SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order) A
Four
North American species previously placed in Ovatus but perhaps more closely related to Myzus (Jensen & Stoetzel 1999). Characteristic features are
the siphunculi with slight subapical swelling and small flange, and the
reticulate dorsal cuticle. Abstrusomyzus
leucocrini (Gillette
& Palmer) Apterae are pale green
to brownish green or black; BL 1.6-1.8 mm. Described as abundant on leaves of
Leucocrinum montanum in Colorado,
USA (Gillette & Palmer 1934), but there are subsequent records from Achillea (including oviparae and
apterous males collected in October) and Potentilla
(A. Jensen, aphidtrek.org)
so its true host associations are still uncertain. Abstrusomyzus
phloxae (Sampson) Plate 17h Apterae are pale apple-green (see aphidtrek.org); BL 1.2-1.8 mm.
Recorded from many plant species and genera in different families (Achillea, Agoseris, Apocynum, Capsella, Carex, Centaurea, Cerastium, Galium, Phacelia, Phlox, Plantago, Polygonum, Ranunculus, Stellaria, Trifolium, Viola). Usually it colonizes the
basal or rosette leaves of low-growing plants. Widely distributed in North
America; in eastern USA it is most common on Plantago, where colonies are often attended and sheltered by ants
(Jensen & Stoetzel 1999). In western USA it is common on Apocynum where it feeds on undersides
of leaves causing discoloration visible from above (A. Jensen, aphidtrek.org) Populations from different parts of North
America in the BMNH collection show differences in ratio of R IV+V to HT II,
but it is unclear whether these are related to host plant or geographical
location. Apparently mostly
anholocyclic, although males have been produced in culture (BMNH, leg. C.-k.
Chan), and oviparae were collected
from Fragaria vesca in Nova Scotia
(BMNH, leg. H.T. Shultz). 2n=18*. Abstrusomyzus
reticulatus (Heie) Apterae are shining back; BL 1.6-1.9 mm.
On undersides of curled leaves of Oxalis sp. (probably stricta) in North Carolina, USA.
Oviparae and alate males occur in October. Abstrusomyzus
valuliae (Robinson) Apterae are blackish, BL c.1.2-1.3 mm. Alatae have 2-7 rhinaria
on ANT III. On Fragaria vesca in
Manitoba and Alberta (Canada).
A
genus of two palaearctic species, with spiracles on abdominal segments 2-5 as
in Phylloxera and Viteus, but characterized by the curious
star-like tubercles borne by immatures.
The life cycle is remarkable in having only two generations per year,
fundatrices and sexuales. The adult fundatrices bear small spiculose
tubercles on head, thorax and abdomen.
Acanthochermes quercus
Kollar Recorded from Quercus
robur and Q. petraea in European Russia, Finland, Sweden, central
Europe (Hungary, Germany), Portugal and the Middle East (Iran). The large fundatrices (BL up to 3 mm)
induce small ring-shaped galls on the upper surface of the leaf, opening on
the underside (see Chen & Qiao 2012a).
Morphology and developmental stages were illustrated by Grassi (1912),
and Tavares (1931) gave an account of the biology. Acanthochermes
similiquercus Jiang,
Huang & Qiao Fundatrices are
yellow dotted with black; BL 1.6-2.3
mm. In small yellow, yellowish green or red galls on leaves of an
unidentified Quercus sp. in Sichuan, China (Jiang et al. 2009).
One
palaearctic species on Atraphaxis
(Polygonaceae) with 5-segmented antennae, no evident cauda and
characteristically shaped siphunculi. A second species previously noted as
belonging to Acaudella has been
described in a new genus (Blackmania). Acaudella
puchovi Nevsky Apterae are dark green, BL c.1.4 mm.
Alatae have 9-19 secondary rhinaria on ANT III, 0-1 on IV. In large numbers
on undersides of leaves of Atraphaxis
buxifolia (= caucasica) in Uzbekistan (Samarkand). It is also
recorded from A. spinosa in both
Kazakhstan (Kadyrbekov 2017a) and Israel (Bodenheimer & Swirski 1957). A
redescription is provided by Kanturski & Wieczorek (2015b). The life
cycle is unknown.
Five
palaearctic species with a broadly rounded cauda living without host
alternation on Centaurea. Holman
(1991) provided a revision and a key
to species. They are adapted for ant attendance. Acaudinum
beheni Remaudičre & Davatchi Apterae are shining dark brown, BL 1.5-1.75 mm. On basal leaves of Centaurea behen in western Asia (Iran,
Turkey, Armenia, Azerbaijan), attended by ants. Oviparae and apterous
(alatiform) males occur in late September (Tuatay & Remaudičre 1964). Acaudinum
bulgaricum Holman Apterae are dull dark brown to black ,
BL 1.4-1.9 mm. On Centaurea
solstitialis in Bulgaria, feeding
on upper and undersides of leaves and also on stem and branches, attended by
ants. An alate male was collected in early September (Holman 1991). Acaudinum
centaureae (Koch) Plate 10h Apterae are shining black (see Influential
Points/Gallery), BL 1.7-2.5 mm. At bases of leaf stalks of Centaurea spp., often in ant shelters
(Stroyan 1950; Holman 1991). Widely distributed in Europe. Oviparae occur in
September-October. 2n=10. Acaudinum
longisetosum Holman Apterae are matt or slightly shiny jet
black, BL 1.9-3.2 mm. At stem base and root collar of Centaurea
spp., especially C. scabiosa,
in spring, later also living on terminal parts of plants, attended by ants.
There are also records from Achillea spp.
(Wojciechowski et al. 2016) that
perhaps require additional confirmation. In
Poland, Czech Republic, Slovakia, Switzerland, Italy and Ukraine.
Oviparae and apterous males occur in October. Acaudinum
roumanicum Holman Apterae are matt or slightly shiny jet
black, BL 1.6-2.7 mm. At stem bases of Centaurea
rhenana in spring, later on terminal parts of plants, attended by ants.
Closely related to A. longisetosum;
the discriminant given in the key is based on work of Holman (1991). Romania, Bulgaria and Slovakia.
Sexual morphs are unknown.
Two
nearctic and one palaearctic species on Compositae, possibly related to Brachycaudus but with secondary
rhinaria in apterae, capitate dorsal hairs and a differently shaped cauda
(Hille Ris Lambers 1956c). Leclant
& Remaudičre (1967) compared and keyed the species. Acuticauda
asterensis Gillette
& Palmer (Fig.15g,h) Apterae are pale yellowish green; BL c.1.3
mm. Alatae have black wing veins. On leaves and stems of Aster and Symphyotrichum
spp. in USA (Colorado, New York and Wisconsin) and Canada (Quebec);
there is one record from Solidago requiring
substantiation. Oviparae and alate males in autumn (Palmer 1952). Acuticauda
erigerontis Leclant
& Remaudičre Apterae are deep
golden yellow with brown antennae and siphunculi; BL 1.2-1.5 mm. Alatae have
dark wing veins. On Erigeron acer in
the French Alps, rolling the leaves of the basal rosettes (Leclant &
Remaudičre 1967). Monoecious holocyclic, with apterous males. Acuticauda
solidaginifoliae (Williams) Plate 10c Apterae are shining reddish or greenish
brown with a black dorsal shield; BL 1.8-2.0 mm. On leaves of Solidago spp., which are folded along
mid-ribs to become pod-like. Widely distributed in northern and western USA.
Monoecious holocyclic, with apterous males in September-October (Palmer
1952).
One
species near Vesiculaphis but with
distinctive siphunculi. Hille Ris Lambers (1973) discussed its taxonomic
position and probable host relationship.. Acutosiphon
obliquoris R.C. Basu,
A.K.Ghosh & Raychaudhuri (Fig.18a) Apterae are dark or reddish brown; BL
1.6-2.1 mm. Secondary rhinaria in alata III 25-28, IV 12-14, V 1-3.
Originally described from Cotula (Compositae),
but this is an unlikely host, and subsequent collections indicate possible
host alternation between Ericaceae (Lyonia
ovalifolia) and Cyperaceae (Carex,
Cyperus); see Hille Ris Lambers (1973b) and Miyazaki (1977). In India
(Assam) and Nepal, and more
recently fully redescribed and illustrated from China (Fang et al. 2006).
A genus of
about 70 palaearctic and 9 nearctic species living without host
alternation on various dicots, particularly Leguminosae/Fabaceae, Rosaceae,
and Euphorbiaceae. They are typically rather
large broadly spindle-shaped, short-haired aphids with long antennae, legs,
siphunculi and cauda, usually green but sometimes brownish, pink, or
yellow. Eastop (1971) gave keys for
the identification of the species then known, and there are accounts for
Fennoscandia and Denmark (Heie 1994), Britain and Ireland (Blackman 2010),
Japan (Miyazaki 1971), India (Raychaudhuri, M.R.Ghosh & R.C. Basu 1978)
and the Russian far east (Pashchenko 2005). Meier (1958) reviewed, and
provided photographs of, the European legume feeders. Nieto Nafría et
al. (2015b) provided a list of species. Acyrthosiphon argus Miyazaki Apterae are pale green, with a dark green spinal stripe; BL 1.8-2.9 mm. Described from Daphne odora in Japan, colonising the undersides of leaves and the flowers. Infested leaves turn yellowish or dark brown, and heavy infestation can cause early defoliation and death of shrubs (original description). Previously collected on Daphne sp. in China (BNMH collection, coll. VFE, 31.v.1985). Populations in Japan (perhaps introduced from the Asian mainland) were apparently anholocyclic. Acyrthosiphon assiniboinensis Robinson Apterae are green; BL 2.2-2.8 mm. Alatae have 15-18 secondary rhinaria on ANT III. On Potentilla fruticosa in Canada (Manitoba, Northwest Territories, Yukon). Acyrthosiphon astragali Eastop Apterae are green, BL 2.2-2.8 mm. In large colonies on stems of Astragalus sp(p)., falling readily when disturbed (Narzikulov 1972). Recorded from Afghanistan, Pakistan, Tajikistan and Kashmir (BMNH collection). Alatae are undescribed. Acyrthosiphon auctum (Walker) Apterae are pale yellow-green, green or pale pink, slightly wax bordered intersegmentally; BL 1.9-2.8 mm. Alatae have a dark head and thorax and reddish abdomen. On various Caryophyllaceae (Cerastium, Honkenya, Silene, Stellaria) and Brassicaceae (Cakile, Capsella), mainly in sand dune habitats, in north Europe, including Iceland (Heie 1994), and Greenland (BMNH collection, leg. J.M. Cotton). Monoecious holocyclic with oviparae and apterous males in September-October (in Scotland; BMNH collection, leg. H.L.G. Stroyan). Acyrthosiphon auriculae Martin Apterae are pale to bright green; BL 1.8-2.85 mm. On upper surfaces of young apical leaves of a cultivated hybrid Primula (section Auricula); experimental host transfers indicated specificity to this section of the genus (Martin 1981). Only known from England, although alatae of a possible geographic variant of this species were trapped in Corsica. Monoecious holocyclic with apterous males. 2n=8. Acyrthosiphon bidenticola Smith Apterae are bright green with mainly dark antennae, legs and siphunculi; BL 1.6-2.0 mm. On various Compositae (Aster, Bidens, Erechtities, Gnaphalium, Helianthus, Lagascea, Tithonia, Wedelia), and populations have also been found on Bocconia frutescens (Mexico, BMNH collection, leg. R. van den Bosch) and Salvia coccinea (Jamaica, Tannice Hall, pers. comm.). Central and South America (Brazil, Cuba, Jamaica, Mexico, Puerto Rico, Venezuela). 2n=8* Acyrthosiphon bistorti Ivanoskaya Apterae are yellowish green; BL c. 4.1 mm. Alatae have about 8 secondary rhinaria on ANT III. In inflorescences of Persicaria bistorta in west Siberia, and also now recorded from Polygonum and Persicaria spp. in Kazakhstan (Kadyrbekov 2012a) and Russia (Altai; Kadyrbekov 2017a). Acyrthosiphon boreale Hille Ris Lambers Apterae are green, yellowish green or yellowish (see influentialpoints.com/Gallery); BL 2.7-3.2 mm. Apterae have 2-9 secondary rhinaria on ANT III. On Potentilla spp. in northern Europe, Greenland, Canada (Baffin Island) and north-east Siberia (Chukotka). There are also records from Switzerland, Hungary and Kazakhstan, although the distinction from other members of the malvae group is not clear enough to be certain about the extent of its distribution. It seems likely that A. malvae ssp. pentaphylloides, described from Potentilla fruticosa in east Siberia (Pashchenko 2005) and also recorded from Kazakhstan (Kadyrbekov 2016b), is a synonym. Monoecious holocyclic, with apterous males. Stekolshchikov & Khruleva (2015) provided detailed descriptions of the male and ovipara. 2n=10. Acyrthosiphon brachysiphon Hille Ris Lambers Apterae are waxy green in life, 1.8-2.8 mm,
with siphunculi hardly longer than cauda, their apices slanted outwards. This aphid has a boreoalpine distribution
on Vaccinium spp. (Blackman &
Eastop 2006), but there is a record of apterae and oviparae from Salix
lapponum in Norway (Heikinheimo 1966, as A. aurlandicum). Acyrthosiphon brevicorne Hille Ris Lambers Colour of apterae unrecorded, probably green, like oviparae; BL of aptera 1.8-2.3 mm. On Dryas spp. Distribution is boreoalpine; northern Canada, Greenland, Sweden and Switzerland. Monoecious holocyclic, with oviparae and apterous males in July-August in the Arctic, and in October in Switzerland (Meier 1972). Strathdee et al. (1995) studied cold-hardiness of eggs under laboratory conditions. A. dryasae described from Sakhalin is possibly this species. Acyrthosiphon capitellum Zhang Apterae are green with dark apices to antennal segments, femora and tibiae, and dark basal parts of siphunculi, BL c. 3.3 mm. On Fragaria vesca in China (G. Zhang & Qiao 1998c, 1999d). Acyrthosiphon caraganae (Cholodkosvsky) Apterae are green with variably developed greyish wax stripes between segments, and often a bright green spinal stripe; BL 2.6-4.3 mm. On leaves and young growth of woody Leguminosae, especially species of Caragana and Colutea, often forming large colonies. Probably originating in the Altai region, it now occurs in parks and gardens in temperate regions throughout the Northern Hemisphere. Monoecious holocyclic; males are apterous in western Europe and alate in Russia, Switzerland and eastern Canada (Eastop 1971). Populations in western Europe, Tajikistan and Sakhalin have been distinguished as subspecies; A. caraganae occidentale Hille Ris Lambers, A. caraganae tadzhikistanicum Narzikulov and A. caraganae sachalinense Pashchenko (2005). 2n=10 (also for ssp. occidentale). Acyrthosiphon (Liporrhinus) chelidonii (Kaltenbach) Apterae are pale green or greenish white, covered with greyish-white wax powder except along segmental borders; BL 1.5-2.3 mm. On Chelidonium majus, forming colonies on upper parts of stems and on young leaves. In Europe, Kazakhstan (Kadyrbekov 2017a), east Siberia, Korea and Japan. Oviparae and alate males in Netherlands in October-November (Hille Ris Lambers 1947). Acyrthosiphon churchillense Robinson Apterae are green; BL 2.1-2.6 mm. On Oxytropis spp. and some other
Leguminosae (Melilotus alba, and
probably Hedysarum boreale ssp. mackenzii) in northern Canada
(northern Manitoba, Yukon, Northwest Territories). Monoecious holocyclic with
oviparae and apterous males in July-August (Robinson 1979 and BMNH
collection, leg. AG Robinson). Close to A.
kondoi; see couplet for separating these species on Oxytropis. Acyrthosiphon corsicae Remaudičre & Leclant Apterae are pale green; BL 2.2-2.6 mm. Alatae have 8-10 rhinaria on ANT III. On Euphorbia spinosa in Corsica (Remaudičre & Leclant 2000). Acyrthosiphon crepidis Holman & Szelegiewicz Apterae are green, with ventral side pruinose; BL 1.7-2.2 mm. Other morphs are unknown. On Crepis (= Youngia) tenuifolia in Mongolia. Acyrthosiphon cyparissiae (Koch) Apterae are green with black front part of head and mainly black appendages or, especially in warmer climates, yellowish green with appendages mainly brown or brownish yellow; BL 2.4-3.4 mm. On Euphorbia spp., feeding mainly on upper sides of upper leaves, often in large numbers. In Europe (but UK records apply to other species), Middle East, Central Asia and China . Monoecious holocyclic with alate males (Hille Ris Lambers 1947). In warmer regions the less-pigmented form A. cyparissiae ssp. propinquum Mordvilko is more common (Eastop 1971), although it needs to be confirmed that this is not merely an effect of the environment. Another nominal subspecies, A. cyparissiae ssp. turkestanicum Nevsky, is reported from Central Asia. Acyrthosiphon daphnidis Ilharco Apterae are green to yellowish green with red eyes, body with greyish white powder accentuating dorsal segmentation; BL 2.4-2.8 mm. In colonies on undersides of leaves of Daphne gnidium; severe infestations can cause yellowing and dropping of leaves. In Portugal, Spain, Canary Islands (Ilharco 1996b), Corsica and northern Italy (Barbagallo et al. 2011), and also reported from Iran (Rezwani & Parvizi 1990). Acyrthosiphon dauricum Szelegiewicz Apterae are green; BL 2.3-2.6 mm. Described from fundatrices and apterae on a composite plant in Mongolia, that was subsequently identified as Saussurea salicifolia (Szelegiewicz 1964a). However, it could not be verified that this was the true host, and subsequent records from Delphinium grandiflorum and Linaria buriatica (Holman & Szelegiewicz 1972) have not helped to clarify the situation. Acyrthosiphon dryasae Pashchenko Apterae are pale green: BL c.2.1 mm (Pashchenko 2005). On Dryas punctata in east Siberia (Kamchatka). Apart from the slightly longer siphunculi, the description does not clearly distinguish this species from A. brevicorne. Acyrthosiphon echinospartii Nieto Nafría & Mier Durante Apterae are pale green, with somewhat darker appendages; BL 2.6-3.1 mm. On Echinospartium lusitanicum in Spain, where it has also subsequently been recorded from Genista sp. (Mier Durante et al. 1995). Acyrthosiphon elaeocarpi
Tao Apterae are whitish yellow with
yellowish brown siphunculi and blackish brown antennae, apices of tibiae,
tarsi, cauda and anal plate; BL c. 3.5 mm.
On Elaeocarpus serratus in Taiwan (Tao 1963). Biology and sexual morphs are unknown. Acyrthosiphon ericetorum Hille Ris Lambers Colour of apterae in life unknown, probably pale green; BL c. 1.9-2.3 mm. On Genista spp. in the Netherlands. Monoecious holocyclic; the original description was of oviparae and apterous males. Apterous viviparae were described by Nieto Nafrěa & Mier Durante (1985). This aphid only seems to differ from A. genistae by its small size, which could be an effect of the host plant. Acyrthosiphon euphorbiae Börner Apterae are green, tinged with yellow; BL 2.3-4.4 mm. On Euphorbia spp., in small colonies on upper sides of leaves, sometimes mixed with E. cyparissiae. In Europe, North Africa, the Middle East and Central Asia. Alatae have 11-25 rhinaria on ANT III. Sexual morphs are undescribed and the life cycle is apparently unknown. Narzikulov (1953) described a population in Tajikistan as a subspecies, A. euphorbiae ssp. tadzhikistanicum. Another subspecies described from western Europe, A. euphorbiae ssp. neerlandicum Hille Ris Lambers, was established to be a synonym by Remaudičre & Leclant (2000). The range of measurements recorded for populations in Kazakhstan described as another subspecies, A. euphorbiae ssp. brevisiphon Kadyrbekov, is within that recorded for A. euphorbiae from other parts of the world (Kadyrbekov 2005e, as A. neerlandicum ssp. brevisiphon). A closely related form, A. pareuphorbiae, occurs in China. Acyrthosiphon evodiae
(Takahashi) Apterae are yellow in
life, about 2.3 mm, with black antennae and legs, brown-black siphunculi and
a pale cauda. Alatae have wings "somewhat
clouded along the veins".
