachyunguis
bonnevillensis Knowlton Apterae are pale bluish green, with pale
appendages (see aphidtrek.org);
BL 1.0-2.1 mm. Alatae have a shiny black head and thorax. In small colonies
on leaves, flower stems and new shoots of Sarcobatus
vermiculatus. It is well camouflaged on lower surfaces of leaves, and has
a dwarf form in summer (Palmer 1952). Also recorded from Atriplex canescens (Remaudičre & Halbert 1996), although
there is possible confusion with B.
tetrapteralis (q.v.). In western USA and Mexico. Brachyunguis
brevisiphon Kadyrbekov, Renxin & Shao Apterae are greenish with a slight grey
wax film; BL c. 1.2 mm. In ant-attended colonies on shoots of Tamarix
spp. in China (Xinjiang-Uygur region; Kadyrbekov et al. 2002), and subsequently recorded from Kazakhstan
(Kadyrbekov 2017a). The description does not provide any clear differences
from B. tamaricophila, and the key character that we have used to
distinguish this species may not be reliable. Brachyunguis
cahuille (Dickson) (Fig.36g) Apterae are dull dark green, BL 0.9-1.4
mm. On leaves of Suaeda spp. in
southern California, Arizona and Mexico (Remaudičre & Halbert 1996; as B. tetrapteralis). Brachysiphon
calligoni (Nevsky) Apterae are pale green, covered in wax
meal; BL 1.3-1.7 mm. On young shoots of Calligonum
spp. in Uzbekistan and Kazakhstan (Kadyrbekov 2017a). Brachyunguis
calotropicus Menon & Pawar Apterae are pale to muddy green, with
sparse wax powder, BL 1.2-1.5 mm. In
dense aggregations on upper or undersides of younger leaves of Calotropis spp. (gigantea, procera), sometimes attended by ants. The life cycle is
unknown. First recorded from the Delhi
region of India without a proper description (Menon & Pawar 1958). B.
ushari Müller & El Tigani 1986 seems to be a synonym, and the present
account is based on that assumption.
There is also a possibility that B. calotropicus may prove to
be a synonym of B. harmalae. In India, south-west Asia (Saudi
Arabia) and Africa (Algeria, Egypt, Sudan).
2n=8 (Kurl 1978). Brachyunguis
convolvulisucta Zhang, Chen, Zhong & Li Appearance in life is unrecorded; BL
c.1.5-1.6 mm. On Convolvulus arvensis
in Xinjiang province, China (G. Zhang 1999). Brachyunguis
cuscutae (Nevsky) Apterae are green, densely covered with
wax; BL 1.2-1.5 mm. Found in immense numbers on flowers and stalks of Cuscuta spp. in early November
(Ivanoskaya 1960). In Uzbekistan, and also recorded from Iran (Rezwani et al. 1994) and Kazakhstan
(Kadyrbekov 2014e). Brachyunguis
cynanchi (Nevsky) Apterae are dull red to dark violaceous,
coated with fine mealy wax; BL 1.2-1.8 mm. On Cynanchum acutum in Uzbekistan, found in large numbers on apical
parts of stems, flower stalks and upper sides of leaves (Ivanoskaya 1960). Also
recorded from Iran (Rezwani 2010) and Kazakhstan (from C. sibiricum; Kadyrbekov
& Aoitzhanova 2005). Brachyunguis
dendrostellerae Kadyrbekov Apterae are greenish without wax
powdering: BL 1.3-1.4 mm. On flowers
and stems of Diarthron (= Dendrostellera) spp. in south-east
Kazakhstan, ant-attended (Kadyrbekov 2014c). Possibly specimens with dark
siphunculi collected on Diarthron
by G. Remaudičre in Iran (BMNH collection) are this species. Brachyunguis flexosiphon Kadyrbekov Apterae are
dark green with red eyes; BL 1.5-1.8 mm. Described from the inflorescence of
an unidentified umbelliferous plant (Kadyrbekov 1999c), and subsequently
recorded from Zosima tordyloides
(= Semenovia rubtzovii), Ferula spp., Schrenkia
?golickeana and Bunium setosum (= setaceum?) (Kadyrbekov 2014d, 2017a). In Kazakhstan. Brachyunguis
harmalae Das (= plotnikovi Nevsky) Plate 7h, fig.36h Apterae are mid- to deep green
anteriorly, more yellowish on
posterior abdomen, thickly dusted with wax powder, and with two longitudinal
rows of submarginal dark green spots; BL 1.2-2.0 mm. Alatae have 3-7 secondary rhinaria on ANT
III. Often
ant-attended. Monoecious holocyclic on Peganum
harmala in Pakistan, where Das (1918) studied its life cycle in detail,
and on Calligonum in Central Asia, where damaging populations have
been reported from cotton (Nevsky 1928a, as plotnikovi). Both apterous
and alate males occur. The populations
that occur in the Middle East, Africa (Tunisia, Eritrea, Kenya, Sudan,
Senegal) and on the Iberian peninsula (Nieto Nafría et al. 2005a), colonising various, mostly xerophytic plants
including Calligonum and Calotropis, are presumably anholocyclic.
Eastop & Raccah (1988) listed 12 hosts in 7 botanical families in
the Arava valley, Israel. An outbreak
is also recorded on Citrus paradisi
(Swirski 1963). B. flavidus (Ivanoskaya), described from Peganum harmala in Turkmenistan, is not clearly distinct. 2n=8. Brachyunguis
kaussarii Remaudičre & Davatchi Colour of apterae in life is unknown; BL
1.3-1.6 mm. Alatae have 7-9 secondary rhinaria on ANT III. On Stocksia brahuica (recorded as “bralmica” in error; G. Remaudičre,
pers. comm.) in Iran. Brachyunguis
letsoniae Das Apterae are green, covered in white wax
powder, BL c. 1.85 mm. Alatae have dark cross bars on abdominal tergites 6-8,
and 6-8 rhinaria on ANT III. Feeding on young growth and inflorescences of “Letsonia scandens” (possibly this is Lettsomia (= Argyreia) splendens?),
in Pakistan in October. It has not been reported since its original
description. Brachyunguis
lycii (Nevsky) Apterae are pale green coated with mealy
wax; BL 1.2-1.6 mm. In large colonies on leaves and fruits of Lycium spp.in Central Asia. Specimens conforming to Nevsky’s
description have also been found on Lycium
in Cyprus, Egypt and Iran (BMNH colln), although it is unclear how this
species differs from B. harmalae
colonising various other plants in these regions. 2n=8*. Brachyunguis
monstratus Kadyrbekov Apterae are dark green with dark red
eyes; BL 1.0-1.3 mm. In inflorescences of Atraphaxis
spp., in east and
south-east Kazakhstan and in Xinjiang, China (Kadyrbekov 1999c, 2004b, 2014c). Brachyunguis
nevskyi (Kreutzberg)
Apterae are dark green; BL c. 1.15 mm. On Pistacia vera in south-west and central Asia, colonising
inflorescences in spring and new shoots, petioles and leaves in autumn
(Ivanoskaya 1960, as Xerophilaphis). Brachyunguis
nurikamalae Kadyrbekov Apterae are greenish with brown head,
without wax powdering: BL 1.5-2.0 mm. On inflorescences of Ziziphora sp. in northern Kazakshtan,
attended by ants (Kadyrbekov 2014c). Alatae have 5-9 secondary rhinaria on
ANT III. Brachyunguis
peucedani (Nevsky) Apterae are pale green covered with wax;
BL 1.6-1.8 mm. On flowers and stems of Peucedanum
pseudoreoselinum in Uzbekistan, and recorded from Ferula spp. in Kazakhstan by Kadyrbekov (2003c). To judge from
the descriptions by Nevsky and by Ivanoskaya (1960), placement of this
species in Semiaphis by Eastop
& Hille Ris Lambers (1976) and Remaudičre & Remaudičre (1997) seems
to have been in error. Brachyunguis
poacynophila Zhang, Chen,
Zhong & Li Appearance of aptera
in life is unrecorded; BL c.1.4 mm. On Apocynum
venetum in north-west China (G.