Described (in Macrosiphum) from specimens found feeding on
young leaves of Euodia (= Melicope)
triphylla in Taiwan (Takahashi 1929). Aphids collected on Acronychia pedunculata in Hong Kong
were provisionally identified as this species (Martin & Lau 2011, as Sinomegoura evodiae). The generic
placement of this species is uncertain; it may belong in Sinomegoura.The life cycle is unknown. Acyrthosiphon extremiorientale Pashchenko Apterae are green, emerald green or orange, with antennae and siphunculi mainly dark; BL 4.2-4.9 mm. The single known alata had 40 rhinaria on ANT III. Feeding on Vicia spp. and Astragalus schelichowii in east Siberia (Pashchenko 2005). The life cycle is unknown. Acyrthosiphon fragariaevescae Nevsky Apterae are brownish or green, with pale antennae, siphunculi and cauda; BL c.2 mm. Alatae have 13-22 rhinaria on ANT III, and ANT PT/BASE c. 4.1. On Fragaria vesca in Central Asia. Kadyrbekov (2005e) provided a redescription of the aptera. Apart from the shorter ANT PT this species seems close to A. rubi. Acyrthosiphon fragum Zhang Apterae are green with black siphunculi; BL c. 3.9 mm. On Fragaria sp. in China (G. Zhang & Qiao 1998c, 1999d). Acyrthosiphon galijae Kadyrbekov Apterae are green; BL 3.3-4.0 mm. On stems and leaves of Sanguisorba officinalis in Kazakhstan (Kadyrbekov 2005e). Very similar to A. sanguisorbae;, and the differences (size, hair length, etc.) could be due to host plant and other environmental factors. Acyrthosiphon genistae Mordvilko Apterae are green to yellow green, with dark brown siphunculi; BL c. 3.4 mm. On Genista tinctoria, at ends of stems and branches. In Russia and Ukraine (Bozhko 1976). A. ericetorum from other Genista spp. in Austria could be small specimens of this aphid. Acyrthosiphon ghanii Eastop Colour of apterae in life unknown; BL 2.0-2.4 mm. On Compositae/Asteraceae (Sonchus, Scorzonera) in Pakistan (Naumann-Etienne & Remaudičre 1995). 2n=10? (Kapoor & Gautam 1994, but aphid was possibly misidentified as host was Medicago). Acyrthosiphon (Xanthomyzus) glaucii (Narzikulov) Apterae are shining greenish white to white; BL 1.8-2.2 mm. On Glaucium spp. in Central Asia. It is also recorded from Hypecoum parviflorum (Kadyrbekov 2017a) [Host plant of type material is given as G. fimbrilligerum in original description, but as G. corniculatum on slide labels of specimens in BMNH collection with same date and locality, leg. Narzikulov.] Acyrthosiphon
gossypicola Shinji Perhaps a synonym of Acyrthosiphon gossypii, but apterae were described as reddish
brown in life, and alatae have only 7-9 secondary rhinaria on basal half of
ANT III, whereas those of A. gossypii
have 10-21 distributed over most of its length (Eastop 1971). On Gossypium herbaceum in Korea. Acyrthosiphon gossypii Mordvilko Plate 23c Green in life, with adults as well as immatures dusted with fine wax (see aphids of Karnataka website); BL of apterae 2.5-3.8 mm. Host plants are mainly Leguminosae/Fabaceae especially tribe Phaseoleae (Dolochos, Phaseolus, Vigna, but not known from Glycine), Malvaceae (including cotton) and Zygophyllaceae; more rarely on other plants including Brassicaceae. Trees recorded from include Hibiscus esculentus, Robinia pseudoacacia and Sophora japonica. Intraspecific variation in host plant preferences and life cycle are discussed by Müller & El Tigani (1986). Populations with particular host preferences may occur in different regions (Blackman & Eastop 2000). In southern Europe, North Africa, Middle East, south-west and central Asia, India, China, and possibly Korea and Japan. Anholocyclic over much of its range, but there is a holocycle on Alhagi camelorum in central Asia. A. umarovi (q.v.) is probably a synonym. 2n=6. Acyrthosiphon heptapotamicum Kadurbekov Apterae are green; BL 3.2-3.9 mm. On stems and leaves of Clematis songoricum in Kazakhstan (Kadyrbekov 2005e). This aphid has long thin siphunculi like those of A. gossypii, but with a longer ANT PT than that species, and more caudal hairs. Acyrthosiphon
hissaricum Umarov Apterae are green; BL c.3.9 mm. Alatae
have BL c. 3.1 mm and 20-30 secondary rhinaria on ANT III. On Cicer songaricum in Tajikistan, and
also reported to occur on Medicago falcata and Hedysarum spp. in Kazakhstan (Kadyrbekov 2012a, 2014d). Similar
to A. rubi, but much larger.. Acyrthosiphon ignotum Mordvilko Apterae are pale green or yellowish green; BL 2.2-3.4 mm. Apterae have 1-8 secondary rhinaria on ANT III, and alatae have 7-13. At shoot tips of Spiraea spp., often in large colonies. Also recorded from Sibiraea laevigata. In northern Europe, Iran, Kazakhstan, north India, Siberia, Mongolia (many records are as A. spiraeae). A. leleji may be a synonym. 2n=10 (Kuznetsova & Shaposhnikov 1973), but Khuda-Bukhsh & Pal (1986b) record 2n=14 for an aphid identified as this species collected on Deutzia corymbosa. Acyrthosiphon ilka Mordvilko (= Acyrthosiphon bidentis Eastop) Apterae are green, often with darker green dorsal cross-bands; BL 1.4-2.7 mm. On Papaveraceae (Glaucium, Papaver), Compositae/Asteraceae (Bidens, Carthamus, Cichorium, Lactuca, Lampsana, Picris, Sonchus), Brassicaceae (Erysimum, Isatis), Linaceae (Linum) and plants in several other families; e.g. Kadyrbekov (2014d) records it (as A. bidentis ssp. montanum) from Kazakhstan on three genera of Campanulaceae (Adenophora, Campanula, Codonopsis). In southern Europe, Middle East, Central Asia, Pakistan (Naumann-Etienne & Remaudičre 1995), south Siberia, China (as A. linisuctum) and Africa. The African populations are found on a very similar range of hosts to the Eurasian ones, but tend to have longer, more slender siphunculi, and were described as a distinct species, A. bidentis Eastop, but this seems only to be a geographical variant of A. ilka (Eastop & Blackman 2005), as also may be A. mordvilkoi Nevsky, described from Linum usitatissimum in Central Asia, and A. papaverisuctum (q.v.). A form described as a subspecies, A. ilka ssp. montanum Kadyrbekov occurs in mountainous regions of Kazakhstan above 1500m on a variety of host plants (Kadyrbekov 2005e, as A. bidentis ssp. montanum); the observed differences might be environmentally determined. Other possible members of this group are A. papaverinum Nevsky ex Pek and A. titovi Mordvilko, both recorded from Papaver sominiferum in Central Asia but neither formally described. Acyrthosiphon
kapustjanae Pashchenko Apterae are green in life; BL 2.7-2.8 mm
(2 specimens). Described from Salix
udensis in Kamchatka, east Siberia (Pashchenko 2005). Further collections
are necessary to determine whether this is the normal host. This species is
very similar to A. dauricum. Acyrthosiphon knechteli (Börner) Apterae are green or red; BL 2.0-3.0 mm. On undersides of leaves and on young shoots of Vaccinium uliginosum in northern and central Europe and west Siberia. Monoecious holocyclic with oviparae and alate males at end of August (Heikinheimo 1990), and apparently specific to V. uliginosum. However, in north-west Russia, Stekolshchikov (2017) and Stekolshchikov & Khureva (2020) have reported populations of an aphid that differed insignificantly in its morphology from A. knechteli, but was found feeding on eight plant species in six families. Further research is needed to clarify the taxonomic status of this form (“Acyrthosiphon sp. aff. knechteli”). Acyrthosiphon kondoi Shinji Plate 23b Apterae are bluish green (see aphids of Karnataka website); BL 2.1-2.9 mm. Alatae have a darker brown thorax than A. pisum. On stems and leaves of Leguminosae/Fabaceae, mainly of tribes Trifoleae (Medicago, Melilotus, Trifolium) and Loteae (Dorycnium, Lotus) but also on Astragalus (Galegeae), and also recorded from Pisum, Vicia and Lens (Vicieae). An important pest of alfalfa (Blackman & Eastop 2000), now widely distributed in Asia, North and South America, South Africa, Australia and New Zealand, with distribution more recently extended to the Mediterranean region (Turkey, Kök & Özdemir 2021, Greece, Tsitsipis et al. 2007 and Corsica, Nieto-Nafría et al. 2007). A record from an unlikely host Fagonia in Algeria (Laamari et al. 2013) needs further confirmation. Monoecious holocyclic in Japan (Kawada 1992), apparently anholocyclic in most other places. 2n=10. Acyrthosiphon (Tlja) lactucae (Passerini) Apterae are pale yellowish green or pink, with pale grey wax bloom (see Influential Points/Gallery); BL 1.7-2.9 mm. On stems and undersides of leaves of Lactuca spp. Europe, Middle East, Kazakhstan (Kadyrbekov 2014d), Pakistan (Naumann-Etienne & Remaudičre 1995), introduced to North America and more recently recorded from Argentina (Mier Durante et al. 2011) and Chile (Nieto Nafría et al. 2018). Monoecious holocyclic, with alate males. 2n=16. A. scariolae (q.v.) is probably a distinct species with 2n=18, but further work is necessary to confirm this. Acyrthosiphon (Xanthomyzus) lambersi Leclant & Remaudičre Apterae are yellowish green, lightly wax-dusted on abdomen; BL 1.4-2.7 mm. On Glaucium spp. in southern Europe, North Africa and the Middle East. Anholocyclic in France according to original description. Acyrthosiphon leleji Pashchenko Apterae are green; BL 2.3-2.5 mm. On shoot apices and leaves of Spiraea salicifolia and Spiraea sp. in east Siberia (Pashchenko 2005). The alatae and life cycle are unknown. It is not clear how this species differs from A. ignotum. Acyrthosiphon leonurae Pashchenko Apterae are pale green; BL 2.6-2.7 mm. Occurring singly on undersides of leaves and on apical shoots of a species of Leonurus in east Siberia (Pashchenko 2005). The alatae and life cycle are unknown. Perhaps this species should be placed in Aulacorthum. Acyrthosiphon lobkovae Pashchenko Colour of apterae in life is unknown; BL 2.9-3.2 mm. On Myrica tomentosa (= M. gale) in east Siberia (Kamchatka). An ovipara was collected in September (Pashchenko 2005). An ovipara collected at the same time and from the same host differed in having relatively shorter ANT PT and longer siphunculi, and was described as another species, A. myricae Pashchenko. Acyrthosiphon loti (Theobald) Apterae are green, frequently with faint greyish transverse stripes (see influentialpoints.com/Gallery), or more rarely pink; BL 1.7-2.9 mm. On Lotus and Dorycnium spp., Anthyllis vulneraria, Hippocrepis comosa, Onobrychis sativa (= viciifolia), and sometimes on certain other herbaceous Leguminosae, including Medicago sativa. In Europe, eastward to west Siberia, Turkey and Iran, and introduced to South America (Argentina: Ortego & Mier Durante 2003; Chile: Nieto Nafría et al. 2018). Monoecious holocyclic, with both apterous and alate males. 2n=10. Acyrthosiphon macrosiphum
(Wilson) Yellow-green in life, with
darker antennae and tibiae (see aphidtrek.org);
1.5-1.9 mm. Holocyclic, with alate
males, on leaves of Amelanchier spp. in USA (California, Colorado,
Utah, Oregon, Montana) and Canada (British Columbia, Saskatchewan). Sambucus canadensis is also listed
as a host, but specimens have not been seen from that plant. References: Palmer 1952, Eastop 1971. 2n=10. A photo of a live fundatrix is available at http://www.flickr.com/photos/sandnine/4545416151. Acyrthosiphon malvae (Mosley) Plate 23a (Fig.30c) Apterae are green, yellowish or greyish green, or pinkish red (see influentialpoints.com/Gallery); BL 1.5-3.2 mm. Apterae have 1-24 secondary rhinaria on ANT III, alatae have 12-31. On many plants, but particularly herbaceous Rosaceae. This is a complex taxon, in which several forms with more specific host plant associations are recognised as subspecies, although slide-mounted specimens cannot usually be identified to subspecies level unless large samples are available. Populations of pale green or red aphids on many Geraniaceae (Geranium, Pelargonium, Erodium) and Malvaceae (Alcea, Malva), as well as many plants in other families, are regarded as A. malvae s. str., and are of almost world-wide distribution, with anholocycly in warmer climates and indoor populations. A. malvae ssp. agrimoniae (Börner) is yellowish green and found in flowerheads or on undersides of leaves of Agrimonia spp. in Europe and western and central Asia. A. malvae ssp. pentaphylloides is green and described from Pentaphylloides (=Potentilla) fruticosa in east Siberia (Pashchenko 2005; probably this is a synonym of A. boreale), A. malvae ssp. poterii Prior & Stroyan is bright salmon pink, yellowish or green and found on Poterium sanguisorba (= Sanguisorba minor), and is only known from England. A. malvae ssp. potha Börner is pale yellowish or greyish green and associated with Alchemilla spp., and possibly also Acaena spp. and Geum rivale, throughout Europe. Records of this subspecies from Potentilla are possibly all referable to the very similar A. boreale (q.v.). A. malvae ssp. rogersii (Theobald) is green or yellow-green, often shiny, and may form large colonies on young leaves of Fragaria in north and west Europe (Blackman & Eastop 2000). Populations in the BMNH collection from England and Switzerland with specificity to Geranium robertianum may also qualify for at least subspecies status.The recognised subspecies are apparently all monoecious holocyclic on their respective host plants; both apterous and alate males occur in this group, and occasional hybridisation probably occurs between host-specific and more polyphagous populations, further confusing the taxonomic situation (Prior & Stroyan 1964; Müller 1972). 2n=10 (incl. ssp. poterii, rogersii). Acyrthosiphon matilei Remaudičre & Leclant Apterae are green, BL 1.7-2.4 mm. Living all year round on Euphorbia spinosa and Eu. flavicoma ssp. occidentalis. In the south of France, Italy, Spain and Croatia. One ovipara was found in southern France in November (Remaudičre & Leclant 2000), but Spanish populations are habitually anholocyclic with large overwintering populations, and can live even under snow at high altitudes (Nieto Nafría & Mier Durante 2013). The description is very close to that of A. pareuphorbiae on Euphorbia in China. Acyrthosiphon moltshanovi Mordvilko Apterae are pale yellow or greenish; BL 2.4-3.7 mm. Described from an unidentified plant in Tajikistan, and subsequently reported from various plants in Pakistan and India; the true host is uncertain but is possibly Selinum wallichianum (= tenuifolium; David & Hameed 1975). Acyrthosiphon nigripes Hille Ris Lambers Apterae are green with black appendages; BL 2.8-4.0 mm. On Laserpitium siler in south and central Europe (Austria, Italy, Hungary). Populations on Peucedanum are regarded as a subspecies, A. nigripes ssp. peucedani (Bohzko); this was described as shiny brown by Szelegiewicz (1967b; specimens from P. officinale), but bright shiny green in the original description (from P. ruthenicum). Another possibly subspecific form occurs on Seseli; this was described and compared with A. nigripes s. str. by Pintera (1957), under the name A. superba Börner (which is a synonym of nigripes). The name A. nigripes ssp. blattnyi Pintera ex Szelegiewicz is applicable to this form. Acyrthosiphon ononis (Koch) Apterae are green or red; BL 2.7-3.9 mm. On Ononis spp. in Europe. Monoecious holocyclic with apterous males (Meier 1958). A close relative of A. pisum, and often regarded as a subspecies. Acyrthosiphon papaverisuctum (Zhang, Chen, Zhong & Li) Apterae are ivory white; BL 2.3-2.8 mm. On Papaver sominiferum in Gansu province, China (Zhang 1999, as Impatientinum). Perhaps a geographical variant of A. ilka. Acyrthosiphon pareuphorbiae Zhang Colour of apterae in life is unrecorded; BL c.2.5 mm. On Euphorbia sp. in Yunnan province, China. Acyrthosiphon parvum Börner Apterae are green to dark green, sometimes with dusky antennae, legs and siphunculi; BL 1.9-2.5 mm. On young growth, and later on undersides of leaves, of Cytisus, Cytisophyllum, Anthyllis and Genista in eastern, central and southern Europe (Poland, Hungary, Bulgaria, Czech Republic, Austria, Switzerland, Italy, Majorca). Acyrthosiphon pentatrichopus Hille Ris Lambers Colour of apterae in life unknown; BL 3.0-3.4 mm. Alatae are undescribed. On Rosa fendleri and Rosa sp. in western USA (Colorado, Utah). Fundatrices were collected on R. fendleri in May (Hille Ris Lambers 1974b). Acyrthosiphon phaseoli Chakrabarti, A.K.Ghosh & Raychaudhuri Apterae are dull white to slightly greyish; BL 2.7-3.0 mm. On undersides of leaves of a Phaseolus sp. in India (HP). Alatae are undescribed. A. pisivorum, described from Pisum sativum in China, is very similar, and may be a synonym (2n=14 for pisivorum; Chen & Zhang 1985a). [If Macrocaudus phaseoli Shinji, described from Phaseolus in Japan is rediscovered and proves to be an Acyrthosiphon, then this species (if not a synonym) will require a replacement name.] Acyrthosiphon pilosum Nieto Nafría, Aldea & Castro Apterae are green; BL 2.4-3.7 mm. On Ononis spp. in Mediterranean coastal areas of France and Spain. Oviparae were collected in mid-October (Nieto Nafría et al. 2015b). Acyrthosiphon pisum (Harris) Pea Aphid Plate 23d Apterae are green or pink, with antennae dark at apices of segments, often lightly wax-dusted (see influentialpoints.com/Gallery); BL 2.5-4.4 (-5.5) mm. On young growth and developing pods of many herbaceous and some shrubby or woody Leguminosae/Fabaceae, and an important pest of peas and alfalfa (Blackman & Eastop 2000, p. 218). Trees recorded from are Albizia julibrissin, Robinia pseudoacacia and Sesbania grandiflora. Certain other plants (Capsella bursapastoris, Reseda spp.) may be colonised under dry conditions. A. pisum is almost cosmopolitan. It is monoecious holocyclic in temperate regions, producing both apterous and alate males in differing proportions that are dependent on host race (Frantz et al. 2010). In Europe and Central Asia, there is a complex of races and subspecies with different host ranges and preferences (Müller 1985), more than one of which has been introduced to other parts of the world, and populations on different host plants (e.g. Vicia, Trifolium, Medicago) have diverged genetically to different extents (Hawthorne & Via 2001; Simon et al. 2003; Blackman & Eastop 2007; Peccoud et al 2008, 2009a,b, 2014; Peccoud & Simon 2010; Nouhaud et al. 2014; Duvaux et al. 2015; Eyres et al. 2017). Recent work has shown that several of the “host races” are reproductively isolated, and that diversification of these occurred much earlier than previously thought, predating Neolithic agriculture (Fazalova et al. 2018; Fazalova & Nevado 2020) The A. pisum complex harbours a rich community of secondary facultative symbionts, and the roles of these in relation to genetic diversity, host specialisation and resistance to parasitism have been the subject of extensive research in Europe and North America (e.g. Ferrari et al. 2011; Russell et al. 2013). The form usually found on Ononis
is here judged to be sufficiently distinct to warrant separate species
status (see A. ononis). Molecular
studies have singled out other host-specific forms which qualify for separate
species or at least subspecies status, on Cytisus
scoparius (and probably other
members of the tribe Genisteae including Spartium
junceum), and on Lathyrus pratensis
(Peccoud et al. 2009a,b). Available names for these are respectively spartii (Koch) and lathyri (Mosley). A. pisum is currently being studied intensively as the species
selected for the aphid genome project (Brisson & Stern 2006, Tagu et
al. 2010); a draft of its genome sequence has
been published (International
Aphid Genomics Consortium 2010 – and see Y. Li et al. 2019 for recent research). 2n=8. Acyrthosiphon
porrifolii (Börner) Apterae are presumably yellow-green; BL
1.5-1.8 mm. On Hieracium porrifolium in
Austria. Males are alate (Börner, 1950). Acyrthosiphon primulae (Theobald) Apterae are pale yellow or greenish yellow, shiny, with apices of antennae and legs dark (see influentialpoints.com/Gallery); BL 2.0-2.5 mm. Alatae have dark dorsal cross-bands and postsiphuncular sclerites. On undersides of leaves of Primula spp. in Europe, and introduced to New Zealand (Cottier 1953, as Aulacorthum), Tasmania (Heie 1994, as Microlophium) and California (BMNH collection, leg. R.C. Dickson). The generic position of this species is uncertain. 2n=16. Acyrthosiphon
pseudodirhodum (Patch) Apterae are pale cream to white; BL
2.2-3.5 mm. On various Rosaceae (Geum, Potentilla, Rosa, Spiraea) in north-eastern USA and
Canada. MacGillivray (1968) redescribed this species. Two other undescribed,
somewhat similar spp. occur in western North America, one on Chamaebataria millefolium in
California (BMNH collection, leg. P. Rude) and the other on Holodiscus discolor (California, BMNH
collection, leg. C. Lagace) and Cercocarpus
parvifolius (Colorado; BMNH collection, leg. F.C. Hottes & D. Hille Ris
Lambers). Both have R IV+V 0.9-1.0 × HT II (0.72-0.82 in pseudodirhodum), 4-6 accessory hairs on R IV (2 in pseudodirhodum), and longer hairs
(46-50 μm) on abdominal tergite 3 (14-20 μm in pseudodirhodum). The Chamaebataria
aphid has siphunculi 0.18-0.2 of body length , whereas the aphid on Holodiscus and Cercocarpus has very thick siphunculi 0.125-0.15 of body length
and only 2.5 –3.6 times longer than their width at midlength (6.0-7.9 ×
longer in pseudodirhodum and
5.3-5.8 times longer in the Chamaebataria
aphid). Life cycles of all three species are unknown, but the sample from Colorado on Cercocarpus included alate males. Acyrthosiphon
purshiae (Palmer)
Apterae are mid-green, shining or slightly frosted, with dark
appendages; BL 2.0-2.5 mm. On leaves and stems of Purshia tridentata in western USA (Palmer 1952). [This species was described in Macrosiphum, and DNA analysis by Foottit et al. (2008) suggests that it should probably be returned to that
genus.]
Acyrthosiphon ranunculum L.K.Ghosh Apterae are yellowish white, the antennae ringed with black, legs mainly pale, siphunculi brown-black , cauda pale; BL 2.2-2.6 mm. On stems, petioles and undersides of leaves of Thalictrum pauciflorum (= dioicum) in Himachal Pradesh, India. Acyrthosiphon rubi Narzikulov (= Metopolophium sonchifoliae Raychaudhuri, L.K. Ghosh & Das) Apterae are shining green; BL 1.6-2.9 mm. Alatae have 18-34 rhinaria on ANT III. On Rubus spp. and Fragaria sp., but apparently with polyphagous tendencies, because there are also rather numerous records from plants outside Rosaceae (Catamixis, Desmodium, Fagopyrum, Hieracium, Polygonum, Rumex, Sonchus), although at least some of these are probably misidentifications. In mountainous regions of Tajikistan, Nepal (BMNH collection, leg. K.C. Sharma) and northern India. Oviparae and alate males were described (on Rubus) from Himachal Pradesh, India by Raychaudhuri et al. (1980). The specimens from Rubus ellipticus in northern India (Uttar Pradesh) described by Stroyan & Nagaich (1964) as ssp. elliptici, are smaller and have a less rugose dorsal cuticle, but other specimens from this host in Nepal are typical rubi. 2n=12. Acyrthosiphon rubifoliae (Raychaudhuri, M.R. Ghosh & R.C. Basu) Apterae are pinkish green; BL 3.3-4.2 mm. Alatae have c.22 secondary rhinaria on ANT III and faint brown dorsal markings. Described from Rubus ellipticus (as a Metopolophium), and subsequently also recorded from Potentilla nepalensis (Raychaudhuri, L.K.Ghosh & Das 1980). In northern India (Himachal Pradesh, Nagaland, Sikkim). Acyrthosiphon rumicis Narzykulov in Narzikulov & Umarov Apterae are shining grass-green, BL c.4.7-4.8 mm. In large colonies on stems and undersides of leaves of Rumex spp. in Tajikistan. Acyrthosiphon sanguisorbae Seccombe Colour of aptera in life is unrecorded; BL 1.8-2.9 mm. On Poterium sanguisorba (= Sanguisorba minor) in Afghanistan. Alatae have 20-22 secondary rhinaria on ANT III. Placed as a subspecies of A. malvae by Remaudičre & Remaudičre (1997), but with several features that justify its status as a full species. Acyrthosiphon saussureae Pashchenko Colour of aptera pale green; BL 2.5-3.0 mm. On a species of Saussurea, feeding on undersides of leaves and readily falling from plant when disturbed (Pashchenko 2005). Other morphs and life cycle are unknown. In east Siberia (Kamchatka). Acyrthosiphon scalare (Richards) Apterae are pale green or yellow; BL 1.7-1.9 mm. Other morphs are unknown. On Potentilla fruticosa in Ontario, Canada (as Chaetosiphon). Acyrthosiphon (Tlja) scariolae Nevsky Apterae are pale yellow-green, finely wax-powdered; BL 2.2-2.8 mm. Described from Lactuca serriola (= scariola) in Central Asia, and specimens assigned to this species have subsequently been collected on Lactuca spp. (saligna, serriola, virosa) and Sonchus arvensis in Europe, Middle East and North America (Canada), but further work is needed to confirm the distinction from A. lactucae (q.v.). 2n=18. Acyrthosiphon shinanonum Miyazaki Apterae are green with black apices to antennae, femora and tibiae; BL 2.5-3.1 mm. On Geum calthifolium (var. nipponicum) in Japan. Acyrthosiphon soldatovi Mordvilko Apterae are green, with dark antennae and dusky yellow, black-tipped siphunculi; BL 1.9-3.2 mm. On Spiraea spp., especially salicifolia, in east Siberia. Also in north India (Simla: BMNH collection, leg. K. Narayanan), Kazakhstan (Kadyrbekov 2009a) and Tajikistan (Narzikulov & Umarov 1969); the latter population was monoecious holocyclic with alate males on S. hypericifolia, and regarded as a subspecies, A. soldatovi tadzhikistanicum. Acyrthosiphon sophorae
Narzikulov & Umarov Apterae are
rather large (about 2.4 mm), pale green with waxy bloom, on Sophora
spp. (alopecuroides, japonica, mollis) in Kazakhstan,
Tajikistan and Pakistan. The life
cycle is unknown. References:
Narzikulov & Umarov 1969, Eastop 1971, Naumann-Etienne & Remaudičre
1995. Acyrthosiphon supranubius Carnero & Nieto Nafría Apterae are powder green with antennae,
legs, siphunculi and cauda darker; BL 2.9-3.5 mm. On Cytisus supranubius, only known from the Canary Islands (Carnero
& Nieto Nafría 1995). The life cycle is unknown. [This species was
regarded as a possible synonym of A.
parvum by Eastop & Blackman (2006), but is now distinguished from
that species in the key to apterae on Cytisus.] Acyrthosiphon svalbardicum Heikinheimo Apterae are very variable in colour, pale pink, yellow, green or brown with darker dorsal markings; BL 1.2-1.9 mm. On Dryas octopetala in Spitzbergen. Monoecious holocyclic with oviparae and apterous males in early August (Strathdee et al. 1993). The alata was described by Simon et al. (2008). Chaubet et al. (2013) described two new aphidiine parasitoids from this aphid, one of them specific to it. Acyrthosiphon thracicum Tashev Apterae are green, BL 2.7-3.2 mm. On Euphorbia platyphyllos in Bulgaria, and also recorded from Czech Republic on E. polychroma (Holman 1965). Remaudičre & Leclant (2000) compared it with the closely related A. euphorbiae. Acyrthosiphon umarovi Narzikulov Apterae are green; BL 2.9-3.8 mm. On upper parts of stems of Astragalus sp. in Afghanistan. Alata is undescribed. It seems probable that this is a synonym of Acyrthosiphon gossypii. Acyrthosiphon vandenboschi Hille Ris Lambers Colour of apterae in life is unknown; BL 3.7-4.4 mm. Alatae have 42-60 secondary rhinaria on ANT III. On Potentilla glandulosa in California, USA. Acyrthosiphon vasiljevi Mordvilko Apterae are green; BL not recorded. On leaves of Cucurbita pepo in Central Asia. Similar to A. caraganae according to the original description, which does not provide enough information to include it in a key. Aphids identified as A. vasiljevi were collected on Sonchus asper by Rusanova (1942). Acyrthosiphon
wasintae (Hottes) Apterae are pale dull green; BL 1.4-1.7
mm. Living singly on undersides of leaves of Potentilla fruticosa in western USA (Colorado, Utah, Idaho,
Wyoming, Montana, Oregon) and Canada (Alberta; BMNH collection, leg. D. Hille
Ris Lambers). Monoecious holocyclic, with alate males (Palmer 1952).