Zhang 1999). Brachyunguis
rhei (Nevsky) Apterae are yellowish brown; BL c. 2 mm.
On Rheum maximoviczii, forming
large colonies on undersides of leaves, and subsequently recorded from
several Rumex spp. In southern
Kazakhstan and Turkmenistan. Brachyunguis
salsolacearum (Nevsky) Apterae are green, covered with fine
pruinose secretion, siphunculi and cauda green, dorsal abdomen with small
dark green spots; BL 1.3-1.9 mm. In large numbers on young growth and stems
of various Amaranthaceae (Anabasis,
Girgensohnia, Halocharis, Salsola)
in Central Asia (Narzikulov & Daniyarova 1990). Brachyunguis
saxaulica (Nevsky) Apterae are brownish yellow, greenish or
dark brown, covered with fine wax; BL 1.6-2.1 mm. At ends of young shoots of Haloxylon spp., forming “cone-like
pseudogalls”, ant-attended. In Kazakhstan, Tajikistan and Turkmenistan
(Ivanoskaya 1960, as Xerophilaphis),
and also recorded from Iran (Rezwani 1987). Brachyunguis
shaposhnikov Ivanoskaya Apterae are green, tinged with orange near siphunculi, without wax; BL
c.2.2 mm. Described from a plant of the Polygonaceae, and recorded from Polygonum aviculare by Kadyrbekov
(2017a). In western Kazakhstan. Brachyunguis
skafi Remaudičre & Talhouk Apterae are very dark, blackish; BL
1.2-1.9 mm. Recorded from three species of Astracantha (previously all three have been placed in Astragalus) in Turkey, Iran and
Lebanon. Monoecious holocyclic with oviparae and apterous males in
September-October (Remaudičre & Talhouk 2000). Brachyunguis
suaedus (Paik) The aptera described from Suaeda glauca in Korea by Paik (1965,
as Hyalopterus) is Clypeoaphis suaedae, and the alatae
may be vagrants from another plant . The length of the antennal terminal
process and black pigmentation suggest that it does not belong in Brachyunguis, where it is currently
placed. Brachyunguis
tamaricis (Lichtenstein)
Plates 13a, b,
fig. 113B Apterae according to the original
description are velvety grey-green, with no mention of wax dust (but
development of wax may depend on age and microclimate); BL is 0.9-1.7 mm. Alatae have secondary rhinaria distributed ANT III
2-8, IV 0-2, V 0-1. Colonies on twigs are inconspicuous,
resembling small leaves or leaf-scales of the host plant. They may be attended by ants (Monomorium). On Tamarix
spp. in south, central and eastern Europe, North Africa, south-west and
central Asia east to Pakistan.
Monoecious holocyclic.
Ivanoskaya (1956) described the fundatrix on T. meyersi in Turkmenistan (as tamaricophila), and specimens from Pakistan on T. kotschyi in the BMNH collection
also seem to be fundatrices. Oviparae
are recorded from Portugal (Ilharco 1996a) and oviparae and apterous males
have been found in France (G. Remaudičre, cited in Nieto Nafría et al. 2005a). 2n=8. [There
is much confusion in the literature on the nomenclature of the Tamarix-feeding species of Brachyunguis. B.
tamaricophila of Ivanoskaya (1956; nec. Nevsky 1928b) seems to be tamaricis, and B. tamaricivorum Narzikulov) (= tamariciarum Ivanoskaya) is not clearly distinct. B.
tamaricifoliae (Hall 1926) is also a synonym of tamaricis. Records of tamaricis from Calotropis procera probably refer to either B. harmalae or B.
calotropicus. See also Nieto Nafría et
al. 2005a. ] Brachyunguis
tamaricophilus (Nevsky) (fig. 113A) Apterae according to the original
description are green, densely covered with mealy wax secretion; BL 0.7-1.7
mm. In large, ant-attended colonies on
young shoots, leaves and flowers of Tamarix
spp. Monoecious holocyclic in
Uzbekistan (Tashkent), oviparae being found, but males are still undescribed
(Nevsky 1928b). Ivanoskaya (1956)
described the fundatrix on T.
karakalensis in Turkmenistan (as tamaricifoliae). It is also recorded from Kazakhstan, Iran,
Israel, Libya and Pakistan (BMNH collection).