Distinguished
by having five pairs of abdominal spiracles, whereas members of the other
genus of Adelgidae, Pineus, have only four distinct pairs. As in Pineus
the primary host of holocylic species is Picea, but the secondary
hosts are Abies, Larix, Pseudotsuga and Tsuga, never
Pinus. The galls on Picea are
cone-like, often resembling miniature pineapples. The complete life cycle of holocyclic
species takes two years (fig. 1), and
involves up to seven different morphs.
Several species are anholocyclic with a reduced number of morphs and
no host alternation, either living on Picea (in which case a gall may
or may not be formed by a "pseudofundatrix"), or living on what was
the original secondary host. The name Chermes was once
commonly applied to Adelges, but is
now suppressed (Eastop 1963). Some
authors have used Adelges in a narrow sense for the A. laricis
group, and placed other species in Aphrastasia, Dreyfusia, Cholodkovskya,
Gilletteella and Sacchiphantes, but this generic classification
is based on rather slight differences, and we use Adelges in its
broader sense to cover all these groups.
Recent molecular work (Havill et al., 2007) provides some
support for evolution in association with secondary host genera, with species
migrating to Abies clustering separately from those going to Larix
and Pseudotsuga, and the species associated with Tsuga apparently
forming a third group. Havelka et al.
(2019) analysed mitochondrial (COI) and nuclear (EF-1α) sequences of the
species in Lithuania, and discussed the taxonomic implications of their
findings. Favret et al.
(2015b) provided a taxonomic and nomenclatural catalogue of the world fauna. Carter
(1971) provided an account of the British species, and Carter (1976) keyed
the galls of British Adelgidae on Sitka spruce. Binazzi (2000) keyed the
winged morphs of Italian species. Other taxonomic accounts are by Börner
(1908), Börner & Heinze (1957), Annand (1928: North America), Heinze
(1962: central Europe), Binazzi & Covassi (1991: subgenus Dreyfusia
in Italy), Inouye (1953: Japan) and A.K. Ghosh (1984a: India). Havill &
Foottit (2007) and Havill et al.
(2007) reviewed adelgid ecology and evolution, and Sano & Ozaki (2012)
discussed the evolution of adelgid life cycles. Foottit et al. (2009a) and Žurovcová et
al. (2010) evaluated the use of DNA bar-coding of the mitochondrial COI
gene for clarifying adelgid taxonomy. Adelges (Sacchiphantes) abietis
(L.) is anholocyclic on Picea spp. (abies, glauca, jezoensis, koyamai, sitchensis),
with only two generations per year; a pseudofundatrix that induces a
pineapple-like gall (fig. 123A,
and see also influentialpoints.com/Gallery),
and yellow to yellow-green alate gallicolae that leave the gall in late
summer but do not generally disperse far, often laying eggs on the same or
neighbouring spruce trees. The galls
are hard, green, and covered with trichomes; Rohfritsch (1990) studied their
development in detail. Ewert (1967) found
differential levels of infestation by A. abietis galls among spruce
clones, Mitchell & Maksymov (1977) studied predators, Flaherty et al. (2010) and Pilichowski et al. (2014) studied factors
influencing gall size and distribution and development of gallicolae, and
Lasota et al. (1983) described a
heavy fungal attack on the galls.
Found throughout Europe, and in North Africa (Morocco), Central Asia
(Kirghizia; Kadyrbekov 2013c), India and North America. Japanese records of abietis should
probably be referred to A japonicus, and most records from Larix
are probably referable to A. viridis.
2n=18 (Steffan 1968). Adelges (Gilletteella) cooleyi
(Gillette) typically host-alternates between Picea spp. (engelmanni,
pungens, sitchensis) and Pseudotsuga spp. (macrocarpa,
menziesii) (fig. 64G). The galls on shoot tips of Picea are characteristically elongate,
with green, pink or red gall chambers and long needles protruding from them (fig. 123B,
and see also bladmineerders.nl/gallen). Apterous exules on needles of Pseudotsuga are covered in white
woolly wax (see influentialpoints.com/Gallery). Native to western
North America, but now occurring throughout Europe and North
America, and in Tasmania (BMNH collection, leg. V.F. Eastop). Cumming (1962)
described a monomorphic, non-galling, anholocyclic form on Picea glauca in Canada. An anholocyclic form also occurs on Pseudotsuga in both Europe and
California, to which Steffan (1970) appled the name coweni Gillette; see under that name for further information.
Parry (1978a,b; 1980) and Parry & Spiers (1982) have made extensive
studies of the population biology of A.
cooleyi in Scotland, Teucher (1955) studied natural enemies, and Stephan
(1987) reported differences in resistance of Pseudotsuga provenances in Germany. Ahern et al. (2008) compared
genetic variation and secondary host preferences of native western and
introduced eastern populations of A.
cooleyi in North America, and Ahern et
al. (2009) studied genetic variation in the indigenous western
populations. 2n=22 (Steffan 1968). Adelges (Gilletteella) coweni
(Gillette) Anholocyclic, living all
year on Pseudotsgua menziesii.
Alate sexuparae develop in spring, but these are all gynoparae and produce
only sexual females, so that without males the holocycle is defunct.
Described from California, and introduced to southern Europe. Roversi &
Binazzi (1996) provided a full account of the life cycle in central Italy,
where it has become a serious pest in stands of Douglas Fir. Adelges
(Gilletteella?) glandulae
Zhang Host-alternating between Picea spp. (brachytyla, likiangensis,
purpurea) and Abies ?faberi in
Sichuan and Yunnan provinces, China (Zhang et al. 1980, Chen &
Qiao 2012a). The galls are ovate, cone-like,
without associated needles; the form of the gall as well as DNA analysis
(Havill et al. 2007) indicate that
this species is wrongly placed in subgenus Gilletteella. The
hiemosistens morph on Abies
overwinters in second instar. This
species is described as anholocyclic, presumably in error as it is both
gall-forming and
host-alternating. The
fundatrix/?pseudofundatrix is undescribed. Adelges
(Dreyfusia) joshii Schneider-Orelli & Schneider (fig. 16G)
Anholocyclic,
with three generations per year (hiemosistens and two aestivosistens) on Abies pindrow in western
Himalaya. Adults are dark brown,
0.7-1.2 mm, covered with wax. Ghani
& Rao (1966) gave a detailed account of the morphology and life cycle. Adelges
(Sacchiphantes) karafutonis Kono & Inouye Galls of this species on young shoots of Picea jezoensis are small, with only
5-15 scales (Inouye 1953). Only the gallicolae are described, and the
secondary host (presumably a Larix
species), is unknown. For differences
from the closely related A. torii
see Eichhorn & Carter (1978).
Recorded from Sakhalin and Japan (Hokkaido). Adelges
(Sacciphantes) kitamiensis
(Inouye) Described from secondary host
morphs attacking Larix kaempferi in
Hokkaido, Japan, sometimes so heavily that the entire trunk of a 10-20
year-old tree is covered in its white wax wool. Hiemosistens immatures and adults are dirty
blue in colour (cf. viridis). For further details and differences from viridis see Inouye (1963). The small
(about 0.6-0.7 mm) sexuparae maturing in May-June leave Larix for a previously unknown primary host; no galls have been
found on Picea abies, even where
this tree occurs in mixed stands with L.
kaempferi. However, the probable
primary host generations of this species have now been identified, producing
pineapple galls on three Picea spp.
in Central Japan, and with a DNA barcode listed as that of “Adelges sp. A” in Foottit et al. (2009). Alate gallicolae
(1.5-2.5 mm) emerging from these galls deposited egg masses in the laboratory
on L. kaempferi. The first instars
hatching from these eggs agreed morphologically with the syntypes and
description of S. kitamiensis, but
were yellow in colour (Sano et al.
2011). Immature and adult fundatrices from galls have not yet been described. Adelges
(Dreyfusia) knucheli Schneider-Orelli & Schneider (fig. 16F)
Host-alternating
between Picea smithiana and Abies spp. (pindrow, spectabilis)
in western Himalaya (Pakistan and Kashmir), or anholocyclic on Abies pindrow where the primary host
is absent, e.g. in Murree Hills of Pakistan.
The pineapple or cone-like galls on P. smithiana vary considerably in size and shape. The sistens generations on Abies pindrow are dark brown with wax
filaments radiating a star-like fashion, and occur on main stems of older
firs as well as on branches of younger trees.
Progrediens apterae and alate sexuparae develop on new-grown needles, the
latter without secreting wax. Ghani
& Rao (1966) provided detailed descriptions of all stages and
morphs. Predators were discussed by
Pschorn-Walcher (1964). [Stebbing's
(1910) account of Chermes himalayensis
probably applies to this adelgid; but see Schneider-Orelli & Schneider
(1954).] Adelges
laricis Vallot (=strobilobius
Kaltenbach) Plate
1a, 1b The oldest-named member of a taxonomically
very difficult complex of adelgids which have host alternation between Picea and Larix, or have an incomplete cycle restricted to either the
primary or the secondary host. Galls
of laricis sensu stricto on Picea are typical of the group (fig. 123D,
and see also bladmineerders.nl/gallen);
they are globular or ovate, with a cream or ivory coloration and waxy
texture, becoming pink or brown just prior to opening. They may be terminal, or have some growth
of the shoot beyond the gall. Gall
development was described in detail by Rohfritsch (1990). Predation within galls by dipterous larvae
was described by Mitchell & Maksymov (1977). Gallicolae of A. laricis (fig. 64F) migrate
to Larix in June-July; they are 1-2
mm, blackish, and secrete little or no wax.
The hiemosistens generation on Larix
in spring is dark grey without wax, but the progrediens apterae left after
departure of the sexuparae in June-September produce abundant wax and
honeydew; these deposits from a heavy infestation cause the foliage to turn
blue (Börner & Heinze 1957; Carter 1971). See influentialpoints.com/Gallery
for further information and photographs. A.
laricis sensu stricto occurs in Europe and North America and is recorded
from Picea abies, koraiensis, mariana, rubens, sitchensis (primary hosts), and from Larix decidua, ×eurolepis, kaempferi,
laricina (secondary hosts). 2n=20
(Steffan 1968). Li & Ts’ai (1973) provided a very
detailed account of an adelgid under the name A. laricis in north-east China, with primary host Picea obovata. Numerous other names have
been applied to members of the laricis
group, some of which were only described from one or two morphs and are
possibly not distinct species: aenigmaticus Annand Only the apterous ?progrediens morph is
described by Annand (1928). It was
collected on Larix laricina in
Maine, USA, and distinguished from laricis
by its smaller, rounded dorsal sclerites bearing round, separate wax pores,
and by the short and thick third antennal segment. Similar specimens were collected by F.W.
Quednau on the same host in Quebec, Canada (F.W. Quednau, pers. comm. 1971). Life cycle is unknown. diversis Annand Progrediens apterae collected on cones of
European larch (Larix decidua) in
Oregon, USA were thinly covered with short wax threads (Annand 1928). Only the first instar ?sistens, progrediens
and immature ?sexupara are described.
The progrediens is distinguished by having no sclerites on abdominal
tergites 5-8 and only small sclerites with few wax pores on the more anterior
tergites. Not recorded since the
original description. geniculatus
(Ratzeburg) This is the name currently applied to
populations that build up large populations on Larix decidua in central Europe (Austria, Germany), causing
discoloration, deformation and eventual loss of the needles and also damaging
buds (Perny 2004). A. geniculatus
had previously been synonymised with A.
laricis, but was again given separate species status by Steffan (1968) on
account of its anholocycly and apparent inability to produce alate sexuparae.
However the assumption that the only populations reaching damaging levels on
larch are anholocyclic and can therefore be identified as geniculatus may be unwarranted, as
successive generations of progredientes of laricis persist on larch and can build up heavy infestations
(Carter 1971). Further work also seems necessary to confirm that A. geniculatus is functioning as a
fully independent, permanent parthenogen, and that its separate species
status is justified.. isedakii
Eichhorn Host-alternating between Picea jezoensis (var. hondoensis) and Larix kaempferi in Japan (Eichhorn & Carter 1978). The gall on Picea is similar to that of laricis,
but larger. The gallicola has a red
body and greenish wings, is smaller (1.0-1.5 mm) than the non-migratory
gallicola of A. japonicus (see
below), and has no wax glands on the head.
Only the first instar hiemosistens (neosistens) has been described
from Larix. japonicus
Monzen (fig. 64L) Anholocyclic,
pseudofundatrices induce cone-like galls on shoot tips of Picea jezoensis and P. sitchensis in Japan and Sakhalin
(Inouye 1953; Paschenko 1988).
Gallicolae emerging from galls in August-September are large (1.9-2.8
mm) and with or without cephalic wax glands (Eichhorn & Carter 1978);
some do not disperse, so that individual trees may develop heavy levels of
infestation (Kamata 1986). The cause of the relatively slow larval
development in the galls was studied by Tabuchi et al. (2009). Sano et al.
(2008) have now shown that in northern Japan there is also a holocyclic form
of this species that migrates to Larix
kaempferi; they reared and described alate gallicolae and first instar
exules of the migratory form, and compared the two life cycles. karamatsui Inouye Anholocyclic on secondary hosts (Larix kaempferi, gmelinii) in Japan, Sakhalin and Korea. Sistens, apterous progrediens and sexupara
are described (Inouye 1953).
Sexuparae, small and dark brown to black with abundant wax, fly to Picea in Japan, but die or produce
eggs which do not hatch (Eichhorn & Carter 1978); an analogous situation
to that with A. tsugae on Tsuga in USA. lapponicus (Cholodkosky) Anholocyclic, pseudofundatrices inducing
galls on both native and exotic Picea
spp. Dark red gallicolae about 1.6 mm
long energe from galls in June-July and lay reddish eggs on needles of Picea with little or no secretion of
wax. In northern and central Europe,
and more recently recorded from Kirghizia (Gabrid 1981). Differential infestation of Picea spp. in northern Finland was
studied by Häggman and Rousi (1986). lariciatus
Patch Host-alternating between Picea spp. (abies, glauca, mariana, pungens) and Larix spp.
(laricina, lyalli) in North America (Alberta, Manitoba, Saskatchewan,
Utah). Galls are globular, often
sub-terminal and to one side of shoot, green when young to red and brown when
mature. Cumming (1968) gave a detailed
account of morphs and life cycle in Canada. oregonensis
Annand Only known from the original
description of sistens and progrediens morphs from Larix occidentalis in north-western USA (Oregon, Washington,
Montana). Differences from laricis were discussed by Annand
(1928). It feeds on the needles and on
the twigs at the needle bases. The
primary host, if any, is unknown. potaninilaricis
Zhang Host-alternating between Picea retroflexa and Larix potaninii in Sichuan,
China. Described as a subspecies of laricis on the basis of differences in
number and distribution of wax glands (Zhang et al. 1980). tardoides (Cholodkovsky) Host-alternating between Picea abies and Larix sibirica in north-eastern Europe. The gallicolae migrate to the secondary
host in July-August, later than laricis,
and unlike those of laricis,
secrete abundant wax (Börner & Heinze 1957, p.346). tardus
(Dreyfus) Anholocyclic on Picea spp. throughout continental
Europe. The gallicolae emerge from
late June to September, later than lapponicus,
and lay eggs on exposed parts of the trees under a covering of white
wax. Due to non-dispersal of the
gallicolae, the galls may become very numerous and can severely affect growth
of young spruce trees. Baurant (1968)
studied the phenology of A. tardus
in relation to Picea abies in
Belgium, and suggested that its synchronisation with the host might be upset
by selection of appropriate spruce varieties.
2n=20 (Steffan 1968). Adelges
(Dreyfusia) merkeri Eichhorn Host-alternating between Picea orientalis and Abies alba, with a rather restricted
distribution in Europe (Germany, southern Sweden, one record from Austria);
origins unknown. Galls on Picea resemble those of A. nordmannianae but are usually
larger (7-22 mm) and open in June-July (Eichhorn 1975). On Abies,
colonies are found on the trunk and crown region of older firs, or on all
parts of young and seedling trees, where damage may be severe with swelling
and stunting of infested branches, especially at their bases. As well as the hiemosistens there are one
or even two aestivosistens generations.
Adults sistentes generally have less wax wool than either nordmannianae or piceae. Pschorn-Walcher
& Zwolfer (1960) provided ecological notes and Eichhorn (1968) studied
population dynamics on Abies,
especially the influence of natural enemies. [ Havill et al. (2020)
propose on morphological and molecular evidence that A. merkeri is a hybrid of A.
piceae and A. nordmanniae, and
therefore that it is an invalid species. However, it is now well established
that new species can arise by hybridisation, and A. merkeri (like A. prelli
– see below) has a viable sexual phase and consistent biological
characteristics, indicating that it is functioning as a good species.]
Adelges
(Dreyfusia) nordmannianae (Eckstein) (=nüsslini Börner) (fig. 10B, fig. 16C)
Host-alternating
between Picea orientalis and Abies spp. (alba, bornmuelleriana, cilicica, nordmanniana, etc.).
Believed to be endemic and relatively uninjurious in the Caucasus and
East Pontus mountains (Russia and Turkey), where there are mixed stands of Picea orientalis and Abies nordmanniana, but causing severe
damage to Abies where non-endemic
in Europe, North America, New Zealand (see Zondag 1982) and Tasmania. Galls on Picea (fig. 123C)
are 2-15 mm long, terminal, globular, pinkish and resembling strawberries
when young, later becoming greenish with red or purple coloration at bases
and tips of scales. They mostly open
in June (Eichhorn 1975). There is
never more than one aestivosistens generation, and even this is lacking in
some populations. On Abies alba there appear to be two
forms of A. nordmannianae:
"typica" which infests branches, twigs and needles of young trees
in sunny dry locations - the attack of the progrediens generation causing
severe deformation of needles and shoots and often killing the tree (see
Varty 1956); and form "schneideri" which infests the trunks of
older firs 30-120 years old in forests (studied by Pschorn-Walcher &
Zwolfer 1958, and Eichhorn & Pschorn-Walcher 1972). The latter form would not transfer to young
trees (Schneider-Orelli, Schaeffer & Wiesman 1929). Colonies of sistentes on stems are black
with a grey wax bloom and only a fringe of wax wool (cf. A. piceae). Eichhorn
(1969a) studied the natural enemies of A.
nordmannianae in Turkey, and also worked on morph determination (Eichhorn
1969b), life cycle (Eichhorn 1973, 1991) and embryonic development (Eichhorn
1970). Most recent ecological studies of this species have been in Austria
(Stürzer-Gilbert 1982; Bauer-Schmid 1983). For a review of natural enemies and potential for biological control see
Ravn et al. (2013). Analysis of
mitochondrial and nuclear DNA sequences failed to reveal any consistent
differences between nordmannianae, piceae and prelli (Ravn et al. 2013).
However, analysis of microsatellite DNA of samples from the Caucasus, Europe
and North America discriminates between nordmannianae
and piceae, and also provides
evidence that hybridisation between these two has occurred to give sexually
viable new taxa such as merkeri and
prelli (Havill et al. 2020). 2n=22 (Steffan 1968). Adelges
(Aphrastasia) pectinatae (Cholodkovsky) (fig. 16A)
Host-alternating between Picea (abies, obovata) and Abies spp. (alba, balsamea, concolor, etc.), in Europe (Sweden,
Finland, Latvia, Ukraine) and east to Siberia. Recorded also from east Norway on 10
species of Abies, although the gall
generations on Picea were not found
there (Austarĺ 1990). The galls (fig. 123E)
are compact, cone-like, dull green turning reddish-brown before opening,
about 10-17 mm long. Reddish-brown
gallicolae fly to Abies in
June-July. Feeding on Abies is restricted to the
needles. Frolowa (1924) studied
cytology of egg and sperm maturation in this species. 2n=20 (Steffan 1968). Populations alternating between Picea spp. (glehni, jezoensis,
etc.) and Abies (sachalinensis, veitchii) in Japan are regarded as a separate subspecies, A. pectinatae ishiharai Inouye
(Eichhorn & Carter 1978). Its
morphs and life cycle are described in detail by Inouye (1953). Fang (1986) described the pest status of
what is possibly this subspecies on Picea
jezoensis, P. koraiensis and Abies nephrolepis in China. An anholocyclic form in the pectinatae group, Adelges pindrowi Yaseen
& Ghani, occurs on Abies pindrow
in Pakistan. Information about its
life cycle is provided by Yaseen &
Ghani (1971). Adelges
(Dreyfusia) piceae (Ratzeburg) Balsam
Woolly Aphid (fig. 16D)
Anholocyclic
on Abies spp. (alba, balsamea, cilicica, fraseri, etc.), with one hiemosistens and usually one
aestivosistens generation. Sexual generations and galls on Picea orientalis have not been described, although there is now
evidence frrom microsatellite DNA
studies that sexual reproduction may sometimes occur if P. orientalis is available, and that hybridisation may occur with
A. nordmannianae (Havill et al. 2020). Throughout Europe, and
introduced to North America, and also to Chile (Fuentes-Contreras et al. 1997). In favourable conditions
a second, or even a third, aestivosistens generation may occur
(Pschorn-Walcher 1964), and in North America there is sometimes also a
generation of progrediens apterae and alatae. On A. alba in Europe this adelgid normally infests the trunk and
larger branches of older trees (30-100 years), and is of little economic
importance. Introduced populations in North America, however, attack stems
and buds of the crown region of Abies
balsamea and A. fraseri,
causing extensive injury. Young trees
of A. balsamea are killed by
"gout disease" (excessive shoot swellings near the buds); heavy stem attack may give rise to severe
transpiration stresses by the
formation of rotholz or compression wood (Balch 1952), causing crown dieback
and a degrade of the timber (Carter 1971).
Hain et al. (1983) studied
the host's defence reaction against stem attack. See influentialpoints.com/Gallery
for illustrations and further information. The economic importance of A. piceae has resulted in an extensive
literature; Schooley & Oldford (1981) provide an annotated bibliography,
and Johnson & Lyon (1988) gave a short, well-illustrated account. Biological control measures up until 1958
were summarised by McGugan & Coppel (1962). Foottit & Mackauer (1983) used
morphometric studies to distinguish and key three forms of A. piceae in North America, thought to
represent separate introductions. They
designated these as subspecies (p.
piceae, p. canadensis and p. occidentalis), although it is
questionable whether the subspecies category is appropriate for anholocyclic
taxa. The form known as canadensis is similar or identical to
populations of piceae causing
compression wood in Abies grandis
in Scotland (Busby 1964). Binazzi
& Covassi (1991) differentiated an anholocyclic population on Abies rosi in Italy as a new species, A. (D.) nebrodensis, which has now
also been given subspecies status, as A.
(D.) piceae nebrodensis (Havill et
al. 2020). 2n is unknown. Adelges
(Dreyfusia) prelli (Grosmann) (fig. 16E)
Host-alternating
between Picea orientalis and Abies nordmanniana or (rarely) Abies cephalonica. Like A.
nordmannianae, A. prelli is
indigenous to Turkey (Eichhorn 1969) and introduced to central and western
Europe. The galls are bluish-green,
larger than those of A. nordmannianae
(6-30 mm long), and open later, in July-August (Eichhorn 1975). On Abies
nordmanniana in western Europe a hiemosistens and one or two
aestivosistens generations occur, almost exclusively attacking the crown
region of mature trees (Eichhorn 1956).
The hiemosistens on young shoots is more sclerotized and has less wax
than that on stems or branches (Eichhorn 1964). North American populations of A. piceae on Abies balsamea closely resemble A. prelli, which has not however been positively identified from
that host (or from Abies alba). [ Havill et al. (2020) propose on morphological and molecular evidence
that A. prelli is a hybrid of A. piceae and A. nordmanniae, and therefore that it is an invalid species.
However, it is now well established that new species can arise by
hybridisation, and A. prelli (like A. merkeri – see above) has a viable
sexual phase and consistent biological features, indicating that it is
functioning as a good species.] Adelges
(Sacciphantes) roseigallis (Li &
Tsai) Described from “red-barked Picea” (P. koraiensis?) in north-east China (Heilungkiang province),
where there is host alternation to a managed species of Larix, presumably L.
gmelinii (Li & Ts’ai 1973). The galls on Picea are bright red, smaller than those of A. viridis and resembling cypress cones; they are attached to
bases of old branches. Alatae gallicolae are lustrous black, not powdered
with wax (cf. viridis). Li &
Ts’ai (1973) described other differences from A. viridis in colour and appearance in life of various stages,
and in life cycle and fecundity. In Gansu province this species is reported
to be anholocyclic on Picea spp. (asperata, crassifolia; Du et al.
1998, Chen & Qiao 2012a). Information is currently insufficient to
include it in the key to aphids on Picea. Adelges
(Sacciphantes) segregis
(Steffan) Anholocyclic on Larix decidua in central Europe, with
two generations per year, one alate and the other a
"pseudohiemosistens". Closely related to A. viridis, but the overwintering first instar larva has a
shorter rostrum than the neosistens of viridis
(Steffan 1961a). Adelges
(Dreyfusia) todomatsui (Inouye) (fig. 16H)
Anholocyclic
on Abies sachalinensis (incl. var. mayriana) and A. veitchii in Hokkaido, Japan, with a hiemosistens and one or
possibly two aestivosistens generations (Eichhorn & Carter 1978). No progrediens (needle-dwelling) morph has been
found. Inouye (1953) observed that two
biological forms occur that are indistinguishable morphologically; a
stem-dwelling form on young and mature
(pole-stage) trees, and a bud-dwelling form that lives under the
scales of the previous year's shoots and causes gall-like enlargement of
buds. Predators were studied by
Pschorn-Walcher (1964). Adelges
(Sacciphantes) torii (Eichhorn) Host-alternating between Picea jezoensis var. hondoensis and Larix kaempferi in Honshu, Japan.
Galls are small and reddish, similar to those of A. karafutonis, opening in August. Eichhorn & Carter (1978) provided a
detailed account. Adelges
(Annandina) tsugae Annand
Hemlock Woolly Aphid (= funitecta of
Blackman & Eastop 1994; see Favret et
al. 2015a)
Populations with host alternation between Picea and Tsuga occur
in Japan and China. The galls and generations on Picea are apparently still not properly described. In Japan the
primary host is Picea polita (and
possibly P. jezoensis hondoensis;
McClure 1996), whereas in China the primary hosts are P. likiangensis and P. brachytyla (Foottit et al. 2009a), with possible A. tsugae galls also reported on P. morrisonicola in Taiwan (Havill et al. 2016). Secondary hosts are Tsuga diversifolia and T. sieboldii in Japan, T. chinensis, T. dumosa and T. forrestii in
China, and T. formosana in Taiwan (Inouye 1953; Havill et al. 2016). A. tsugae
is recorded from North America, India, Japan, China and Taiwan. The
introduced populations in North America are anholocyclic on both native and
introduced Tsuga spp. DNA analysis
has now provided evidence of separate introductions to western and eastern
North America, the more recent introduction being particularly damaging to
the native eastern species T.
canadensis and T. caroliniana
(Havill et al. 2016; and see
McCarty & Addesso 2019 for a comprehensive review). McClure (1989a)
described sistens and apterous progrediens morphs and also alate sexuparae
from populations on T. canadensis in
Connecticut, USA; the sexuparae migrated to Picea spp. and laid eggs, but sexuales failed to develop, even on
Picea polita, the recorded primary
host in Japan. Joseph et al. (2011) and Tobin & Turcotte
(2018) reported on the phenology of A. tsugae respectively in northern
Georgia and in the Central Appalachians. Analysis of mitochondrial and
microsatellite DNA has shown that the lineages that are severely damaging to
the two indigenous Tsuga spp. in eastern North America probably
originated from southern Honshu, Japan, where the native secondary host is T. sieboldii. Other lineages, some of
which had associations with particular Tsuga spp., were found
elsewhere in Japan and in mainland Asia and western North America, where
trees are not significantly damaged (Havill et al. 2006, 2007, 2016). Allozyme analyses have also been used to compare
populations in eastern and western USA with those in China (Sánchez &
Keena 2009). Evidence is accumulating that at least Chinese and Japanese
populations should be regarded as distinct species. An anthocorid bug, Tetraphleps galchanoides, has been studied in China
as a potential biocontrol agent (L. Li et al. 2011), and there is an
ongoing biological control programme in north-western USA to find adelgid
predators to use against A. tsugae (Ross et al. 2017, Rose et al. 2019). Adelges
(Cholodkovskaya) viridanus
(Cholodkovsky) Anholocyclic on Larix spp. (decidua, x eurolepis, gmelinii, kaempferi, sibirica) in
Europe (Sweden, UK, France, Germany, Italy, Ukraine) and east to China, Korea
and Japan, although somewhat sporadic or local in occurrence. The life cycle is variable, probably
depending on latitude and/or temperature.
The overwintering first instar sistens is yellowish-green to green and
usually under bark on the stem. The
large (1.8-2.5 mm) adult sistentes, yellowish-green and surrounded by wax
secretion, occur in May-June. Their
eggs develop either as overwintering hiemosistentes or as alate
progredientes; first instars of the latter morph are very active and move to
tender young extension shoots to feed at the needle bases. The adult alate progrediens is quite large
(2.0-2.6 mm), grey-green, and mostly non-migratory, laying eggs on the
needles. These mostly develop as
hiemosistentes but, in France and Germany (Gaumont 1954, Steffan 1964), some
of them develop into a second generation of alate progredientes, and in
suitable conditions these may even give rise to a third partial alate
progrediens generation. 2n=24 (Steffan
1968). In northern Russia, populations of the
closely-related A. (C.) viridulus
(Cholodkovsky) occur on Larix sibirica,
and these seem always to have a single annual (hiemosistens) generation, no
alatae being known. Its overwintering
first instars have fewer wax pores on posterior abdominal segments than A. viridanus. Adelges
(Sacchiphantes) viridis (Ratzeburg) Host-alternating between Picea spp. (abies, koraiensis, orientalis, sitchensis, etc.; see Weiss 1955), and Larix spp. (decidua, x eurolepis, gmelinii, kaempferi). Steffan (1961a) made a detailed study of
this adelgid in Germany (but see also Eichhorn 1989), and Li & Ts’ai
(1973) described its life-cycle in north-east China.. The pineapple galls on Picea are similar to those of A.
abietis but open earlier, in July or early August (see influentialpoints.com/Gallery). Miszta (1987) has studied their volatile
oil composition in comparison to normal plant tissue. Immatures within the gall, and the alate
gallicolae when they emerge, are reddish-yellow to brown, powdered with white
wax (cf. A. abietis and A. roseigallis). Hiemosistens immatures and adults are green
or brown. Sexuparae developing on Larix the following spring are pale
greenish with sparse wax, and their feeding causes yellowing and kinking of
the newly-grown needles (Carter 1971).
The return migration to Picea
is quite early, in May-June. It occurs
throughout Europe, and is also recorded from China where it migrates from Picea koraiensis to Larix gmelinii (Fang et al. 1983). 2n=18 (Steffan 1968).
Aiceona is often
placed in the Anoeciinae, but it is not closely related to Anoecia and requires a separate
subfamily. There are about 14
recognised species, all in east and south-east Asia, mostly associated with
Lauraceae. Several species are
described from unidentified hosts.
Unusually, oviparae as well as males are alate, and they may be found
in colonies at almost any time of year.
Siphunculi are well-developed as hairy cones in all female morphs, but
strangely absent in males. Accounts of
Aiceona are available for India
(M.R. Ghosh & Raychaudhuri 1973, A.K. Ghosh 1988), north-east India
(Raychaudhuri et al. 1980d), China (Qiao & Zhang 2002c) and Japan
(Takahashi 1960a). The genus is
clearly still in need of further revision. Aiceona
actinodaphnis Takahashi Apterae are 2-3 mm, purplish-black dusted with
white wax powder, living under leaves and on new growth of various Lauraceae
(Actinodaphne, Cinnamomum, Lindera, Litsea, Phoebe). Alatae have dark-veined smoky wings with a
pale area between the tip of the pterostigma and Rs. Sexual morphs are unknown. Recorded from China, Okinawa and Japan
(Tseng & Tao 1938). A related
species with paler wings, A. siamensis,
is described from unidentified Lauraceae in Thailand (Takahashi 1941a). Aiceona
himalaica Miyazaki
Appearance in life unrecorded, apterae 2.4-2.8 mm; cleared and mounted
apterae have a pale body contrasting with the dark hind tibiae. Apterous viviparae, alate oviparae and
males are described. Collected in July
in Nepal, from Engelhardtia spicata
(Juglandaceae), an unusual host association for this genus (Miyazaki 1977).
It needs to be established whether this species is distinct from A.
pseudosugii (see A.K. Ghosh, 1988) Aiceona
japonica Takahashi
Apterae are about 3 mm, blackish-brown, covered with white wax powder,
on leaves and young growth of Lauraceae (Actinodaphne
lancifolia, Cinnamomum camphora,
?Litsea sp.) in Japan, and also
recorded from Korea (Paik 1972). The
wings of the alata are uniformly dusky without any clear area. Alate oviparae
and males occur along with apterous and alate viviparae in May (Takahashi
1960), although Miyazaki (1977) expressed some doubt about Takahashi's
description of the ovipara of this species. The male genitalia were described
and illustrated by Wieczorek et al.
(2012). Aiceona
malayana Takahashi
Apterae pale greenish-yellow, 2.2-2.5 mm, attacking young leaves of ?Actinodaphne sp. near Kualar Lumpur,
Malaya. Alatae have wings clear or
only faintly dusky. Distinguished by
its short antennal hairs, this species is only known from the original
collection, when it was recorded erroneously as A. osugii (Takahashi 1950,1960a).
Sexual morphs and life cycle are unknown. Aiceona
(Subaiceona) manipurensis (Singh
& Raychaudhuri) Apterae
yellowish, BL 2.0-2.9 mm. Developing large ant-attended colonies in February
to May on Persea odoratissima in Manipur, India. A.K. Ghosh (1988)
provided a redescription. Sexual morphs and life cycle unknown. Aiceona
osugii Takahashi
Apterae are yellowish green, with pale antennae and legs, BL about 2.5
mm. Described as feeding on undersides of leaves of an undetermined tree in
Taiwan, and subsequently recorded from Lindera communis (Tao 1999).
Alatae have dusky wings and dark spinal sclerites or short cross-bands on ABD
TERG 1-5. Alate oviparae and males were described from Sichuan province,
China, collected on Lindera sp. at
the end of October (Qiao & Zhang 2002c). Alate oviparae (with spinal
sclerites merged to form a broad dark abdominal spinal stripe) have also been
collected in May in Taiwan on Persea
sp., and (presumably vagrants) on a plant of the Amaranthaceae (BMNH
collection, leg. S.E. Halbert). A. parvicornis may be a synonym. Aiceona
pallida A.K. Ghosh & Raychaudhuri Apterae are creamy yellow to dirty grey,
BL 2.1-2.7 mm. In ant-attended colonies on tender shoots and undersides of
young leaves which are rolled longitudinally. Alatae have pale wings with
dark pterostigma. The only identified host is Machilus glaucescens (= Persea villosa) (A.K. Ghosh 1988). In West
Bengal and Meghalaya, India. Sexual morphs and life cycle unknown. Aiceona
parvicornis Miyazaki Appearance in life is unknown; mounted
apterae (about 2.7 mm) have dark head, prothorax, coxae and femoro-tibial
joints. Alatae have dusky wings. Collected in May from Lindera pulcherrima in Nepal (Miyazaki 1977). Sexuales and life cycle are unknown. It
needs to be established whether this species is distinct from A. osugii
Takahashi. Aiceona
pseudosugii David, Sekhon & Bindra Apterae are 2-3 mm, pale yellow with dark
hind tibiae, occurring in small colonies on undersides of young leaves, with
attendant ants. Alatae have
unpigmented wings. The only identified
host is Persea odoratissima (David,
Sekhon & Bindra 1970). Recorded from Himachal Pradesh and West Bengal,
India. Sexuales and life cycle are
unknown. [Differences from A. longisetosa M.R. Ghosh & Raychaudhuri, described
from an unidentified plant in West Bengal and synonymised with pseudosugii
by Eastop & Hille Ris Lambers (1976), are listed by A.K. Ghosh (1988). Another
possible synonym is A. robustiseta
(q.v.).] Aiceona
retipennis David, Narayanan & Rajasingh Apterae are 2.5-2.8 mm, yellowish-white,
on undersides of leaves and young shoots (L.K. Ghosh 1972). Alatae have very dark wings with a clear
patch between the pterostigma and Rs like that in A. actinodaphnis, and spinal sclerites
of short cross-bands on ABD TERG 1-5.
Distinguishing features of this species are the very long fine
antennal hairs and short last rostral segment (R IV+V) only 0.6-0.7 times HT
II. Pal & Raychaudhuri (1977)
describe the alate male, collected in February. Alate oviparae (with spinal
sclerites often merged between segments to give a broad longitudinal spinal
stripe) have been collected in July
(BMNH collection, leg. V.D. Verma).
Identified hosts in the Lauraceae are Machilus gamblei (BMNH collection, 1 aptera leg. K. Narayanan), Persea sp. (Chakrabarti, Saha &
Mondal 1988), and Phoebe lanceolata; the various records from plants
in other families seem doubtful or due to casual occurrences. Widely distributed in northern India. 2n=18 (Khuda-Bukhsh 1980). Aiceona
robustiseta M.R. Ghosh & Raychaudhuri Apterae are 2.0-2.5 mm, creamy-yellow with
a light dusting of wax, in colonies on young growth attended by ants (M.R.
Ghosh & Raychaudhuri 1973). Alatae
have unpigmented wings. Alate
oviparae, collected in February and June, were described by Pal &
Raychaudhuri (1977). A robustiseta was described from Litsea polyantha (= monopetala) in West Bengal, India.
[Specimens from Machilus chekiangensis collected
in Hong Kong, from Machilus gamblei (=
Persea bombycina) in
Assam, India and from Eurya nitida in
Thailand, have been provisionally identified as this species (Martin &
Lau 2011). However , apart from the lower number of secondary rhinaria in
alatae, the differences from A.
pseudosugii, a pale species with dark hind tibiae described from Persea odoratissima, are minimal.] Aiceona
titabarensis (Raychaudhuri & A.K. Ghosh) Plate 5c, d Apterae are 2.1-2.7 mm, dirty brownish to
grey, dusted with wax powder, often heavily infesting the undersides of
leaves and young shoots of Litsea
polyantha (Basu & Hille Ris Lambers 1968), with or without attendant
ants. Alatae have smoky wings as in A. actinodaphnis and A. retipennis, but the clear area
adjacent to the pterostigma is narrower and less clearly defined. The species was originally described (in Lachnus) from Heteropanax fragrans (Araliaceae), a host which is unusual enough
to need confirmation. Alate oviparae
can be found in colonies with apterous and alate viviparae at least from June
to December (M.R. Ghosh & Raychaudhuri 1973, as A. litseae). Recorded from Sikkim and West Bengal, India,
and from Hong Kong and Vietnam (BMNH collection).
An
east Asian genus with Phorodon-like
projections on the antennal tubercles, but the head is not spiculose, the spiracular
apertures on abdominal segments 1-5 are remarkably small, there are typically
paired processes on abdominal tergite 7 and a larger median one on tergite 8,
the subgenital plate has a posterior projection, and the cauda has a
characteristic knobbed shape. Alatae look very different from apterae and
have numerous tuberculate secondary rhinaria, peculiar hypha-like projections
associated with the primary rhinaria, and much thinner, clavate siphunculi.
Most species are only known from Polygonaceae,
but one is known to have a sexual phase on Enkianthus. M.R. Ghosh et al. (1976) reviewed the species in
east India, the Japanese species were reviewed by Takahashi (1961d) and
Miyazaki (1971), and the Chinese species were reviewed by Qiao et al. (2006c). Akkaia
bengalensis A.N.
Basu (Fig.42u) Apterae are orange with black siphunculi;
BL 2.1-2.5 mm. Alatae have secondary rhinaria distributed III 25-37, IV 9-11
(M.R. Ghosh et al., 1976). On undersides
of leaves of Polygonum and Persicaria spp. in India (Meghalaya,
Sikkim, West Bengal) and probably Japan, as the aphid illustrated and
described by Moritsu (1983) as A.
odaiensis may be this species. Close to A. taiwana, and the distinction between the two needs additional
verification. Akkaia
neopolygoni M.R.Ghosh,
A.K.Ghosh & Raychaudhuri (Fig.42w) Apterae are green to deep green; BL c.
2.4-2.5 mm. Alatae are undescribed. On apical parts and undersides of leaves
of Polygonum and Persicaria spp. in India (Arunachal Pradesh, Sikkim, W Bengal). Akkaia
odaiensis Takahashi Apterae are yellow with all appendages
pale; BL 1.8-2.0 mm. Alatae are undescribed. On Polygonum thunbergii in Japan, and Tao (1999) records its
occurrence in China. Moritsu’s (1983) account refers to another species,
probably A. bengalensis. Akkaia
polygoni Takahashi Plate 20b (Fig.42s,v) Apterae are yellow or orange-yellow,
appendages concolorous with body; BL 1.9-2.4 mm. Alatae have secondary
rhinaria distributed III 40-48, IV 14-17, V 0-3. On stems and upper sides of
leaves of Polygonum and Persicaria spp. in Japan, China, Taiwan and Korea (BMNH
collection, leg. K.S. Boo). Heteroecious, with sexual phase on Enkianthus spp. and Pieris japonica (Takahashi 1961d,
Miyazaki 1971). 2n=24. Akkaia
pseudopolygoni Qiao, Jiang
& Ren Colour of aptera in life
unrecorded; BL 1.7-2.0 mm. Alatae are unknown. On an unidentified Polygonum sp. in Sichuan province,
China (Qiao et al. 2006c). Akkaia
sikkimensis Agarwala
& Raychaudhuri Colour of aptera
unknown; BL 2.5-2.6 mm. Described from Raphanus
sativus, but this will not be the true host. India (Sikkim). Akkaia
taiwana Takahashi (Fig.42r,t) Apterae are pale brownish white with jet
black siph.; BL 1.8-2.1 mm. Alatae have secondary rhinaria distributed III
22-32, IV 6-13 (6-segmented antennae) or III 28-34 (5-segmented antennae). On
Polygonum thunbergii and Polygonum sp. in southern Japan,
Taiwan, China, Sri Lanka (BMNH collection, leg. Lund University Ceylon Expedition
1962), and alatae have been trapped in New South Wales, Australia (Carver
1976).
A
little-known genus with eight nominal species in eastern Asia. Molecular data
has cast some doubt about whether it belongs in Cerataphidini (Stern et
al. 1997). It has aleyrodiform apterae with a complete crenulate margin
of wax glands, as in Cerataphis, but
distinguished from that genus by absence of frontal horns and a more evident
division between prothorax and mesothorax.
The biology is still very unclear. Host-alternation from Styrax was proposed, but the species
involved, takenouchii Takahashi, is
no longer thought to belong to this genus, and is now placed in Tuberaphis. One species of Aleurodaphis has now been found to
produce leaf-roll galls on Stuartia (Theaceae) in Japan, and another
was obtained from boat-shaped galls on Sinojackia
xylocarpa (Styracaceae) in China.
Others are described from Compositae/Asteraceae (three species), Bambusa, Impatiens and Ligularia. Two of the
species on Compositae/Asteraceae in Japan are monoecious and anholocyclic
(Takahashi & Sorin 1958). The Japanese species were reviewed by Sorin
& Miyazaki (2004), and the Chinese species by Jiang & Qiao (2011) and
Qiao et al. (2018). Aleurodaphis
antennata Chakrabarti & Maity Only the aleyrodiform aptera (BL 1.3-1.5
mm) is described, from Bambusa sp.
in Uttar Pradesh, India (Chakrabarti & Maity 1982). The last rostral segment (RIV+V) is shorter
than in other Aleurodaphis (1.1-1.5
× HT II). Appearance in life is
unknown, presumably dark with marginal wax. Aleurodaphis asteris Takahashi & Sorin Plate 3a Apterae are black, aleyrodiform, with a fringe of wax; BL 1.0-1.1 mm. On Aster spp. in Japan and Korea, living all year on stems at or just below ground level, but young larvae move to distal parts of stems in spring and summer. Also recorded from China, where it was also found on Carpesium abrotanoides (G. Zhang 1999). Anholocyclic, with seven apparently non-overlapping generations per year on Aster (= Gymnaster) savatieri (Takahashi & Sorin 1958). Alatae are unknown. 2n= c.32. Aleurodaphis blumeae van der Goot (= A. sinisalicis Zhang; synonymy by Jiang & Qiao 2011) Apterae are pinkish purple to red-black, not as black as A. asteris, with glassy marginal rays of wax, and developing distinctive white spinal wax markings (Moritsu 1983); BL 0.9-1.4 mm. In dense colonies on stems, and also on undersides of young leaves of hosts, causing slight leaf-curl (Calilung 1967 and J. Martin, pers.comm.). Recorded from various Compositae/Asteraceae (Adenocaulon, Aster, Blumea, Carpesium, Chrysanthemum, Cynoglossum, Erechtites, Kalimeris), and from Mazus miquelii (BMNH collection, leg. W. H. Paik). A record from Salix (as A. sinisalicis Zhang) is possibly erroneous (Jiang & Qiao 2011). In east and south-east Asia (China, Japan, Korea, Vietnam, Taiwan, Assam, Philippines, Java, Malaysia). Aleurodaphis
impatientis Sorin & Miyazaki Apterae are blackish brown to dark
purple brown with marginal white wax; BL 1.2-1.3 mm. On leaves and stems of Impatiens spp. in Japan. Anholocyclic,
overwintering at stem bases (Sorin & Miyazaki 2004). 2n= c.30*. Aleurodaphis ligulariae Sorin & Miyazaki Apterae are blackish brown to dark purple brown with marginal white wax; BL 1.9-2.1 mm. On undersides of leaves, stem apices and flowerstalks of Ligularia fischeri in Japan. Anholocyclic, overwintering on basal parts of Ligularia and in leaf litter (Sorin & Miyazaki 2004). Aleurodaphis mikaniae Takahashi Apterae are reddish brown, with a fringe of wax; BL 1.3-1.6 mm. In dense colonies on stems of Mikania scandens in Taiwan, and there are subsequent records including one from a Parasenecio sp. in China (Shaanxi, Sichuan; Huang et al. 2012). 2n= c.30. Aleurodaphis
sinojackiae Jiang & Qiao Apterae are aleyrodiform, with marginal
wax: BL 1.8-2.1 mm. On Sinojackia
xylocarpa, inducing the leaves
to curl and form boat-shaped galls (Jiang & Qiao 2011). Alatae were
collected in June. In Zhejiang and Jiangsu provinces, China. The life cycle
is unknown. Aleurodaphis
stewartiae Sorin & Miyazaki Colour in life unrecorded, presumably
rather dark; BL of aptera (progeny of fundatrix?) about 1.5 mm. On Stuartia
monadelpha in Japan (Honshű), rolling the leaf margin upwards to form a
crescent-shaped gall (Sorin & Miyazaki 2004). Emigrant alatae emerge in
late July/early August; two were collected on an Aster sp., which
could possibly be the secondary host.