B. tamaricifoliae of
Ivanoskaya (1956; nec. Hall 1926) seems to be this species. [The account of this species from Sicily by Barbagallo
& Stroyan (1982) applies to B.
tamaricis.] Brachyunguis tausaghyz (Nevsky ex Ivanoskaya)
Apterae are greenish yellow, with pruinose covering, and pale
siphunculi; BL c.1.6 mm. On Scorzonera
tausaghyz and Seriphidium nitrosum in
Kazakhstan, colonising flower heads (Narzikulov et al. 1971). Kadyrbekov
(2017a) reported it also from Chondrilla
brevirostris. B. afghanica,
described from Artemisia turanica in
Afghanistan (Narzikulov &
Umarov 1972), does not seem to be distinguishable from this species on the
basis of the published descriptions. Brachyunguis
tetrapteralis (Cockerell) Apterae are pale greyish green to
blue-green, with antennae dusky/dark distally; BL 1.0-1.7 mm. On stems and
leaves of Atriplex spp in western
and south-western USA, and Mexico, and there is also a record from a Chenopodium sp. Usually ant-attended. Monoecious and holocyclic, with sexuales in
early October (aphidtrek.org).
Records from Suaeda (e.g.
Remaudičre & Halbert, 1996) should perhaps all be referred to B. cahuille, which has a dark, very swollen clypeus, shorter siphunculi and shorter antennal
terminal process. Records from Sarcobatus
are possibly all referable to B.
bonnevillensis, and records
from plants outside the Amaranthaceae also need to be treated warily. The synonymy of Aphis piutapa (Hottes & Wehrle 1951b, on Lycium parviflorum) with B.
tetrapteralis is possibly incorrect, as the description of A. piutapa differs in certain respects
such as the shape of the cauda. The palaearctic species B. harmalae is very similar, but has slightly longer dorsal body
hairs. Brachyunguis
tigrahaudus Kadyrbekov Apterae are greenish with brown head,
without wax powdering; BL 1.0-1.4 mm. On flowerheads of Helichrysum arenarium and Jurinea
cyanoides in Aral region of south-west Kazakhstan (Kadyrbekov 2014c). Brachyunguis
transaralensis Kadyrbekov Apterae greenish, without wax powder; BL
1.5-1.8 mm. Alatae have 5-9 secondary rhinaria on ANT III. On flowerheads of Chondrilla brevirostris, attended by
ants, in Aral region of south-east Kazakhstan (Kadyrbekov 2014c). Brachyunguis
zawadovskii (Nevsky) Apterae are green, covered with mealy
wax; BL 1.6-1.9 mm. In large colonies on upper sides of leaves of Lycium depressum (= turcomanicum) in Central Asia. Brachyunguis
zygophylli (Nevsky) Apterae are yellow, yellow-green or green
to dark green, in life covered with dense mealy wax; BL 1.6-2.0 mm. In large
colonies on stems, flowers and undersides of leaves of Zygophyllum fabago. In south-west (Turkey, Iran) and Central
Asia. The record from Spain
(Gómez-Menor 1951, as Brachyunguis
zygophilli) should be referred to B.
harmalae (Garcia Prieto et al.
2004, Nieto
Nafría et al. 2005a).
About 12 nominal taxa living without host alternation on Eriogonum in western USA. Related to Aphis but characterised by a
combination of short antennal terminal
process, short siphunculi, and usually a very short broad cauda. The dorsal
cuticle is strongly reticulated and often has variably developed dark sclerotisation.
Apterae often have secondary rhinaria on ANT III or III-V. K.S.Pike et al. (2009) reviewed the genus and keyed the species, studying
mitochondrial DNA sequence variation in six species and describing two new
ones. They also provided information on primary and secondary parasitoids.
Hille Ris Lambers (1966a) described subspecies of three of the species on the
basis of small morphological differences, but further work is needed to
assess the significance of the observed variation.