One east Asian species closely related to Aphis but with differences in chaetotaxy of dorsal body, R IV+V and second tarsal segments, and in the position of the marginal tubercles on ABD TERG 7 in fundatrices (Takahashi 1966). Alatae have dark-bordered wing veins. Aleurosiphon smilacifoliae (Takahashi) Apterae are dark green, thickly covered with wax powder; BL 1.4-2.1 mm. On Smilax spp., feeding along main veins on underside of leaves. Japan, Korea, China and Taiwan. Monoecious holocyclic with oviparae and alate males in October (Takahashi 1923). 2n=8.
About 12 species with narrowly
elliptical body-shape, differing from other Saltusaphidinae in having
setiform empodial hairs. Previously these species were treated as a subgenus
(Trichocallis) of Thripsaphis, but Quednau (2010)
restored its full generic status, establishing the correct type species as Allaphis caricicola Mordvilko (= Thripsaphis
producta Gillette). Allaphis californica (Hille Ris Lambers) (Fig.19j) Apterae are very pale, rather shiny
yellowish white with glassy transparent legs and sides of body, but black
antennae; BL 2.2-2.7 mm. Alatae are unknown. On Carex ?comosa in
California, USA, and also collected on C. amplifolia in Washington
(BMNH collection, leg. D. Carroll).
2n=10. Allaphis cyperi (Walker) (Fig. 19l) Apterae are described as yellowish green,
covered with bluish white wax, with dark appendages, and extensive greyish
dorsal sclerotisation leaving a pale spinal stripe (but see also aphidtrek.org); BL 2.3-3.1 mm.
Apterae have 0-5 secondary rhinaria on ANT III; alatae have 8-13 on III and
0-3 on IV, and dorsal abdomen with broad dark bars, usually separated between
tergites. On leaves of Carex spp., aften on river banks.
Widely distributed in Europe and North America, and also recorded from
Kazakhstan (Kadyrbekov 2014f) and east and west Siberia (Quednau &
Shaposhnikov 1988; Stekolshchikov & Khureva 2020). Oviparae and apterous
males occur in September (UK, BMNH collection) to late October (Spain, Nieto
Nafría & Mier Durante 1998) in Europe, but an oviparae collected in early
July in Wisconsin USA was depicted by Quednau (2010). A darker form with
shorter appendages and a subarctic distribution in Greenland, Iceland,
northern Scandinavia and northern Russia is now regarded (Quednau 2010) as a
subspecies, A. cyperi ssp. amurensis
(Mordvilko) (= Thripsaphis vibei in
Blackman & Eastop 2006), with Thripsaphis
vibei ssp. arctica Hille Ris
Lambers now regarded as a seasonal
variant of this subspecies). Oviparae and apterous males of this subspecies
were collected in late July in Greenland, and in early August in Iceland
(Hille Ris Lambers 1960b, as Thripsaphis
vibei ssp. arctica). However Stekolshchikov (2015) has reported
the nominal subspecies (A. cyperi
cyperi) as occurring in north-east Siberia (Chukotka), and Stekolshchikov
(2017) reported that specimens from the Nenets region in north-west Russia
had characters of both subspecies.. Allaphis daviaulti (Quednau) (Fig.19g) Apterae are very elongate, pale whitish
yellow, with pale legs, and antennae black distally; BL 1.9-2.3 mm. Alatae
are unknown. In leaf sheaths of Carex ssp.
(trichocarpa, haydenii) in eastern Canada (Quebec, Ontario). The sexual morphs
are undescribed; the record of an ovipara by Blackman & Eastop (2006) is
unsubstantiated. Allaphis foxtonensis (Cottier) Apterae are pale green to yellowish
green; BL 2.4-2.7 mm. Alatae have 12-17 rhinaria on ANT III, and dark
transverse bars on dorsal abdomen (cf. T.
producta). On Carex spp. (ternaria, gaudichaudiana), especially colonising bases of young leaves in
centre of plant. In New Zealand and
Australia (NSW), but without related species in the southern hemisphere and
almost certainly an introduced species. It appears closely related to A. producta; the differences were
discussed by Hille Ris Lambers (1974). 2n=10*. Allaphis hybrida (Hille Ris
Lambers) Colour of apterae in life
is unknown; BL 2.2-2.5 mm. On Carex sp.
in California, USA. The possibility that this population was a hybrid between
A. ossiannilssoni and A. verrucosa was discussed in the
description of this species (Hille Ris Lambers 1974b), but it was regarded as
a good species by Quednau (2010). Allaphis ossiannilssoni (Hille Ris
Lambers) (Fig.19i) Apterae are very elongate, pale greyish
yellow to greyish brown with darker marginal sclerites, and a dark grey
eighth abdominal tergite, secreting bluish wax wool especially at sides and
end of abdomen; BL c.1.7-1.9 mm. Alatae have 8-13 secondary rhinaria on ANT
III and 1-3 on IV. On Carex spp.,
widely distributed in Europe and across Asia to Japan (Higuchi 1972) and
China (as Thripsaphis ossiannilssoni ssp.
hebeiensis Zhang & Zhang, synonymised with the nominate species by
Quednau 2010; the host in China is recorded as Cyperus hebeiensis, but this is not a recognised plant name). Allaphis pacifica (Hille Ris Lambers) Similar to A. ossiannilssoni, and described as a subspecies of that species,
but raised to specific rank by Quednau (2010). Alatae are unknown. On Carex sp. in California, USA. Allaphis producta (Gillette) (= caricicola Mordvilko 1921, = cyperi
ssp. wulingshanensis Zhang & Zhang 1993; synonymy by Quednau
2010) (Fig.19e,h) Apterae are very elongate, yellowish
white to pale greyish brown, secreting bluish white wax, especially at end of
abdomen; BL 1.8-2.5 mm. Alatae have 10-14 rhinaria on ANT III, and broad
black cross bars that are fused into a solid patch on abdominal tergites
3-6. On leaves of Carex
spp. in North America, Europe, and across Asia to east Siberia ; it is
now considered that palaearctic populations formerly known as Thripsaphis caricis (Mordvilko) are
this species. There are oviparae in BMNH collected in mid-July in Iceland,
and in mid-August to September in UK, and oviparae and apterous males were
found in Spain in mid-October (Nieto Nafría & Mier Durante 1998). Allaphis scabra (Hille Ris
Lambers) (Fig.19m) Colour of apterae in life is unknown; BL
2.4-2.8 mm. Alatae have 9-11 secondary rhinaria on ANT III. On Carex sp. in western USA. Allaphis stromi Quednau Colour of apterae in
life is unknown; BL 1.5-2.6 mm. Alatae
are unknown. On Carex spp. in
central USA (Quednau 2010). Allaphis utahensis (Knowlton & Hall) (Fig.19k) Colour in life is unknown, BL of apt. 1.3(?)-2.6 mm. An alata has c.9 secondary rhinaria on ANT
III and broad dark cross bands on the dorsal abdomen. On Carex nebraskensis in western USA (Idaho, Utah). Oviparae and
apterous males occur in October in Utah. Allaphis verrucosa (Gillette)
(Fig.19n) Apterae are yellow or pale greenish
yellow, with antennae distally black and legs mainly pale; BL 2.4-2.8
mm. Alatae have 9-20 sec. rhin. on ANT
III, and dorsal abdomen with broad dark bars partially separated between
tergites. On Carex spp. in Europe, west Siberia, and widely distributed in
North America. Hille Ris Lambers (1974) distinguished populations in
Labrador/Alaska and south-west California as verrucosa ssp. nodulosa and verrucosa ssp. subverrucosa respectively, and these subspecies
are retained by Quednau (2010), although it still needs to be demonstrated
that the described differences are not environmentally induced. Sexuales
occur in late August-September in Sweden (Ossiannilsson 1959). 2n=10.
A monotypic genus for a hairy
aphid with elongate siphunculi and a short cauda. Allocotaphis
quaestionis (Börner) Plate 10d Apterae are rather large, green with a waxy
bloom, rolling apple leaves (Malus spp.)
in spring. The second generation are all alatae; green, with rust-brown
thorax, a black dorsal abdominal patch, and very numerous tuberculate
secondary rhinaria on ANT III-V. They
migrate for the summer to Senecio
doronicum. The gynoparae returning to apple initially give rise to
single-parent colonies on the undersides of the leaves (Hille Ris Lambers
& Wildbolz 1958). In Austria, north-west Russia, Iran and Kazakhstan. However, it may also be in China, as a gynopara
probably of this species was recorded by Zhang (1999) from Pyrus xerophila, and described as a “sp. dubia” under the name A. minensis Zhang, Chen, Zhong &
Li. A record from Tunisia (Ben
Halima-Karmel 2012) also needs further confirmation.
The apterae of the only known
species have long marginal prosomal hairs like Thoracaphis but lack siphuncular pores and marginal hairs on the
reduced ABD TERG 2-7. They have a less
sclerotic dorsum than Thoracaphis,
with an irregular mosaic-like ornamentation. Allothoracaphis
piyananensis Takahashi Apterae are pale yellow to
yellowish-brown, flattened dorsoventrally, on the undersides of leaves along
the main veins. Recorded from Quercus glauca and Q. morii in Japan and Taiwan
(Takahashi 1958a). The alate morph is
unknown, and the species is apparently anholocyclic.
A
genus for three Fagaceae-feeding species, narrow-bodied and with long
siphunculi like many Eutrichosiphum,
but differing in the presence of distinctly capitate or spatulate hairs. Qiao
et al. (2006a) reviewed the genus. Allotrichosiphum
assamense Raychaudhuri, Ghosh, Banerjee & Ghosh Apterae about 1.6 mm, appearance in life
unrecorded but probably green or pale brown; known only from one apterous
vivipara and one immature collected on Quercus
dealbata (= Q. floribunda? or Lithocarpus dealbatus?) in Assam,
India (Raychaudhuri et al.
1973). (Note that this is a different
species from Eutrichosiphum assamense). Allotrichosiphum
cyclobalanopsidis Qiao, Jiang & Martin Appearance in life unrecorded, BL of
aptera 1.9-2.3 mm. Collected in December on Quercus (Cyclobalanopsis)
spp. (neglecta, bella)
in China (Hong Kong, Hainan, Yunnan). The description (Qiao et al.
2006a) includes an alata and first and
fourth instar nymphs. Allotrichosiphum
kashicola (Kurisaki)
Apterae (fundatrices) 2.3-2.8 mm, colour in life unrecorded, probably
green or pale brown. Alatae dark
greenish-brown, clear-winged, with siphunculi nearly as long as body. On young foliage of Quercus (Cyclobalanopsis) spp. (acuta, glauca, myrsinifolia) in Honshu and Kyushu,
Japan. The life cycle is greatly
abbreviated, with sexual morphs (greenish alate oviparae and yellow alate
males) appearing in May. The rather
flat eggs are laid on the undersides of leaves. At Osaka there are only two generations per
year (fundatrices and sexuales), although at Tokyo additional parthenogenetic
generations have been observed (Takahashi & Sorin 1957, 1959). Apterous viviparae other than fundatrices
have not been detected.
A
genus with only one known species, probably of African origin. Absence
of siphunculi, short, 5‑segmented antennae, elongate rostrum and
characteristic habitat and appearance in life make confusion with other
aphids unlikely. Hille Ris Lambers
(1954a) redescribed Aloephagus and
discussed its taxonomic position. Aloephagus
myersi Essig Plate 1e, fig. 81F Apterae
are pale or variably pigmnted and secrete a flocculent wax; BL 1.8‑2.5
mm. On Aloe spp., living under the
leaf bases, attended by ants, and also reported from the related genera Haworthia and Astroloba. Photos of live
aphids and damage to Aloe can be
found at:- http://www.forestryimages.org/browse/TaxThumb.cfm?fam=30&genus=Aloephagus. Widespread in Africa, especially south of the
Sahara, where it is probably native, but originally described from
California. It is also found in glasshouses in Europe, has been found on A. arborescens in Japan (Sano &
Matsumoto 2005), and is reported to be a major pest of cultivated aloes in
Australia (Forster 1998). It is presumed to have a sexual phase on Pistacia in Africa, as emigrant
alatae suspected to be this species, BL about 1.8 mm, have been collected
from galls on a Pistacia sp.
(presumably P. aethiopica) in Kenya
(D. Hille Ris Lambers, pers. comm.; specimens in BMNH collection). Anholocyclic on Aloe in Europe and North America (Blackman & Eastop 2000,
p.220). The galls and the emigrant
alatae are as yet undescribed. 2n=22.
One species on Lonicera in North America related to Hyadaphis but with thinner siphunculi and hind wings of alata having only one oblique vein. Aphids collected on Eupatorium sp., at ground level with ants, in Washington DC (BMNH collection, leg. A. Jensen), and on Symphoricarpos albus in Idaho (A. Jensen colln - aphidtrek.org) may also belong to this genus. Alphitoaphis
lonicericola (Williams) Apterae yellowish brown (Hottes 1926b) or
reddish purple (original description), dusted with wax powder; BL 1.6-2.1 mm.
On young growth of Lonicera spp.,
causing leaf-curl. In USA (Minnesota, Nebraska, Illinois, Kansas). Alatae
have very numerous secondary rhinaria, distributed III 35-47, IV 24-30, V
14-17 (Hottes, 1926b). Life cycle is unknown.
A
genus for one species on Berberis in
Central Asia, characterised by unusual features of the siphunculi and cauda. Amegosiphon platicaudum Narzukulov (Fig.17a,d) Apterae are rather elongate-bodied with
long appendages, pale green to yellowish with an irregular red spinal stripe;
BL 1.6-2.7 mm. On undersides of leaves of Berberis
spp. in Iran, Kazakhstan and Tajikistan. Oviparae and alatiform
(brachypterous) males occur in October-November (Remaudičre & Davatchi
1959, as Elbourzaphis behboudii).
Mehrparvar (2016) provided a redescription.
One palaearctic species on Umbelliferae/Apiaceae with well developed marginal tubercles on abdominal tergites 1-(3-) 5 and apterae with secondary rhinaria. Ammiaphis
sii (Koch) Plate 10e Apterae are pale green or yellow-green, lightly
wax-dusted, with black siphunculi; BL 1.9-2.1 mm. On Falcaria vulgaris, in summer colonies on stems and in
leaf-sheaths, causing swelling of sheaths and twisting and curling of leaf
blades, and in spring and autumn at bases of stems and on young shoots. Central and eastern Europe, south-west and
central Asia (Azerbaijan; Rusanova 1948, as Aphis falcarii; Kazakhstan).
Monoecious holocyclic according to Börner (1952).
Ten species in Asia and North America mostly associated with Lonicera, characterised by having secondary rhinaria in apterae and a very short broadly rounded cauda. Amphicercidus alticola (Narzikulov & Mukhamediev) Apterae are pale green with dark appendages; BL 2.3-2.6 mm. Alatae have 24-32 secondary rhinaria on ANT III. On Lonicera nummularifolia in Central Asia (Tajikistan). A population with apterae having more rhinaria on ANT III was described as a subspecies, A. alticola ssp. tschatcalica (Mukhamediev & Akhmedov 1982). Amphicercidus flocculosus (Gillette & Palmer) Plate 10i Apterae have a blackish brown thorax and abdomen, dull orange to clay-coloured head, sides of body and bases of siphunculi, and are wax-powdered; BL 2.1-2.8 mm. Alatae have secondary rhinaria distributed III c.40, IV 8-16. On leaves and twigs of Lonicera and Symphoricarpos in western USA and Mexico. Monoecious holocyclic, with oviparae and alate males in October-November (Palmer 1952). Amphicercidus forsythiae Zhang, Zhong & Zhang Apterae are greyish green with white powder; BL 3.1-3.2 mm. Alatae have broad dark dorsal cross-bands tending to merge across ABD TERG 4-6, and secondary rhinaria distributed III 113-116, IV 0-2. On Forsythia suspensa in Yunnan, China. Apart from the small number of secondary rhinaria on ANT III of apterae – a character which is likely to depend on the degree of alatiformity - this could be A. japonicus. Amphicercidus japonicus (Hori) Apterae are dark reddish to greenish or yellowish brown, with abundant white wax; BL 2.5-3.4 mm. Alatae have secondary rhinaria distributed III 80-152, IV 0-4. On young shoots and twigs of Lonicera spp. in east Asia (northern India, China, Korea, east Siberia, Japan). Monoecious holocyclic, with oviparae and alate males in October in Japan (Miyazaki 1971), but perhaps anholocyclic in India (Hille Ris Lambers & A.N. Basu 1966, as A. indicus). 2n=8 (Chen & Zhang 1985b). Amphicercidus laniger (Takahashi) Apterae are brownish green, densely covered in white wax powder; BL 1.9-2.5 mm. Alatae have 2-5 secondary rhinaria in the middle region of ANT III. On young stems of Stauntonia hebandra (= S. obovata) in Taiwan. The alata was described by Takahashi (1929). Amphicercidus lonicerae Maity & Chakrabarti Apterae are wax-covered; BL 3.2-3.6 mm. Alatae have secondary rhinaria distributed III 55-75, IV 4-6. In downwardly-rolled leaves of Lonicera quinquelocularis in north India. 2n=8? (Khuda-Bukhsh 1980, as Amphicercidus sp.; 2n=18 was recorded by Khuda-Bukhsh & Pal 1983b, but this was probably an error). Amphicercidus lonicericola (Mukhamediev) Apterae are pale green, wax-covered; BL 1.8-2.2 mm. On Lonicera altmannii in Central Asia. Amphicercidus pulverulens (Gillette) Apterae are dull greenish brown with abundant wax; BL 2.5-3.1 mm. Alatae have secondary rhinaria distributed III c.40, IV 0. On bark of Symphoricarpus spp., at or below ground level. In western USA and Canada. Monoecious holocyclic, with oviparae and apterous males in October-November (Palmer 1952). Amphicercidus sinilonicericola Zhang Colour of apterae in life is unrecorded, presumably wax-covered; BL c. 3.0 mm. On Lonicera japonica in China. Possibly a synonym of A. japonicus. Amphicercidus
tuberculatus David,
Narayanan & Rajasingh Apterae
are green, wax-covered; BL 2.1-2.6 mm. Alatae have
secondary rhinaria distributed III 50-63,
IV 0-3 (David et al. 1972). On Lonicera quinquelocularis in north
India. 2n=6 (3 samples; Chauhan & Kurl 1990) or 12 (Pal &
Khuda-Bukhsh 1984).
About 27 species, mostly nearctic, with a few east and west palaearctic. About half live on Rubus. A few have become fern feeders. They are all medium-sized to large aphids, usually greenish and rather pale, with long appendages bearing short to medium-length hairs. The siphunculi are long, pale or dusky and slightly to moderately swollen on distal half. Most species on Rubus occur in loose colonies on shoots or solitarily on leaves. Mostly monoecious holocyclic as far as known, with many alatae produced in the third parthenogenetic generation. Heie (1995) and Blackman (2010) reviewed the species of north-west Europe. The nearctic species need revision (see aphidtrek.org). The great variation in chromosome number is discussed by Blackman (1980). Amphorophora agathonica Hottes Apterae are pale green with siphunculi pale at base, becoming darker towards apices; BL 2.4-4.7 mm. On young stems and undersides of leaves of Rubus idaeus var. strigosus, and occasionally in small numbers on other Rubus spp. Widely distributed in North America north of about latitude 38˚N. See Kennedy & Schaefers (1974) for population studies. Monoecious holocyclic with alate males. 2n=14. Amphorophora ampullata Buckton (Fig.43e) Apterae are green, with dark-tipped siphunculi; BL 3.0-5.0 mm. On undersides of fronds of ferns in numerous genera (Aspidium, Asplenium, Athyrium, Cystopteris, Dryopteris, Matteuccia, Onoclea, Polypodium, Polystichum, Thelypteris). In Europe, Asia and North America. Monoecious holocyclic with alate males. Originally described from Cystopteris montana, the name is probably being applied to a complex of species with different fern associations. In USA, populations mainly associated with Onoclea sensibilis and Matteuccia spp. (Onocleae) are regarded as a subspecies, A. ampullata ssp. laingi Mason. Populations colonising various ferns in north-east India and Nepal also have distinctive features and have also been given subspecies status, as A. ampullata ssp. bengalensis Hille Ris Lambers & A.N. Basu, which form has now been recognised as occurring in China (Su et al. 2014). Populations on Athyrium felix-femina in Netherlands and UK have a longer R IV+V and other morphological differences as well as a different karyotype (see below), and thus represent an undescribed species. Clearly there is a need for biological, morphometric and molecular studies to clarify relationships within this group. 2n=12 for samples from Dryopteris and Polystichum in UK and Japan (RLB) for ssp. bengalensis from north-east India (Kurl & Chauhan 1986a), and for ssp. laingi from Canada; 2n=10 for a sample from Athyrium filix-femina in UK. Amphorophora amurensis (Mordvilko) Apterae are green with antennae, legs and siphunculi blackish, and a pale cauda; BL 3.0-3.5 mm. On Rubus spp. in east Asia. Takahashi (1961e) provided a description of this species as A. rubiphaga. 2n=14. Amphorophora (Galiaphis) annae (Ossiannilsson) (Fig.29f) Apterae are whitish yellow, pale yellowish green or pale green, with darker apices to antennal segments, legs and siphunculi; BL 1.8-2.3 mm. On stems and undersides of leaves of Galium boreale in northern Europe (Sweden, Finland, eastern Germany). Oviparae occur in August-September, apterous males in September (Ossiannilsson 1954). Amphorophora
catharinae (Nevsky) Apterae are green, eyes black, antennal
segments white with black apices, siphunculi pale with dusky apices; BL 2-3
mm. Alatae have 10-12 secondary rhinaria on ANT III. At shoot tips and on
undersides of leaves of Rosa spp.