Braggia agathona
(Hottes) Apterae are pinkish brown, often with a
dark central spot on the abdomen, the whole body covered with white wax; BL
1.4-1.8 mm. On undersides of leaves of Eriogonum
corymbosum in Colorado and Utah, USA.
Monoecious holocyclic, with oviparae and alate males in October
(original description). Braggia
columbiana Pike Apterae are black or blackish brown with a
white or whitish grey waxy dorsal pattern (see http://www.flickr.com/photos/sandnine/4606202941/);
BL 1.4-2.6 mm. On Eriogonum spp., predominantly E.
compositum, in Oregon and Washington, USA (K.S.Pike et al. 2009). Apterous and alate viviparae occur from about
mid-May to mid-September, sexual morphs are undescribed. Braggia
deserticola Hille Ris Lambers Apterae are shining black dorsally, grey
and wax-powdered ventrally, with black appendages; BL 1.1-1.8 mm. Found in small numbers in flowerheads of Eriogonum fasciculatum, incl. ssp. polifolium, in California and Utah,
USA, and in Mexico. A sample from Eriogonum
sp. at a higher altitude in California with shorter R IV+V and paler
tibiae was distinguished as a subspecies, B.
deserticola thanatophila Hille
Ris Lambers. Braggia
echinata Gillette & Palmer (Fig.27e) Apterae are greyish olive-green, usually
with a slight rusty area at base of each siphunculus, and with white dorsal
reticulations and pale, thick hairs giving a frosted appearance; BL 1.3-1.9 mm.
On leaves and stems of Eriogonum
corymbosum and Eriogonum sp. in
western USA (Colorado, Utah). Oviparae and alate males in October (Palmer
1952). Braggia
eriogoni (Cowen) Plate 8d (Fig.27a,b,g) Apterae are black or brownish black with a
dorsal pattern of white reticulation, accentuated by a wax secretion; BL
1.1-2.1 mm. On leaves, stems and
flowerheads of Eriogonum spp. in western USA. Hille Ris Lambers (1966a)
distinguished several subspecies on the basis of differences in hair length,
siphuncular shape and extent of dorsal pigmentation, but further work is
needed to investigate whether any of the observed variation is correlated
with host plant, season or geographical distribution. Sexuales have not been
described. Braggia
longicauda Pike Apterae are black, blackish grey or
blackish brown, with white wax powdering; BL 1.5-2.9 mm. On Eriogonum spp., predominantly E. elatum, in western USA (Washington,
Oregon, California). Oviparae and alate males occur in September-October
(K.S.Pike et al. 2009). Braggia
uncompahgrensis Hottes (Fig.27f) Apterae are pale grey-green, with frosty
appearance due to thick pale dorsal hairs; BL 0.9-1.4 mm. On Eriogonum corymbosum in Colorado and
Utah, USA. Oviparae and alate males occur in October (Palmer 1952).
Doubtfully distinct from B. echinata. Braggia
urovaneta (Hottes) (Fig.27c,d) Apterae are black, appearing grey due to
powdering with wax; BL 0.9-1.9 mm. On flower stems and upper parts of Eriogonum sp(p). in western USA.
Oviparae and alate males were collected in Colorado on E. corymbosum in early October (Hottes 1950). Hille Ris Lambers (1966a) distinguished a
population with a shorter R IV+V on E.
latifolium in California as a subspecies, B. urovaneta ssp. pachysiphon.
Eight palaearctic and two
nearctic species all except one of which live without host alternation on
Brassicaceae. The exception is a Salix
feeder and should possibly be given its own genus. Brevicoryne are related to Lipaphis
and characterised by very low rounded antennal tubercles, short
siphunculi and broad triangular cauda. Eastop in Hodjat (1981) gave a key to
alate viviparae of Middle Eastern species. Brevicoryne
arctica Richards Apterae are very dark blue or green, with
wax; BL 1.7-2.mm. Alatae have secondary rhinaria distributed III 12-13, IV 5.
On Lesquerella arctica in northern
Canada (Ellesmere Island). Monoecious holocyclic with oviparae and apterous
males in late July (original description). Brevicoryne
barbareae Nevsky Apterae are dark green, dark brown to
black, with appendages dark except for pale ANT III and tibiae (black at apices); BL 1.5-2.0 mm.
Alatae have secondary rhinaria distributed III 17-20, IV 3-5. On undersides
of leaves of Barbarea vulgaris in
Central Asia. This name has been applied to a species on Barbarea and Nasturtium
in north-west India that differs from Nevsky’s species in several respects,
having apterae with extensive dorsal sclerotisation, dark tibiae, a much
longer cauda, and alatae (brachypterae) with rhinaria distributed III 35-36,
IV 0-1 (David & Hameed 1975). Alate males have been described of this
Indian species (L.K. Ghosh et al.
1980, as B. barbareae). Brevicoryne
brassicae (L.) Cabbage
Aphid or Mealy Cabbage Aphid Plate 12f Apterae are greyish green or dull
mid-green with dark head and dark dorsal thoracic and abdominal markings,
densely coated in greyish white mealy wax (see influentialpoints.com/Gallery);
BL 1.6-2.6 mm. Alatae have a dark head and thorax and dark cross-bands on
dorsal abdomen. It occurs on many genera and species of Brassicaceae, and is
a major pest of field crops (Blackman & Eastop 2000) in all temperate and
warm temperate parts of the world. Feeding is stimulated by the presence of
mustard oil glycosides, so other plants with these compounds, e.g. Reseda, are also colonised. Monoecious holocyclic with alate males in
colder regions, anholocyclic wherever winters are mild. Mustaţă
et
al. (2000) studied the large
complex of natural enemies of B.
brassicae in Romania. Ruiz-Montoya et
al. (2003) used enzyme electrophoresis to look for host-related genetic
structure in Mexican populations, and Ruiz-Montoya et al. (2005) and Ruiz-Montoya
&
Núńez-Farfán (2013) studied phenotypic differences between populations on
two sympatric Brassica species. The finding by Rebijith et al. (2013) of CO-I barcode differences indicative of possible
cryptic species within Indian populations of B. brassicae on Raphanus
sativus requires further
investigation. 2n=16. Brevicoryne
crambe Bozhko (= Brevicoryne
crambinistataricae Bozhko)
Apterae are pale green, wax-dusted, BL 1.9-2.8 mm. On terminal shoots
and stems of Crambe tatarica, and also recorded from species in other crucifer
genera (e.g. Barbarea, Isatis, Diplotaxis, Sisymbrium). In Hungary, Moldova, Armenia, Bulgaria,
Turkey, Ukraine and Iran. Brevicoryne
fraterna (Strom) Apterae are pale green to yellow-green;
BL 1.2-1.8 mm. On leaves and young twigs of unidentified species of Salix in USA (Illinois, Ohio, Utah,
Colorado). Monoecious holocyclic with oviparae and apterous males in October
(Palmer 1952). Brevicoryne
jiayuguanensis (Zhang,
Chen, Zhong & Li) Apterae are
silvery whitish green, powdered with wax; BL 1.5-1.8 mm. On Armoracia rusticana in Gansu province,
China (Zhang 1999, as Brachycolus).
The broadly triangular cauda and many (15-19) thick hairs on abdominal
tergite 8 as well as the host association indicate that the species belongs
in this genus. Brevicoryne
lonicerina Mukhamediev
& Akhmedov Apterae are pale
green with brown appendages; BL 2.3-2.6 mm. On Lonicera spp. in Central Asia. Brevicoryne
nigrisiphunculata Hodjat Apterae are yellow to dusky green, with
short black siphunculi; BL 2.1-2.7 mm.
On undersides of older leaves of
Crambe and Brassica in Turkey, Iran, Armenia
(Szelegiewicz 1979, as B. crambinistataricae)
and Ukraine. Brevicoryne
shaposhnikovi Narzikulov Apterae are pale green dusted with grey
wax; BL c.1.6-1.7 mm. On Lonicera spp.
in Central Asia (Mukhamediev & Akhmedov 1982).