in Uzbekistan, and also in Iran, Afghanistan, Kazakhstan (Kadyrbekov &
Aoitzhanova 2005), Tajikistan and Kashmir (BMNH collection). Oviparae and alate males were described from R. ×damascena in November (Narzikulov & Umarov 1969). Amphorophora coloutensis Smith & Knowlton Apterae are pale whitish with a greenish brown abdominal spinal stripe; BL 2.9-3.4 mm. On Geranium spp. (wild) in western USA (Colorado and Utah). Monoecious holocyclic, with oviparae and alate males in late July/August (original description). Amphorophora (Galiaphis) cryptotaeniae (Takahashi) Apterae are pale, siphunculi dark-tipped; BL 1.0-1.3 mm. On Cryptotaenia japonica in Japan. Its generic position is uncertain. Amphorophora filipendulae Miyazaki Apterae are green or greenish yellow, with mainly dark siphunculi; BL 2.5-3.9 mm. On undersides of leaves and young shoots of Filipendula camtschatica in Japan (Miyazaki 1971) and Kamtschatica (Pashchenko 1988a). Monoecious holocyclic. Amphorophora forbesi Richards (= Aulacorthum capilanoense Robinson) Apterae are pale green or whitish (see aphidtrek.org); BL 2.7-3.8 mm. On undersides of leaves of Rubus spectabilis in western North America (British Columbia, Washington, Oregon). This species has an unusual combination of characters (large size, spiculose ventral side to head, steep-sided slightly scabrous antennal tubercles, no rhinaria on ANT III, slightly swollen siphunculi and a rather hairy cauda), so its generic position is not at all clear. 2n=12. Amphorophora gei (Börner) Apterae are pale green or yellowish green, with dusky or dark siphunculi and cauda; BL 2.2-3.9 mm. On undersides of leaves of Geum spp., especially G. rivale in moist, shady places (Heie 1995). Also recorded from Alchemilla vulgaris and Filipendula ulmaria. Widespread in Europe, and introduced to western USA (Idaho: BMNH collection, leg. J. H. Martin). Monoecious holocyclic on Geum; Hille Ris Lambers (1947) described the fundatrix, but sexuales have apparently never been found. 2n=12. Amphorophora geranii Gillette & Palmer Apteraea are “dull greenish yellow to pinkish, mottled with sooty or grass-green”; BL 2.4-3.7 mm. Alatae were described in Palmer (1952). On leaves of Geranium richardsoni and G. bicknellii in western USA (Colorado, Idaho, Utah, Wyoming). Amphorophora idaei (Börner) Apterae are pale green, rather shiny, with pale siphunculi (see influentialpoints.com/Gallery); BL 2.6-4.1 mm. On undersides of leaves of Rubus idaeus s.str., and cultivated varieties derived from it. Europe, and also recorded from Central Asia (Kazakhstan; Kadyrbekov 2005c) and west Siberia (Stekolshchikov & Khureva 2020). Monoecious holocyclic, with alate males. Blackman et al. (1977) distinguished this species from A. rubi, under which name it was often previously included. McMenemy et al. (2009) reviewed its biology, and Mitchell et al. (2010) investigated biocontrol using a combination of plant resistance and the parasitoid Aphidius ervi. 2n=18. Amphorophora (Galiaphis) japonica (Takahashi) Apterae are green; BL c.1.5 mm. Host recorded as “Anemone (?)”. Japan. Host and generic position are uncertain. Amphorophora kesocqua Hottes Apterae are bluish green, siphunculi becoming darker distally; BL c.3 mm. On Rubus sp. in Colorado, USA. Possibly these are fundatrices of A. tigwatensa. Amphorophora pacifica Hill Apterae are shining very pale greenish white with a darker green rather wavy spinal stripe, somewhat interrupted posteriorly; BL 2.1-3.0 mm. On undersides of leaves of Rubus parviflorus in California, USA, where it is anholocyclic, overwintering on young foliage at apices of older shoots (original description). 2n=18. Amphorophora
parviflori Hill Apterae are pale green; BL 3.1-4.3 mm. On
Rubus parviflorus, and sometimes in
small numbers on certain other Rubus spp., in British Columbia, Canada. 2n=12. Amphorophora pawtincae Hottes Apterae are pea-green; BL 2.5-3.7 mm. On leaves, stems and flowers of Primula parryi in western USA (Colorado, Oregon) (Palmer 1952). Amphorophora rossi Hottes & Frison Apterae are whitish green with a darker green spinal stripe; BL 1.9-3.0 mm. On Geum spp., especially G. canadense, in north-east USA and eastern Canada, and also recorded from G. macrophyllum in western USA (Washington, Oregon; A. Jensen colln - aphidtrek.org). Monoecious holocyclic; the original description included the fundatrix. 2n=46*. Amphorophora rubi (Kaltenbach) Apterae are shiny pale yellowish green to green with pale siphunculi (see influentialpoints.com/Gallery); BL 2.2-4.7 mm. On undersides of leaves of Rubus spp., especially fruticosus s. lat., in Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), west Siberia (Stekolshchikov & Khureva 2020), south-west and central Asia, Mongolia (Kadyrbekov & Aoitzhanova 2005; Kadyrbekov 2017a), and introduced to Australia and New Zealand. Monoecious holocyclic with alate males, or anholocyclic in regions with mild winters. Records of A. rubi on raspberries mostly refer to A. idaei in Europe and to A. agathonica in North America. 2n=20. Amphorophora rubicumberlandi Knowlton & Allen Colour of apterae in life unrecorded; BL 2.6-3.7 mm. On canes of wild and cultivated black raspberries (Rubus occidentalis, R. leucodermis) in north-western USA (Oregon, Washington). Amphorophora rubitoxica Knowlton Plate 24g Apterae are shiny deep green, or yellow-green with a darker green spinal stripe, with siphunculi dusky-dark; BL 2.3-3.5 mm. On young shoots and under leaves of many Rubus spp., and causing lesions on leaves of R. occidentalis, but not colonising R. idaeus. Western North America. Life cycle has not been studied. 2n=30. Amphorophora scabripes Miyazaki (Fig.43d) Apterae are pale green with two longitudinal dark green pleural stripes, and black-tipped siphunculi; BL 4.8-5.7 mm. On Thelypteris (= Lastrea) quelpaertensis in Japan. Smaller yellow aphids (BL 3.6-4.8 mm) with dark hind femora on Matteuccia intermedia (= Pentarhizidium intermedium) in China were described as a subspecies, A. scabripes ssp. galba by Su et al. (2014), but are perhaps a separate taxon closer to the A. ampullata group (q.v.). Amphorophora sensoriata Mason Apterae are very pale bluish green with dark antennae, and siphunculi dusky towards apices; BL 2.8-3.5 mm. On stems of Rubus occidentalis, dropping off readily when disturbed. north-eastern North America, as far south and west as Kansas. 2n=72. Amphorophora stachyophila Hille Ris Lambers (Fig.50c) Apterae are rather shiny pale green, rarely greenish pink; BL 2.5-3.5 mm. On stems and undersides of leaves of Stachys spp, esp rigida, in western North America (California, Oregon, Washington, British Columbia). 2n=12. Amphorophora stolonis Robinson Apterae are pale green; BL 2.6-3.1 mm. On stems or at ends of runners of Rubus pubescens in Manitoba, Canada. Monoecious holocyclic with alate males (original description). 2n=48. Amphorophora tigwatensa Hottes Apterae are pinkish brown or greenish, with antennae, legs and siphunculi mainly dark brown, cauda paler; BL 2.8-4.1 mm. On undersides of leaves and young stems of Rubus procerus (Leonard 1974), and also on R. occidentalis (A.Jensen colln - aphidtrek.org). In western North America. 2n=40*. Amphorophora tuberculata Brown & Blackman (Fig.30d) Apterae are shining deep green; BL 2.2-3.4 mm. On leaves and flowerheads of Geranium macrorrhizum in south-east England and Bulgaria; specific to this host, and presumably indigenous to montane areas of continental Europe where this plant is native. Monoecious holocyclic, with both apterous and alate males (Brown & Blackman 1985). The low chromosome number has allowed detailed studies of this aphid’s spermatogenesis (Blackman 1985) and sex determination (Blackman & Hales 1986). 2n=4. Amphorophora urtica Essig (Fig.55a) Apterae are dark red to greyish; BL 3.2-3.6 mm. On undersides of larger leaves of Urtica spp. in western North America (California, Utah, British Columbia). Life cycle is unknown, but presumably anholocyclic in California, as apterous and alate viviparae were abundant in December-January (Essig 1942).
One
European species related to Amphorophora
but with ante- and post-siphuncular sclerites and a long, very hairy
rostrum. Amphorosiphon
pulmonariae (Börner) Plate 24h Apterae are pale to dark green with black
siphunculi and posterior dorsal abdominal markings; BL 2.5-3.0 mm. On
undersides of leaves and petioles of Pulmonaria
spp. in Europe (Netherlands, Germany, Bulgaria, Czech Republic, Hungary,
Ukraine). Monoecious holocyclic with apterous males (Hille Ris Lambers 1949).
A genus for two oriental species close to Aulacorthum, but with 4 hairs on the first tarsal segments. Anaulacorthum fagopyri A.K.Ghosh & Raychaudhuri (Fig.28a) Apterae are pale brown; BL 2.3-2.8 mm. On undersides of young and old leaves of Fagopyrum cymosum in West Bengal, India. Alatae are undescribed. Anaulacorthum
zhangjiajiense (Zhang) Apterae are
pale red with black dorsal markings; BL c.2.6 mm. On Sonchus oleraceus in Hunan Province, China. Described as a
subspecies of Neomyzus circumflexus,
and transferred to this genus by Eastop & Blackman (2005). Su & Qiao
(2010b) provided a redescription.
A
distinctive holarctic genus of uncertain taxonomic position, with about 24
species, many of which are poorly known.
Some have host alternation from Cornus
to roots of Poaceae (in one case, Onagraceae). Others are entirely subterranean on Poaceae
or Cyperaceae. Adult apterae of most species on grass roots are
medium‑sized greenish grey or grey in colour, with a sclerotic dorsal
abdominal plate. They have compound
eyes and are fairly mobile when disturbed.
Immatures are paler, white or cream in colour. The alate sexuparae,
distinguished by their dark posteriodorsal abdominal patch and the large
black pterostigmal spot on the forewing, are a common sight on Cornus leaves in the autumn, alighting
and depositing ant-attended clusters of small, yellow and/or brown sexual
morphs. The oviparae lay their eggs on
the bark of the trunk. Different
species utilise the native Cornus
in North America, Europe and eastern Asia.
Accounts are available for
central Europe (Zwölfer 1958), north-west Europe (Heie 1980b), the UK
(Blackman et al. 2019b), the
Iberian peninsula (Nieto Nafría & Mier Durante 1998), India (Chakrabarti et al. 1982, A.K. Ghosh 1988), Japan
(Sorin 1999a), China (J. Yang et al.
2008) and North American Rocky Mountain region (Palmer 1952). Halbert (1991)
keyed the North American species. This is a difficult group in
need of further revision. In the BMNH collection there are undescribed
species from Cornus mas in Italy,
and from Cornus spp. in Korea. Aclitus
obscuripennis Foerster may be a specialised aphidiine parasitoid. See
also Blackman & Eastop 2000 pp. 222-3. Anoecia
caricis Pergande Colour of aptera not noted; BL c. 1.4 mm.
On roots of Carex sp. in USA. Anoecia
corni (Fabricius)
Plate 5a, b Host-alternating between Cornus sanguinea and roots of Poaceae
in Europe and North America; anholocyclic on roots of numerous species of
grasses and some cereals in Europe, Asia, Africa, Argentina (Ortego 1998a)
and North America. Apterae on grass roots are greenish grey to brown
with sclerotized parts dark grey; BL 1.9-2.8 mm. Apterae in spring on Cornus are dark brown; fundatrices
have reduced, 3-facetted eyes and 5-segmented antennae, whereas subsequent
generations have large compound eyes and 6-segmented antennae. Both spring migrant alatae and the
sexuparae returning to Cornus in
autumn have a dark patch covering ABD TERG 3-6. BL of alata 1.9-3.0 mm. Wieczorek (2008a)
described the reproductive system of the dwarf male, and Wieczorek & Świątek
(2009) compared it with that of Glyphina
betulae. See Knechtel & Manolache (1943)
and Heie (1980) for further information. 2n = 6. An excellent photo of a
sexupara of A. corni can be seen
at: http://www.flickr.com/photos/dioctria/3929122241/,
and see also influentialpoints.com/Gallery Anoecia
cornicola (Walsh)
Host-alternating between Cornus
spp. (amomum, stolonifera) and roots of Poaceae in North America, and also
recorded from Brazil (BMNH collection) where it is presumably anholocyclic on
grass roots. Apterae
on grass roots (Andropogon, Digitaria, Eleusine, Muehlenbergia,
Panicum, Poa, Setaria) are
whitish or yellowish with cross-bands, wax-powdered; BL c. 1.7 mm. Alate including sexuparae
produced on grasses in August-October have a dark dorsal abdominal patch, but
this is apparently not present in spring migrants (Palmer 1952, as A. querci). BL of alata 2.0-2.5 mm. 2n=10. Anoecia
cornimaris Bozhko Apterae are pale brown; BL c. 2.4 mm. On Cornus mas and C. sanguinea in eastern Europe (Moldova, Ukraine). Life cycle is
unknown. Anoecia
equiseti Halbert Apterae are cream-coloured with grey
sclerotic areas; BL 2.1-2.4 mm. Late summer alatae commonly have vestigeal
wings. In ant-attended colonies on roots of Equisetum laevigatum in Idaho, USA, and Manitoba, Canada (BMNH
collection, leg. A. G. Robinson). Life cycle is uncertain, monoecious
holocyclic or anholocyclic; oviparae were produced in late August, and eggs
laid but no males were observed and eggs may have been inviable (Halbert
1991) . Anoecia
fulviabdominalis (Sasaki) Host-alternating between Cornus spp. (brachypoda, controversa)
and roots of Poaceae in Japan; also anholocyclic populations occur more
widely on roots of grasses, and it is a major pest of upland rice in east and
south-east Asia. Apterae on grass roots are grey‑green; BL
2.1-2.5 mm. Apterae
in spring on Cornus blackish, BL about 2 mm. Both spring migrant
alatae and the returning sexuparae have dark dorsal abdominal patches. Tanaka (1961) gave
an account (in Japanese) of its biology in Japan, summarized in Yano et al. (1983). Sorin (1999a) provided descriptions of the
morphs produced on Cornus, and described fundatrices and emigrant
alatae of a variant form as a subspecies, A. fulviabdominalis similis. Anoecia furcata (Theobald)
(= A. nemoralis Börner) Apterae are dark green to blackish brown;
BL 1.4-2.4 mm. Alatae have a black dorsal abdominal patch. On roots of
numerous grasses (Alopecurus, Arrhenatherum, Bromus, Deschampsia, Elymus, Festuca, Panicum, Phleum, Poa, Sieglingia),
attended by ants. In Europe, Central Asia, India, China. Mainly anholocyclic
on grass roots; part of the population may go through a sexual phase on Cornus sanguinea, but this has not
been clearly established. Zwölfer (1957) and Paul (1977) studied variation in
frequency and distribution of spatulate hairs. 2n=12. Anoecia graminis Gillette
& Palmer Apterae are dull black
with dusky bands, wax-dusted; BL 1.6-2.5 mm. Alatae have no black dorsal
abdominal patch. On roots of Hordeum sp.
in Colorado, USA. Apparently monoecious holocyclic, with oviparae in
September (Palmer 1952). Closely related to the European A. furcata. 2n=8. Anoecia haupti Börner Apterae on grass roots are greenish to
brown with sclerotic parts dark grey; BL 1.9-2.6 mm. Alatae have a dark
dorsal abdominal patch; BL 1.4-2.2 mm. On roots of grasses (Bromus, Eragrostis, Hordeum, Poa, Setaria) in Europe (Germany, Spain, Portugal, Italy, Turkey), and
there is also a record from an undetermined grass species in China (Inner
Mongolia; J. Yang et al. 2008).
Heteroecious holocyclic with a sexual phase on Cornus sanguinea (Zwölfer 1957), but the Cornus-feeding
morphs and sexuparae are undescribed. 2n=8. Anoecia
himalayensis Chakrabarti & Maity Colour
of apterae in life unrecorded, probably greenish or brown with grey sclerotic
parts; BL 1.8-2.0 mm. On roots of unidentified grass species in northern
India (Himachal Pradesh, Uttar Pradesh). Spring
colonies comprising apterous fundatrigeniae and immature spring migrant
alatae, in crinkled and crumpled leaves of Cornus macrophylla at 2550m
altitude in Pakistan, were provisionally assigned to this species by
Naumann-Etienne & Remaudičre (1995). Anoecia
ilicicola Sorin
Alatae described under this name from Ilex nipponica in Japan by Sorin (1999) are presumably vagrant
alate sexuparae from populations on grass roots. Alatae identified as the
same species have been collected in China on Cornus (=Bothrocaryum) controversa (J. Yang et al. 2008), as well as being
recorded as presumed vagrants on other plants (Artemisia, Kalopanax). Anoecia
japonica Sorin
Fundatrix, spring migrant alata, sexupara and ovipara are described
from Cornus controversa and Cornus sp. in Hokkaido, Japan
(Sorin 1999a). Sexuparae, presumably vagrants, were also collected on Pteryocarya
sp. Both spring migrants and sexuparae have a black dorsal abdominal patch.
Appearance in life and populations on secondary hosts (presumably grasses)
are unrecorded. Anoecia krizusi (Börner) Apterae are blue-green with dark grey
dorsal sclerotisation; BL 2.1-2.5 mm. Alatae are unknown. On roots of Elymus (Agropyron) caninum,
with tentative identifications also from Agrostis
sp., Dactylis glomerata, Lolium perenne and Sorghum bicolor. In Germany, ?UK
(Stroyan 1964), and also recorded from China (J. Yang et al. 2008). Monoecious holocyclic on grass roots (Zwölfer 1957,
as Neanoecia). It seems closely
related to A. haupti. Anoecia major Börner Apterae are brown with extensive dark
grey sclerotisation; BL 2.2-3.0 mm. Alatae (BL 2.3-3.2 mm) have a large black
dorsal abdominal patch. On roots of grasses, especially Phalaris arundinacea (also Brachypodium
pinnatum, Calamagrostis).
Europe (UK, Sweden, Netherlands, Germany, Spain). Probably with a sexual
phase on Cornus; not yet
found on Cornus in nature, but
Börner (1950) transferred sexuparae to C.
sanguinea and obtained 2-3 generations of fundatrigeniae. Very close to A. corni and hybrids may occur. 2n=8 (2n=7 in possible hybrids
with corni; Blackman 1980). Anoecia mirae Narzikulov Apterae are brown, with broad dark
abdominal cross-bands; BL c. 2.7 mm. On Sorghum
halepense in southern Tajikistan. Probably a synonym of A. major. Anoecia
oenotherae Wilson Apterae are yellowish, with dusky
cross-bands; BL c.1.5 mm. On roots of Oenothera
biennis, apparently widely distributed in North America. Alate
sexuparae produced on Oenothera roots
in autumn are small (1.4-1.8 mm), with 0-4 secondary rhinaria on III, and may
have dark cross-bands on ABD TERG 3-7, but never a solid black patch.
Migration occurs to Cornus spp. (amomum, paniculata, stolonifera).
Records of other Anoecia spp. from Oenothera
roots may be due to misidentification. See also
Palmer (1952). Anoecia
(Paranoecia) pskovica Mordvilko Apterae are greyish white; BL 2.4-2.7 mm.
Alatae are unknown. On roots and subterranean runners of Cyperaceae (Carex, Eriophorum). In Europe (UK, Denmark, Sweden, Germany, northern
Russia) and Central Asia (Kazakhstan; Kadyrbekov & Aoitzhanova 2005) .
Monoecious holocyclic with oviparae and apterous males from mid-October.
Usually ant-attended, but eggs are covered in protective wax and not
collected and stored by ants (Zwölfer 1957). Anoecia
radiciphaga Pal &
Raychaudhuri Colour of apterae in
life is unrecorded; BL 1.7-2.1 mm. On roots of Eragrostis nigra and unidentified grass species in north-east
India (West Bengal), and also recorded from various grasses and cereals
including Alopecurus aequalis, Avena fatua, Imperata koenigii (= cylindrica) in China (J. Yang et al. 2008). Anoecia
setariae Gillette & Palmer Apterae are pale orange-yellow with
brownish head and dusky cross-band on posterior abdomen; BL 1.6-2.0 mm. Alatae commonly have vestigeal wings. In
ant-attended colonies on roots of
grasses (Echinochloa, Setaria), apparently widely distributed
in North America (Smith & Parron 1978). Probably monoecious holocyclic;
ants were observed to gather eggs in September (Halbert 1991). Anoecia
similiradiciphaga Qiao &
Jiang Apterae are presumably dark
brown, BL 1.5-2.0 mm. On roots of Triticum
aestivum in China (J. Yang et al. 2008), and apterae and alatae are also recorded from (roots of?) Cyanotis vaga and Spiraea salicifolia, which are
unlikely hosts. Very similar to A.
fulviabdominalis, the differences requiring further confirmation. Anoecia
stipae Mamontova Apterae are pale brown with blackish
head, legs and tips of antennae, and dark cross-bands on abdiominal tergites
7 and 8; BL c. 2 mm. On roots of Stipa
capillata in chalky ground in Ukraine. Anoecia
takahashii Sorin
Apterae are presumably blackish brown, BL about 1.8 mm. Spring migrant
alatae have a black dorsal abdominal patch (Sorin 1999a; the illustration is
stated to be of the sexupara, but presumably this is in error, as that morph
is not described) . On Cornus sp. in Japan. Anoecia
tanakai Sorin Only
the sexupara is described, BL 2.1-2.5 mm, with an extensive black dorsal
abdominal patch. In autumn on Cornus spp. in Japan (Sorin 1999a). Sexuparae, presumably vagrants,
were also collected on Pterocarya rhoifolia. Anoecia
vagans Koch (= Anoecia willcocksi Theobald; Zwolfer
1957) Host-alternating between Cornus sanguinea and roots of Poaceae.
Fundatrices on Cornus are blackish;
BL c.5.0-5.8 mm (Depa & Mróz 2014). All their progeny are spring migrant
alatae (BL 2.3-2.8 mm) which, unlike those of A. corni, have no dark posterior abdominal patch. Apterae
on grass roots are yellowish, pale green or olive green; BL 2.1-2.7 mm. They
feed on roots of numerous grass and cereal species (Aegilops, Agropyron, Arrhenatherum,
Bromus, Cynodon, Elymus, Eragrostis, Festuca, Hordeum, Lolium, Panicum, Poa, Triticum). Alatae including sexuparae
produced on grass roots have a black dorsal abdominal patch (BL of alate virginoparae
2.1-2.5 mm, of sexuparae 2.5-3.9 mm; Heie 1980b). In Europe, Tunisia, Egypt,
Israel, Iran, Turkey, India, east Siberia, and alatae apparently of this
species have been trapped in USA (South Carolina, Florida; Skvarla et al. 2017). The male genitalia
were described and illustrated by Wieczorek et al. (2012). 2n=12. Anoecia
zirnitsi Mordvilko Apterae are greyish green or brownish
green; BL 1.5-2.1 mm. Alatae are unknown. On thin roots of various grasses (Agrostis, Brachypodium, Calamagrostis, Festuca, Lolium). In Europe (UK, Denmark, Sweden, Finland, Germany,
Poland, Latvia, west Russia), the Middle East and Central Asia (Kazakhstan;
Kadyrbekov 2017a). Monoecious holocyclic with oviparae in October-December,
eggs being stored by Lasius flavus
(Zwölfer 1957). Males unknown; Zwölfer’s suggestion that oviparae may be
parthenogenetic requires further investigation.
A
genus for two Australian species living on Myrtaceae (one on Agonis and the other on Leptospermum). The antennae of both apterae and alatae are
5-segmented, and abdominal tergites 7 and 8 each bear slender hair-bearing
processes. The siphunculi are
cylindrical or tapering, with a ring of 3-8 hairs on the distal half, and the
cauda is broadly rounded. Anomalaphis
casimiri Carver
Apterae are small (BL 1.3-1.7 mm) and brown. Distinguished from the only other described
species by the presence of polygonal reticulation on the siphunculi distal to
the ring of hairs, and by dorsal hairs of aptera mostly arising from
tubercles. Apterae and alatae
occurring together with intermediate forms on Leptospermum scoparium
var. Red Damask in New South Wales, Australia (Carver 1971). Appearance of colonies in life and biology
are not recorded. Anomalaphis
comperei Pergande ex Baker Plate 5f Apterae are small, dark brownish or greyish green with a yellowish
spinal stripe; BL 1.2-1.6 mm. Alatae have shiny black thoracic lobes and a
black dorsal abdominal patch. In
Western Australia on Agonis flexuosa;
this may be the only true host, although the original records of this aphid
were from Acacia and Eucalyptus. Monoecious holocyclic, with oviparae in
February (Carver 1971).
Nine
species associated with woody, mostly shrubby Connaraceae, Leguminosae and Xanthophyllaceae
in east and south-east Asia. The hosts
of two species are unknown. The
apterae have 4-segmented antennae, bear slender hair-bearing processes on
abdominal tergites 7 and 8, and have a cauda tipped with a stylus. Alatae have 5-segmented antennae, much
smaller processes on the posterior tergites, and the caudal process often
indistinct. Both oviparae and males
are alate in the one species (A.
mendeli Quednau & Martin) of which the sexuales are known, obtained
by canopy fogging from an unknown host. The male genitalia of this species
were described and illustrated by Wieczorek et al. (2012). Agarwala & Martin (1994) revised the genus,
and Quednau & Martin (2006) provided keys to apterae and alatae. Anomalosiphum
indigoferae A.K. Ghosh, M.R. Ghosh &
Raychaudhuri Apterae in life are dark
brown to black, BL 1.1-1.4 mm. Alatae
are larger, BL 1.4-1.7 mm. Described
from young twigs of Indigofera sp.
and Mimosa sp. in India (A.K. Ghosh et al. 1971e). There
are also records from Vigna sp. and
Phyllanthus sp.; the latter record
is of alatae only, probably vagrants. Recorded fom
Sikkim and West Bengal. The life cycle
is unknown. 2n=18. Anomalosiphum
murphyi Martin & Agarwala Plate 11a, b Colour in life unrecorded, apterae probably
dark-pigmented; BL 1.2-1.6 mm. On Rourea
sp. and Xanthophyllum stipitatum in
Singapore, and on ?Sindora sp. in
Sarawak. Alatae were collected in
Singapore in January. The life cycle
is unknown (Martin & Agarwala 1994). Anomalosiphum
philippinense Martin & Agarwala Colour of aptera in life is unknown,
probably dark; BL about 1.3-1.4 mm. Alatae are larger, 1.5-1.8 mm. On Dalbergia
ferruginea (= stipulacea)
in the Philippines, and alatae have been trapped in Papua New
Guinea (Martin & Agarwala 1994). Apterae and alatae were collected in
February-April. The life cycle is unknown. Anomalosiphum
pithecolobii Takahashi Apterae are probably green in life; BL
1.3-1.4 mm. Alatae are greenish yellow. Described from alatae collected on
young leaves and shoots of Pithecellobium
(= Archidendron) lucidum in Taiwan. Subsequently apterae believed to be the
same species were described from Canton on Albizia lebbek, although
the alatae from Albizia showed some
differences from the Taiwanese specimens (Takahashi 1936a). Specimens previously thought to be this
species from the Philippines and Papua New Guinea were distinguished as A.
philippense (Martin &
Agarwala, 1994) Anomalosiphum
scleroticum Qiao & Zhang Only the alata is described, BL about 1.6
mm, collected in April from Dalbergia hupeana in southern China
(Guanxi) (Qiao & Zhang 2001a). Anomalosiphum
takahashii Tao
Apterae are pale yellow; BL 1.2-1.5 mm. Described fom an unknown host
in China, and recently collected from Dalbergia
millettii in Zhejiang province (Q. Liu et al. 2013). Martin & Agarwala (1994) provided a
redescription of the aptera. Anomalosiphum tiomanensis
Martin & Agarwala Apterae are
pale green; BL 1.3-1.5 mm. On new
apical growth of Dalbergia torta (= D. candenatensis) in
west Malaysia. Also collected from China on Dalbergia sp. in Hong Kong (Quednau & Martin 2006), and it is
also recorded from Phyllanthus emblica
in Guanxi province (Q. Liu et al. 2013).
Apterae and alatae were collected in February. The life cycle is
unknown.
A holarctic genus for two species feeding on runners of Potentilla, with subapical reticulation on the siphunculi, extensive dark dorsal sclerotisation and ventral spiculosity on the head. Closely related to Macrosiphum, the differences possibly being due to the different microhabitat. Anthracosiphon crystleae Smith Apterae are reddish brown, sclerotic areas shiny, with black antennae and siphunculi; BL 1.8-2.4 mm. Alatae have 11-16 secondary rhinaria on ANT III. On runners of Potentilla canadensis and Potentilla sp. in eastern North America (North Carolina, Pennsylvania, Quebec). Presumably monoecious holocyclic, with alate males (original description). Anthracosiphon hertae Hille Ris Lambers Plate 26i Apterae are shiny black dorsally, lead-coloured with rosy hue ventrally, appendages blackish; BL 2.3-2.8 mm. Alatae have 20-34 secondary rhinaria on ANT III. On runners of Potentilla spp. (anserina, palustris) in northern Europe. Monoecious holocyclic, with alate males and oviparae in September (original description and Prior 1971).
One species in China with
strong similarity to Sitobion and possibly not justifying separate
generic status. Antimacrosiphum
bullacaudatum Zhang
Apterae colour in life unrecorded, probably pale with dark siphunculi;
BL about 2.2 mm. Feeding on Phyllanthus emblica in Yunnan, China (G.
Zhang & Qiao 1998c).
A
palaearctic genus of 10 species with fundatrices that typically crumple or
roll the leaves of Pyrus in spring
into “pseudogalls”. The progeny of the fundatrix are all
alate and migrate to subterranean parts of Umbelliferae/Apiaceae or
Compositae/Asteraceae, where they
are ant-attended. The name Anuraphis has been applied more widely
in the past to include species currently placed in Brachycaudus and Dysaphis.
See Heie (1992) and Blackman (2010) for accounts of the three species in
north-west Europe, Barbagallo & Cocuzza (2003) for morphometric studies
and a key to western palaearctic species, and Kolesova (1970, 1972) for
ecological studies of four species in Crimea. Anuraphis cachryos Barbagallo & Stroyan Apterae are very dark brown to black, BL 2.0-3.0 mm. On main root and leaf bases below soil level of Cachrys spp. Also recorded from Echinophora tenuifolia and Ferula communis. In southern Italy (where it is apparently anholocyclic), and also recorded from Iran and Turkey. Anuraphis capparidis Nevsky Apterae are brown; BL c. 1.9 mm. On Capparis spinosa in Central Asia. Presence of marginal tubercles on abdominal tergites 1 and 7 indicates that this is not an Anuraphis; perhaps it is an immature Aphis sp. Anuraphis catonii Hille Ris Lambers Fundatrices fold leaves of Pyrus communis in spring, their all-alate progeny migrating to form colonies at ground level on Pimpinella spp. Apterae on the secondary host are bright yellowish or greenish, with brown legs and antenae; BL 1.6-2.4 mm. Alatae are much darker, with a large dark dorsal patch. Kolesova (1972) studied its ecology in Crimea. Secondary host populations occur throughout much of Europe, and in south-west and Central Asia (Kazakhstan; Kadyrbekov 2017a), but the pear-feeding generations are reported only from Crimea and Switzerland (Lampel & Meier 2007). 2n=26 (Kuznetsova 1975, by implication). Anuraphis cortusae Nevsky Apterae are pale green with black eyes and dark tips to siphunculi; BL c. 1.9 mm. On Cortusa matthiola in Central Asia. Anuraphis farfarae (Koch) Pear-Colt’s Foot Aphid. The large (BL 2.7-3.7 mm) plump-bodied brown fundatrices roll or fold leaves of Pyrus communis in spring, producing green progeny that all develop into alatae. These migrate to basal and subterranean parts of Tussilago, or sometimes they may occur on Petasites or Hieracium. Apterae on secondary hosts are dark greyish green often tinged with red laterally and with a yellowish spinal stripe, and with dark antennae, legs and siphunculi; BL 2.2-3.1 mm. Immatures are paler, yellowish white, reddish, or pale green with a yellowish white spinal stripe. Alatae have a broad dark patch on abdominal tergites 4-6, almost solid in spring migrants but smaller and with a large window in alatae produced on the secondary host. The oviparae on pear in autumn are relatively small and pale (BL 1.6-1.8 mm) with hind tibiae swollen along the entire length and bearing numerous scent glands. Males are relatively large (BL 2.3-2.6 mm) with secondary rhinaria distributed III c.140-150, IV 42-51, V 13-29, base VI 0-1. Part of the population probably also overwinters parthenogenetically on Tussilago roots (Shaposhnikov & Sharov 1977). In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), west Siberia (Stekolshchikov et al. 2008a) and south-west and Central Asia; a report of its occurrence in North America is unsupported (Foottit et al. 2006). 2n=12. Anuraphis ferulae Shaposhnikov Colour of apterae in life unknown: BL c.3.3 mm. On sheaths of lower leaves of Ferula spp. in Tajikistan. Probably migrating from “pseudogalls” on Pyrus (Shaposhnikov 1995), but the primary host generations have not been described. Anuraphis pyrilaseri Shaposhnikov Apterae are pale ochre-yellow with pale green cross-bands; BL 2.3-3.1 mm. On basal parts of many genera of Apiaceae/Umbelliferae (Cachrys, Ferula, Laser, Laserpitium, Magydaris, Opopanax, Thapsia). In south, central and eastern Europe, Russia (northern Caucasus) and Kazakhstan (Kadyrbekov 2017a). Heteroecious holocyclic in eastern Europe (Crimea), where pseudogalls were found in spring on Pyrus communis (Kolesova 1972), and in Switzerland it is recorded from P. communis ssp. pyraster (Lampel & Meier 2007). Anholocylic on Ferula in southern Italy (Barbagallo & Stroyan 1982), and probably elsewhere. 2n=12. Anuraphis shaposhnikovi Barbagallo & Cocuzza Apterae are yellowish-ochreous to brownish; BL 1.8-2.6 mm. On root collar and leaf bases of Magydaris pastinacea and Opopanax chironium in southern Italy. Closely related to A. subterranea, but with relatively shorter R IV+V and fewer spinal tubercles (Barbagallo & Cocuzza 2003). Anuraphis
subterranea (Walker, 1852) Pear-Hogweed
Aphid Plate 9h Plump-bodied
brown fundatrices (BL 2.6-3.0 mm) roll and crumple the leaves of pear trees
in spring like A. farfarae, but
their progeny are brown (see influentialpoints.com/Gallery),
developing as alatae which migrate in May to basal parts of certain Umbelliferae/Apiaceae, especially Heracleum and Pastinaca.
Apterae on secondary hosts are brown,
BL 2.0-3.5 mm, and their immature progeny are pale brown. They form large
ant-attended colonies inside lower leaf sheaths, at stem bases and on
subterranean parts; the mutualistic relationship with Lasius on Heracleum was
studied by Hansen et al. (2006).
The return migration to pear occurs in September. Alatae have a similar
dorsal sclerotic pattern to A. farfarae.
Oviparae on pear are small (BL 1.4-1.8 mm), relatively pale, with strongly
swollen hind tibiae bearing numerous scent glands. Males are relatively large (2.3-2.7 mm) with secondary
rhinaria distributed III c.120-140, IV 28-57, V 5-12, base VI 0-2. Throughout Europe, North Africa
(Tunisia; Boukhris-Bouhachem et
al.
2007), Iran, western Siberia and Kazakhstan (Kadyrbekov 2014f). Cited to have
been found in Colorado, USA, but there are no North American specimens in
USNM (Foottit et al. 2006). 2n=26.
One species on Cotoneaster, related to Dysaphis but with extensive dorsal sclerotisation and differences in the siphunculi and cauda (Shaposhnikov 1959a). Anuromyzus cotoneasteris (Shaposhnikov) (Fig.24a,b) Apterae are very shiny dark brown to black; BL 2.2-2.5 mm. Secondary rhinaria of alatae (in spring) are distributed III 63-80, IV 18-23, V 3-6. On undersides of curled leaves of Cotoneaster spp. Known from Georgia, southern Ukraine (BMNH collection, leg G. Shaposhnikov) and the south of France (BMNH collection, leg. G. Remaudičre). Monoecious holocyclic, with oviparae and alate males in October (original description). 2n=12.
A
genus differing from Phylloxera in the absence of spiracles from
abdominal segments 2-5. Two species
are described from Pyrus, and
are of economic importance (see Blackman & Eastop 2000, pp.224-5). Aphanostigma iaksuiense
(Kishida) Very small yellow
phylloxerids laying eggs in bark crevices of Pyrus pyrifolia in Japan
and Korea, and also in Sichuan province, China, where its ecology was studied
by Fang (1980). Aphanostigma piri
(Cholodkovsky) Apterae are very small
(BL 0.8-1.0 mm), yellow, laying eggs in bark crevices of Pyrus communis,
also attacking and sometimes destroying the buds, and in some pear varieties
causing necrosis around the calyx of the fruit. In southern Europe, Ukraine,
Georgia, Crimea, Israel, Lebanon and also now in Thailand. A sexual phase is
reported from southern France and Crimea, but it is apparently anholocyclic
in Israel.
A monotypic genus characterised
by the single developed tarsal claw of apterae on the secondary host. Emigrant alatae from galls on Ulmus have
short antennae, forewings with unbranched media and small siphuncular pores
without associated hairs. A cladistic study (Sano & Akimoto 2011) has
indicated a close relationship with another monotypic east Asian genus Schizoneurella, which despite some
obvious morphological differences has a remarkably similar life cycle. Aphidounguis
mali
Takahashi (fig. 117A) Host-alternates between Ulmus parvifolia
and roots of Malus spp.
in Japan. Alatae have also been
collected from U. parvifolia in
China (Hangzhou), and from U. japonica
and U. davidiana var. mandshurica in Korea. It was recently reported from North
Carolina, USA (Footit et al. 2006).
The Ulmus galls consist of several
rolled leaves on one shoot; initially only one of these is occupied by the
fundatrix, but later its progeny migrate to fill the empty leaf-rolls
(Akimoto 1983). Alate emigrants, BL
1.5-1.7 mm, have dark brown head, thorax, legs and antennae, and dark bands
between the abdominal tergites.
Apterae on roots of Malus
are small white aphids secreting wax wool, to which the name Watabura nishyae Matsumura has been
frequently but wrongly applied. The alatae collected on apple roots in
October in China described as A.
pomiradicola Zhang & Hu (G. Zhang et
al. 1999) are likely to be the sexuparae of A. mali. Apterae found causing economic damage to apple roots in
Yunnan, China, and identified as Schizoneurella
indica (Qiao et al. 2005d), may
also be this species. [Note: the record of A. mali from Cydonia in Blackman & Eastop (2000) is in error.]
A palaearctic genus of about 24 species, mostly in southern Europe or central Asia. They feed mainly at stem apices of Caryophyllaceae, but are little studied and life cycles are mostly unknown. Some species may host-alternate from Rosaceae, as one species (A. bozkhoae) is described from Prunus, and two from Prinsepia. They are Myzus-like aphids, but the presence of a pair of mesosternal processes indicates a relationship with Brachycaudus subgenus Acaudus; other Brachycaudus-like characters are the tendency for extensive dorsal sclerotisation, and the subapical annular incision of the siphunculi. Kadyrbekov (2013) described six new species from Kazakhstan. Nieto Nafría et al. (2013a, 2014) described more new species, reviewed the genus and provided a key to apterae, which was updated by Barjadze (2019). Aphidura acanthophylli Remaudičre Apterae are pale; BL 1.4-1.7 mm. Alatae have secondary rhinaria distributed III 15-22, IV 3-11, V 0-4. This species differs from other Aphidura in having apterae with dorsal hairs arising from conical tubercles. On an Acanthophyllum sp. in Iran (Remaudičre 1989b). Nieto Nafría et al. (2013a) provided a photomicrograph of a mounted specimen. Aphidura
alatavica Kadyrbekov Apterae have variable and often extensive
black dorsal sclerotisation, with unsclerotised areas yellowish green, and
brown siphunculi and cauda; BL 1.7-2.0 mm. Living on Cerastium cerastoides
and C. holosteoides (= fontanum ssp. vulgare), forming
loose colonies on stems under flowers. In south-east Kazakhstan (Kadyrbekov
2013). Aphidura amphorosiphon Nieto Nafría, Mier Durante & Remaudičre Colour of apterae in life unknown; BL
1.3-1.6 mm. Alatae have 16-21 secondary rhinaria on ANT III, and a black
dorsal abdominal patch. On Silene
sp. and Eremogone rigida in Iran
(Nieto Nafría et al. 2013a,
Barjadze et al. 2017b). An alate
male was collected in mid-September. Aphidura
bozhkoae (Narzikulov)
Apterae are shiny black aphids, often with black siphunculi, BL
1.7-2.0 mm, infesting apical leaves of Prunus
spp. (but not P. avium or P. cerasus), which curl up and turn
red (Narzikulov 1958, 1965). Kadyrbekov
(2013) also recorded it from Aflatunia (=
Louiseania) ulmifolia. In mountainous areas of central Asia (Tajikistan,
Kazakhstan, Kirgizia, Georgia, Iran). A single aptera collected on Hordeum in Pakistan (leg. C.I.B.C.,
13.ii.63) also seems to be this species. The variation of key features of the
morphology of this species (dorsal sclerotisation, pigmentation of
siphunculi, development of mesosternal processes) was discussed and
illustrated by Nieto
Nafría et
al.
(2014). The life cycle is unknown. Aphidura
corsicensis Nieto
Nafría, Blackman & Martin
Apterae are shining dark green;
BL 2.0-2.1 mm. On Cerastium
soleirolii in the mountains of
Corsica at 1100m (Nieto Nafría et al. 2014). Other morphs and biology are
unknown. Aphidura
delmasi Remaudičre & Leclant Apterae are shining dark brown with
black siphunculi and pale cauda; BL 1.3-2.0 mm. Alatae
have 3-8 secondary rhinaria on ANT III only. On Silene italica and Silene sp. in southern Europe (France, Sicily, Greece). Monoecious holocyclic, with oviparae and apterous males
occuring in alpine France in late October; these morphs and the fundatrix
were described by Nieto Nafría et al.
(2013a). Aphidura gallica Nieto Nafría, Mier Durante & Remaudičre Apterae with variable black dorsal
sclerotisation and black siphunculi and pale cauda, general colour in life
unknown; BL 1.5-2.3 mm. Alatae have 16-34 secondary rhinaria on ANT III. On Silene spp. (gallica, paradoxa) in
France (Nieto Nafría et al. 2013a).
The life cycle is unknown. Aphidura graeca Nieto Nafría, Mier Durante & Remaudičre Colour of apterae in life is unknown; BL
c.1.8 mm. Alatae are undescribed. On Gypsophila
sp. in Greece (Nieto Nafría et al.
2013a). Aphidura
gypsophilae Mamontova-Solukha
Apterae are yellow, BL c.1.9-2.1 mm. Alatae have 10-15 secondary
rhinaria on ANT III only. On undersides of leaves and stems of Gypsophila spp. Recorded from Ukraine, Hungary, Slovakia, Russia (west
Siberia), Iran (Rezwani et al.
1994) and Kazakhstan (Nieto Nafría et
al. 2013a). The life cycle is unknown. Aphidura iranensis Nieto Nafría, Mier Durante & Remaudičre Apterae with black dorsal
sclerotisation, colour in life unknown; BL 1.1-1.3 mm. On an unidentified
species of Prunus in Iran (Nieto
Nafría et al. 2013a). The life cycle
is unknown. Aphidura jimoi Barjadze Colour of apterae in life
is unknown; BL 1.2-1.6 mm. On a species of Dianthus in Georgia (Barjadze 2019). Other morphs and life cycle
are unknown. Aphidura libanensis Nieto Nafría, Blackman & Martin
Apterae with black dorsal sclerotisation, colour in lfe unknown; BL
c.1.8-1.9 mm. On Prunus prostrata
in Lebanon (Nieto Nafría et al.
2014). Other morphs and life cycle are unknown. Aphidura massagetica Kadyrbekov Apterae have extensive black dorsal markings with unsclerotised areas yellowish green; BL 1.9-2.4 mm. An alatiform aptera had 22-24 secondary rhinaria on ANT III. In loose colonies on peduncles of Silene lithophila and S. brahuica in south and south-east Kazakhstan (Kadyrbekov 2013). Aphidura
melandrii Kadyrbekov Apterae have extensive black dorsal
sclerotisation with unsclerotised areas yellowish green, and brown siphunculi
and cauda; BL 1.9-2.3 mm. Living on Silene
latifolia (= Melandrium album),
forming loose colonies on stems under flowers. In east and south-east
Kazakhstan (Kadyrbekov 2013). Very similar to A. ornatella. Aphidura mordvilkoi Shaposhnikov Apterae are green or brownish green; BL c.1.8 mm. On shoot apices and petioles of Prinsepia sinensis in east Siberia. A. prinsepiae Pashchenko is a synonym (Kadyrbekov 2013). Aphidura naimanica Kadyrbekov Apterae have variable black dorsal sclerotisation with unsclerotised areas yellowish green; BL 1.6-2.2 mm. The one described alata has 26-27 secondary rhinaria on ANT III. In loose colonies on stem under flowers of Gypsophila spp., in east and south-east Kazakhstan (Kadyrbekov 2013). Aphidura nomadica Kadyrbekov Apterae have a black dorsal abdominal patch, with unsclerotised areas green; BL 1.7-2.3 mm. In loose colonies on peduncles of Silene spp. in Kazakhstan (Kadyrbekov 2013). Aphidura ornata Hille Ris Lambers Apterae are shining black with unsclerotized areas pale greenish grey, siphunculi black and cauda greenish yellow; BL 1.8-2.2 mm. In rather dense colonies on stem apices and peduncles of Silene spp., sometimes visited by ants. Europe (Italy, France, Switzerland, Hungary, Russia, Ukraine) and Iran (Rezwani 1990). Aphidura
ornatella Narzikulov & Winkler Apterae are dark brown to black with
unsclerotized areas green; BL 1.5-2.6 mm. Kadyrbekov (2013) provided a
redescription, and described an alata with 29-30 secondary rhinaria on ANT
III. Living singly or in loose colonies on shoot apices and peduncles of Silene spp. and Saponaria sp., in Tajikistan, Kazakhstan and also in Iran
(Rezwani 2010), Pakistan
(Nieto Nafría et al. 2013a) and north-west India (David et
al. 1970, as A. bharatia).