A genus
with two species resembling Brachyunguis
but with protruberant eyes lacking ocular tubercles. [Placed in Macrosiphini
by Remaudičre & Remaudičre (1997), perhaps because of the absence of
marginal tubercles, but the spiracles on abdominal segments 1 and 2 are
widely spaced, which is a more reliable indicator that it should be placed in
Aphidini.] Brevicorynella
quadrimaculata Nevsky Apterae brown, dark brown or dark green
covered with a fine pruinose secretion, antennae pale yellow with articulations and all of last segment
black, abdomen with four blackish-brown dorsal spots; BL 1.1-1.6 mm. The aphids cluster on the ends of shoots of
Tamarix spp., and are attended by
ants (Crematogaster). In central Asia (Kazakhstan, Tajikistan,
Uzbekistan) (Nevsky 1928b), Iran (Barkhordari et al. 1981), and also recorded from Tamarix chinensis (= T.
ramosissima) in China
(Xinjiang; G. Qiao, pers. comm.). Brevicorynella
sexmaculata Qiao & Zhang Apterae are green covered with a fine
pruinose secretion, with three pairs of dark patches on dorsal abdomen; BL
1.3-1.6 mm. Alatae are undescribed. On Tamarix
chinensis (= T. ramosissima) in Xinjiang province, China (Qiao et al. 2004b).
A
genus for one Artemisia-feeding
species in north-east China, related to Microsiphum
and Macrosiphoniella but with an
unusual form of cauda and long fine hairs on body and appendages. Alatae are
unknown. Brevisiphonaphis
hirsutissima Stekolshchikov & Qiao Colour of apterae in life is unrecorded,
BL 1.6-1.8 mm. On Artemisia argyi
in Liaoning prov., China (Stekolshchikov & Qiao 2008a, b). Life cycle is
unknown.
One African species
superficially resembling Paczoskia,
but with R IV+V having numerous short stiff hairs and very long siphunculi
without subapical reticulation. Alatae are unknown. Burundiaphis
autriquei Remaudičre Apterae are shining greenish black; BL
2.2-2.9 mm. On Davallia
chaerophylloides ( = denticulata)
epiphytic on oil-palms, often in mixed colonies with Micromyzella. Only
known from Burundi (Remaudičre & Autrique 1985).
Two
species described from galls on the primary host (Zelkova), one of which is now known to migrate to Carex.
Alate spring migrants, emerging from pseudogalls on Zelkova, have an unbranched media,
only one oblique vein in the hind wing, and inconspicuous ring-like
siphunculi. Resembling Kaltenbachiella in many characters,
but in Byrsocryptoides the
secondary rhinaria are less annular and the processus terminalis is
relatively longer, and a detailed cladistic study by Sano & Akimoto
(2011) placed this genus in a basal position in the tribe Eriosomatini. Dzhibladze (1965) provided keys to
fundatrices, apterous fundatrigeniae and spring migrants of the species on Zelkova. Byrsocryptoides
zelkovae Dzhibladze
The galls (pseudogalls) on Z.
carpinifolia in Georgia are formed by rolling, distortion and blistering
of a terminal or subterminal leaf, starting at the base. Alatae (fig.
120D)
leaving the galls in June are small (BL about 1.2-1.3 mm), rather
long-bodied, with dark brown head, meso- and metathorax and pale yellow to
green prothorax and abdomen (Dzhibladze 1960b). They migrate to aerial parts of Cyperaceae;
apterae of elongate form with a long last rostral segment collected on Carex riparia in Germany (BMNH collection, leg. F.P. Müller) are
provisionally assigned to this species, and Barjadze & Gratiashvili
(2009) obtained experimental colonisation of Carex capitata in Georgia. Byrsocryptoides
zelkovaecola Dzhibladze Galls (fig. 125A)
are similar to those of B. zelkovae,
and occur on Zelkova carpinifolia
in Georgia. Alatae (fig. 120C) of
BL 1.1-1.4 mm emerge in June-July. The
life cycle is unknown (Dzhibladze 1965b).
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