Differences from A. bharatia noted
by Kadyrbekov (2013) may be due to intraspecific variation. Aphidura pakistanensis Nieto Nafría, Mier Durante & Remaudičre Colour of apterae in life is unknown; BL
1.6-2.0 mm. Alatae have secondary rhinaria distributed ANT III 32-48, IV
2-10, V 0 (Barjadze 2019). On unidentified Dianthus sp(p). Described initially from Pakistan (Nieto Nafría et al. 2013a), and also now recorded
from Iran (Barjadze et al. 2017b),
and from Georgia (Barjadze 2019). Barjadze (2019) compared the morphology of
apterae from different locations and discussed intraspecific variation. Aphidura
pannonica Szelegiewicz Apterae are green with black dorsal
markings, black siphunculi and dark green cauda; BL 1.7-2.3 mm. Alatae have
14-22 secondary rhinaria on ANT III only. On Silene spp. in eastern Europe (Hungary, Slovakia, Ukraine), also
in Kazakhstan (Kadyrbekov 2017a), and there is also a record from Gypsophila
paniculata (Nieto Nafría et al. 2013a). The Ukrainian population was given
subspecies status (A. pannonica ssp.
cretacea Mamontova-Solukha), but it
is difficult to see the justification for this. 2n=12. Aphidura picta
Hille Ris Lambers (= Aphidura
mingens Pintera) Plate 11i Apterae are yellowish to green with pale
brown head and prothorax, very variable shiny brown dorsal abdominal markings,
and dusky to brown-black siphunculi; BL 1.2-2.1 mm. Alatae have 25-39
secondary rhinaria on ANT III only. On flower-stalks of Silene spp. and Dianthus spp.
in southern and eastern Europe (Sicily, Greece, Bulgaria, former Yugoslavia, Israel,
Turkey) and south-west and central
Asia (Iran, west Siberia, Afghanistan, Tajikistan, Pakistan). Narzikulov &
Umarov (1969) gave subspecies status (A.
picta ssp. pamirica) to a
population in western Pamir. Aphidura prinsepiae Pashchenko Apterae are green or brownish green; BL c.1.8 mm. On shoot apices and petioles of Prinsepia sinensis in east Siberia (Pashchenko 1988a). Probably synonymous with A. mordvilkoi. Aphidura
pujoli (Gómez Menor) Apterae are greenish, with pale/dusky
siphunculi and cauda; BL 1.5-2.1 mm. Alatae have
22-25 secondary rhinaria on ANT III only. On
Dianthus spp., especially D. caryophyllus, in southern Europe (Spain, Portugal,
southern France, Corsica, Italy, Switzerland) and Ukraine, and also
tentatively recorded from Silene
parviflora (Bozhko 1976). The record from Pakistan (Naumann-Etienne
& Remaudičre 1995) should be applied to A.
pakistanensis (Nieto Nafría et al. 2013a). Monoecious holocyclic in Corsica, where oviparae
and alate males were collected in October
(BMNH collection, leg. F. Leclant). Aphidura
togaica Kadyrbekov Apterae are pale yellow with paired brown
dorsal markings; BL 1.6-2.0 mm. Alatae have 22-36 secondary rhinaria on ANT III
and 0-1 on IV. In loose colonies on upper and lower sides of leaves of Gypsophila perfoliata and G. altissima in south-east Kazakhstan
(Kadyrbekov 2013), and also now recorded from G. polyclada in Iran (Barjadze et al. 2017b). Aphidura urmiensis Nieto Nafría, Mier Durante & Remaudičre Colour of apterae in life is unknown; BL
1.9-2.1 mm. Alatae are undescribed. On
Spergularia marina in Iran (Nieto
Nafría et al. 2013a). The life
cycle is unknown.
One rather large species on Rosa in Central Asia with an unusual combination of dark dorsal pigmentation, pale siphunculi, blackish cauda, and numerous secondary rhinaria in apterae as well as alatae, with no clear relationship to other rose-feeding aphids (Umarov & Ibraimova 1967). Aphiduromyzus rosae Umarov & Ibraimova Apterae are dark green or brown with ANT I, II and VI, femora and cauda blackish, siphunculi pale; BL c.2.8 mm. Apterae have 20-25 secondary rhinaria on ANT III only, and alatae have them distributed III 30-35, IV c.6, V c.2. Alatae have an extensive black dorsal abdominal patch. On Rosa sp. in Tyan-Shan Mountains, Central Asia.
The largest aphid genus containing more than 500 species,
most of which occur in the northern hemisphere, but a few are native to South
America, New Zealand, and Australia.
Mostly they are small to medium‑sized aphids, often greenish but
varying from pale to dark green, and sometimes yellow, brown, reddish, or
black. Most species live on shrubs or
herbs, with relatively few on trees.
They usually live on the young foliage and often distort the leaves,
but some species live on young twigs or at the base of the stem, or on the
roots. Most species are attended by
ants which may construct earthen shelters for the species living just above
soil level. There are evident species groups associated with particular
groups of plants, e.g. subgen. Bursaphis
host-alternating between Ribes and
Onagraceae, but there seems to be no overall pattern of host association and
aphid morphology on which to base a subgeneric classification with world-wide
applicability. However, phylogenetic studies using both molecular (Coeur
d’acier et al. 2007b, Kim & Lee
2008) and morphometric methods (Kim et
al. 2010a) have confirmed that certain sub-groupings apply within the
genus across the palaearctic region, and Lagos et al. (2014) have shown that they also apply to North American
species. The primary hosts of the relatively few Aphis species that host-alternate provide no evidence of
primitive host associations. An interesting possible explanation might lie in
the suggestion that northern Aphis species
might be derived from southern hemisphere ancestors (von Dohlen & Teulon
2003).
Accounts are available for the British Isles (Stroyan 1984),
Fennoscandia/Denmark (Heie 1986), Switzerland (Lampel & Meier 2007), the
Iberian peninsula (Nieto Nafría et al.
2005a), India (A.K. Ghosh & L.K. Ghosh 2006), Mongolia (Holman & Szelegiewicz, 1971), east
Siberia (Pashchenko 1992, 1993a-c, 1994a-c) and Japan (Takahashi 1966).
Pashchenko (1997a,b) provided keys to the east Siberian species. Brown (1989)
keyed alatae of the north-western European species. Cook (1984b) reviewed and
keyed the North American Compositae/Asteraceae-feeding species. Lagos et al. (2014) analysed morphological
and molecular data for Aphis spp.
in midwestern USA and identified four main species groups. Lagos-Kutz et al. (2016) reviewed and keyed the
species of the “asclepiadis group”,
which have a primary host asssociation with Cornus. An interactive key to Aphis
species of midwestern USA is available online at http://imperialis.inhs.illinois.edu/lagos/key.asp?key=Aphis&lng=En&i=1&keyN=1.
Remaudičre (1994) keyed the South American species. Rusanova (1948) reviewed
and keyed species in Azerbaijan, and described several new species that have
not subsequently been recognised.
Subgenus Bursaphis was
revised by Remaudičre (1993b), and subsequently studied intensively by
Rakauskas and co-workers (e.g. Rakauskas 1998b, Rakauskas et al. 2011). Martin (1991)
provided a key to Toxoptera, now
placed as a subgenus of Aphis, and
Qiao et al. (2008) reviewed and
keyed the species of Toxoptera in
China. Xu et al. (2021) provided a key to Aphis
spp. with stridulatory spines on the hind tibiae. With
some reservations we follow Kadyrbekov (2001c) in transferring all the species
previously placed in Aphis subgenus
Absinthaphis to Xerobion. We have previously treated Protaphis as a subgenus of Aphis, but now give it separate generic status, in accordance
with Aphid Species File. Aphis aba Miller
Apterae have deep brown head and thorax, paler brown to pale red
abdomen with dark brown marginal spots; BL 1.2-1.4 mm. Alatae have secondary
rhinaria distributed III 18-22, IV 4-6, V 2-3. On leaf petioles and midribs
of Iva
xanthifolia in Massachusetts, USA.
Aphis acaenaevora
Mier
Durante & Ortego Apterae are
greenish yellow, lightly wax-dusted, with dark brown or black head, thorax,
siphunculi and cauda; BL 1.8-2.4 mm. Apterae have 1-8 secondary rhinaria on
ANT III, alatae are unknown. On Acaena
splendens, forming colonies at base of stem at or slightly below ground
level. In Argentina (Andean region of Patagonia). Monoecious holocyclic with
oviparae and apterous males in late April (Mier Durante & Ortego 1998). Aphis acaenovinae Eastop Apterae are bright apple-green with three
rusty orange blotches on abdomen, one at base of each siphunculus and the
other in between; BL 1.4-2.0 mm. Apterae have 0-5 secondary rhinaria on ANT
III, alatae have them distributed III 3-8, IV 0-2. On Acaena ovina, living at ground level in rosette, attended by
ants. In South-east Australia. 2n=8 (D.F. Hales, pers. comm.). Aphis acanthoidis (Börner) Apterae are dark green; BL 1.1-1.6 mm. On
leaf rosettes and developing flower
stems of Carduus spp. In Netherlands,
central and eastern Europe, Italy (Barbagallo & Massimino Cocuzza 2014),
western Siberia, and Kazakhstan, where it is also recorded from Cousinia sp. and Jurinea sp. (Kadyrbekov 2017a),. A record from Carthamus (= Carduncellus) plumosus in Algeria (Laamari et al. 2013) needs further comfirmation.
Aphis acanthopanaci
Matsumura Apterae have BL 2.0-2.3 mm
and are dark green according to original description of specimens from Acanthopanax ricinifolium (= Kalopanax septemlobus) (Matsumura 1917).
Midsummer specimens (from Aralia
cordata) have BL about 1.3 mm and are yellow according to Takahashi
(1966), but he possibly had A. spiraecola. Only known from Japan. Alatae are undescribed, and the life cycle
is unknown. Aphis
acetosae L. Apterae are dark green to black; BL 1.5-2.7
mm. Alatae have secondary rhinaria distributed III 4-21, IV 0-2. On stems and
inflorescences of Rumex spp., especially R.
acetosa, often in large
ant-attended colonies. It has also been found on Rheum spp. (Pashchenko 1988a).
In northern, central and eastern
Europe, Iran, Kazakhstan, east and west Siberia, and now also recorded from
Algeria (Laamari et al. 2013). Also
recorded from USA, but this needs confirmation (Footit et al. 2006). Aphis (Toxopterina) achillearadicis Pashchenko Apterae are dark green; BL c.1.4 mm. In
small dense colonies on roots of Achillea
ptarmicifolia. East Siberia (Maritime Territory). Clearly it is closely
related to A. vandergooti, and
therefore we have transferred it to subgenus Toxopterina. Aphis
achyranthi Theobald Apterae are dull yellowish brown with
dark brown siphunculi and cauda (see aphids of
Karnataka website), immatures
yellow with white wax (David, 1958); BL 1.2-1.7 mm. Alatae have
secondary rhinaria distributed III 16-20,
IV 9-12, V 3-5. Described from Achyranthes
sp. in southern India (Theobald 1929), and there are samples from A. aspera in BMNH collection from
Pakistan (leg. M.A. Ghani). David (1958) redescribed it from Punica granatum in southern India,
colonising tender shoots, not attended by ants, and dropping readily when
disturbed. 2n=8 (Kurl & Chauhan 1986b). Aphis acrita
Smith Apterae are greyish pulverulent, with
dark siphunculi and cauda; BL 1.7-1.9 mm. On Sedum sp. in Ohio, USA. Monoecious holocyclic, with both apterous
and alate males (original description). Aphis
acuminata Nieto Nafría
& von Dohlen Apterae are shining
brown to black; BL 1.2-1.7 mm. On stems of Adesmia sp. in Salta province, Argentina (Nieto Nafría et al. 2016c). Other morphs and life
cycle are unknown. Aphis adesmiae Delfino Apterae are shining
black, without wax powder; BL 1.3-2.1 mm. On Adesmia boronioides in Argentina, forming dense colonies mainly
on the stems and inflorescences (Delfino et
al. 2009). It was recently also recorded from Chile (Nieto Nafría et al. 2018). Alatae were described by
Nieto Nafría et al. (2016c). Probably monoecious holocyclic, but sexual morphs are
unknown. Aphis
affinis Del Guercio Apterae are pale yellow (small specimens)
to dark grey-green, often with narrow dark intersegmental cross bands,
immatures dusted with wax powder (see aphids of
Karnataka website); BL 0.6-2.1
mm. Alatae
have secondary rhinaria distributed III
3-9, IV 0. On Mentha spp., usually
clustered on stem apices, attended by ants. In Europe,
southern Russia, Middle East, Central Asia, India and Pakistan. Very small
apterae (BL 0.57-0.83 mm) collected on M.
suaveolens in Azores (BMNH collection, leg. F. Ilharco) are probably this
species. There are
single records from two other labiates, Rosmarinus
officinalis in Tuscany (Barbagallo et
al. 2011) and Marrubium alternidens (= anisodon) in Iran (Mohktari et
al. 2012). Monoecious holocyclic with
apterous males (Tuatay & Remaudičre 1964). Ilharco (1987) redescribed
apterae and alate viviparae. 2n=8. Aphis
agastachyos Hille Ris
Lambers Apterae are oval, shining
dark green (see aphidtrek.org),
BL 1.5-2.0 mm, curling leaves of Agastache
urticifolia and Agastache sp. in western USA (Idaho, Utah, Oregon, Washington). Possibly this species also
colonises Monardella odoratissima
(A. Jensen, pers. com.). Alatae have large dark marginal and postsiphuncular
sclerites. Apterous males and oviparae were collected along with viviparae in
July (original description), indicating
that the life cycle is abbreviated. Aphis
agrariae Bozhko Apterae are dark brown, lightly
wax-dusted, with black dorsal abdominal cross-bands and black appendages; BL
c.2.2-2.3 mm. On Euphorbia agraria,
colonising stems, flower stalks and leaves, in Ukraine. Aphis
albella Nevsky Apterae are white or pale yellow, with
appendages only slightly dusky; BL c.1.8-2.1 mm. On undersides of leaves
of Cichorium
intybus, and also recorded in smaller numbers from Portulaca, Hibiscus and
Foeniculum officinale. In southern
Kazakhstan/Uzbekistan. Apart from the
absence of pigmentation, which might be due to a genetic aberration or
unusual environmental conditions, this species could be A. gossypii. Aphis
alchemillae Börner Apterae are shiny, dark green to black,
with pale legs and antennae; BL 1.2-2.2 mm. On Alchemilla spp. (conjuncta,
glaberrima, vulgaris) colonising stems, petioles and undersides of leaves,
and sometimes in inflorescences (Börner 1952). In Germany and Austria. [From
examination of specimens in the BMNH collection there appear to be two Aphis species on Alchemilla under this name, one collected from ant shelters at
the stem bases of an unidentified species in Austria (leg. R.N.B. Prior), and
the other from A. conjuncta in
Frankfurt Botanical Garden (leg. M. Mackauer). Further work is necessary to
see whether either of these is alchemillae
Börner.] Aphis
alienus Theobald Apterae are green to dark green; BL
1.0-1.9 mm. Alatae
have secondary rhinaria distributed III
4-7, IV 0 -3. On rhizomes and basal parts of Teucrium spp., especially T.
scorodonia, in ant shelters. Known
from various countries in Europe (England, France, Spain, Portugal, Italy,
Czech Republic), and it has also been found in Pakistan (Naumann-Etienne
& Remaudičre 1995). Most pre-1983 records of this species are as A. scorodoniae (see Stroyan 1984). Aphis
alstroemeriae Essig Colour of apterae in life unknown,
probably dark; BL 1.6-2.1 mm. On an unidentified species of Alstroemeria (Chilean Lily) in
Aconcagua Province, Chile. Alatae have secondary rhinaria
distributed III 5-11, IV 0-2. Remaudičre
(1994) redescribed the paratypes in the BMNH collection. Aphis
althaeae (Nevsky) (= davletshinae Hille Ris Lambers of Blackman & Eastop 2006; see García
Prieto et al. 2004) Apterae are pale green, BL 1.6-1.8 mm, on
undersides of leaves and stems of Alcea
and Althaea spp., often forming
large colonies which curl leaves and stunt young growth. Alatae
have secondary rhinaria distributed III
6-15, IV 0-4, V 0-1. Nevsky (1929) also recorded it from Malva and Convolvulus, and Kadyrbekov (2004b) found it on Lavatera thuringiaca. Specimens
collected on Salvia sclarea in
Afghanistan were described as a subspecies, A. althaeae spp. afghanica Narzikulov & Umarov. In Europe (Germany, Poland, Russia,
Spain, Italy, Greece, ?Sweden) and south-west and central Asia (Iran, Turkey,
Lebanon, Kazakhstan, Uzbekistan). Monoecious holocyclic (Börner & Heinze
1957). Aphis
amaranthi Holman Apterae are dull olive green or reddish
brown, BL 1.5-2.1 mm. On Amaranthus spp., colonising the roots and stem bases,
and sometimes on the leaves or in the influorescences, attended by ants. Alatae have no dorsal abdominal
sclerotization except laterally and at bases of siphunculi, and secondary
rhinaria distributed III 8-22,
IV 0-8, V 0-2, those on III being distributed irregularly (not in a row). In
Brazil, Argentina, Cuba and USA
(California – BMNH collection, leg. H. Walker, 1975; Florida – Halbert
et al. 2000; Georgia and Louisiana
-trapped alatae in BMNH collection).
Life cycle is unknown; most common in winter in Cuba. 2n=8. Aphis
angelicae Lee & Seo, nec. Koch
1854 Apterae are pale yellow; BL not
recorded. On Angelica decursiva, in
ant shelter on stem. In Korea. Perhaps not an Aphis; apart from the 5-segmented antennae, it seems to have many
features in common with the species described from A. sinensis in China under the name Dysaphis angelicophaga (q.v). [Note: if these two should prove to
be synonymous, then angelicophaga becomes
the valid name, as Aphis angelicae is
preoccupied.] Aphis
antherici Holman (= liliago
Müller) Apterae dark bluish
green, olive brown or reddish brown to black, with basal part of antennal
flagellum, and tibiae for most of their lengths, whitish; BL 1.1-2.2 mm. Alatae have secondary rhinaria
distributed III 8-21, IV 0-3. On basal parts
of Anthericum spp., living in ant
shelters. In Czech Republic, Slovakia, Germany and Poland. Monoecious
holocyclic with apterous or brachypterous males. Holman (1966) and Müller
(1968a, as A. liliago) both gave
full accounts of this aphid. Aphis apocynicola
Holman Apterae are olive to brownish green with
black head and siphunculi, and pale cauda; BL 1.9-2.4 mm. Large colonies were
found on the terminal parts of the shoots of Apocynum scabrum (= A.
venetum var. scabrum?)in
Uzbekistan, and on A. lancifolium in Kazakhstan (Kadyrbekov &
Aoitzhanova 2005). Records of A.
apocyni Koch from Apocynum in
Kazakhstan, Uzbekistan and Tadzhikistan are referable to apocynicola,
and A. apocyni from China could be
hot weather dwarfs of this species. The life cycle is unknown. Aphis
aquilonalis
Stekolshchikov & Khruleva Apterae
are dark brown-black or dark dirty yellowish brown with sparse wax
pulverulence; BL 2.0-2.6 mm. On Rumex
arcticus (= R. aquaticus ssp. arcticus) in north-east Siberia
(Chukotka), on undersides of leaves and in inflorescences. The life cycle is
abbreviated; a collection in late July included old fundatrices, apterous and
alate viviparae, oviparae and apterous males. Stekolshchikov & Khruleva
(20150 provided descriptions of all morphs. Aphis araliaeradicis
Strom Apterae are pale brown, BL c.1.6 mm.
Apterae have 0-5 secondary rhinaria on ANT III, alatae have 8-11. Described
from Aralia nudicaulis in
Wisconsin, USA, forming ant-attended colonies on roots and stem-bases. Also
recorded from California (Smith & Parron 1978). Presumably monoecious
holocyclic, but males are undescribed. Aphis arbuti
Ferrari Apterae are wine red; BL 1.7-2.7 mm. Alatae
have secondary rhinaria distributed III
10-19, IV 0-6, V 0. On woody Ericaceae (Arbutus
unedo, Erica arborea) in
southern Europe, and also reported from Turkey (Görür et al. 2011a). A member of the taxonomically
difficult A. fabae group, and only
recognisable as a distinct species by its colour and host association. Mier Durante & Nieto Nafria (1986) provided
morphometric descriptions of apterae and alatae. Apparently monoecious and holocyclic
(Barbagallo & Stroyan 1982), but sexual morphs are still undescribed. Aphis
arctiumi Bozhko Apterae are yellow-green, wax-dusted; BL
c. 1.5 mm. On Arctium tomentosum in
Ukraine. Aphis
argrimoniae (Shinji) Apterae are green, with pale legs and
antennae, BL 1.2-1.8 mm. Alatae have dark antennae with secondary rhinaria
distributed III 12-24, IV 9-17, V 4-8.
On Agrimonia spp. in Asia
(Japan, Korea, east and west Siberia). Bozhko (1976a) described a similar aphid
from Agrimonia eupatoria in Ukraine
as a subspecies, A. argrimoniae ucrainica.
Aphis armata
Hausmann Apterae are black in life, resembling A. fabae (see influentialpoints.com/Gallery);
BL 2.2-2.9 mm. Alatae have secondary rhinaria distributed III 9-24, IV 0-8, V 0. On flower stems of Digitalis spp., vigorously attended by
ants. In Europe, from UK to Italy (Barbagallo & Patti 1998), and
introduced to British Columbia, Canada (BMNH collection, leg. C.-k. Chan).
Its distribution is uncertain, as morphology of parthenogenetic generations
overlaps with that of long-haired members of the A. fabae group. Monoecious
holocyclic, with oviparae having hind tibiae much less swollen than A. fabae. See Jacob (1947) for a detailed account. 2n=8. Aphis
artemisiphaga Holman Apterae are green or dark green, mottled,
matt; BL 1.1-1.5 mm. At base of stem of Artemisia
spp., attended by ants, in east Siberia. Aphis
artemisiphila Holman Apterae are green, similar to A. artemisiphaga; BL 1.5-1.8 mm. On
roots of an unidentified Artemisia sp.,
attended by ants, in east Siberia. Holman (original description) noted that
differences from A. artemisiphaga
could be seasonal and require confirmation. Aphis
(Pseudoprotaphis) artemisivora
Holman Apterae are dull green, lightly
wax-dusted, with a dark dorsal pattern; BL 1.9-2.4 mm. In ant-attended
colonies on roots of an unidentified Artemisia sp. in Mongolia. Aphis asclepiadis Fitch (= A.
helianthi Monell, = A. carduella
Walsh) Plate 6i Apterae on secondary hosts are pale green,
yellowish green to deep olive green mottled with yellowish green, with black
siphunculi (see influentialpoints.com/Gallery
and aphidtrek.org); BL 1.4-2.5
mm. Alatae have secondary rhinaria
distributed III 15-40, IV 0-2. Colonies occur in late spring and summer on
stems and leaves of many genera and species, especially Compositae/Asteraceae
and Umbelliferae/Apiaceae, and also able to colonise Apocynaceae. Heteroecious holocyclic, with a sexual phase on Cornus spp. Spring
populations twist and curl the leaves of Cornus
(Hottes & Frison 1931, as helianthi);
apterae on Cornus in spring are
1.5-1.8 mm, dark yellow to green, with siphunculi variably pigmented from
green to black. Oviparae in autumn
have considerably swollen hind tibiae. In
USA and Canada, and also recorded from Brazil on Yucca (assuming A. yuccicola Wilson is a synonym), Peru (Vilca Mallqui &
Vergara Cobián 2011) and Argentina (Gonzáles Rodríguez et al. 2016). Records
from Turkey (Akyürek et al. 2010, as helianthi) and
Kazakhstan (Kadyrbekov 2017a, as carduella)
need further confirmation. Lagos et al.
(2014) using both morphological and molecular data identified a clade of 10 species as “the asclepiadis group” (asclepiadis, cornifoliae, decepta, impatientis, neogillettei, nigratibialis,
salicariae, saniculae, thaspii, viburniphila), five of which have Cornus as their primary or only host.
Lagos-Kutz et al. (2016) reviewed
and keyed this group, and provided conclusive molecular evidence that A. carduella should be treated as a
junior synonym of A. asclepiadis.
2n=8. [This species, found on numerous host plants in
North America and now also introduced to South America, has a confused
nomenclatural history. The name A.
helianthi Monell was widely used in the North American literature,
especially for populations on Compositae/Asteraceae, until Cook (1984b)
established its synonymy with the earlier name A. carduella. Earlier, Addicott
(1981) had synonymised A. heraclella,
the name previously often applied to populations on Umbelliferae/Apiaceae,
with A. carduella (as helianthi) on the basis of host transfers and morphological studies
involving Cornus,
Umbelliferae/Apiaceae and Epilobium (Onagraceae),
but not Compositae/Asteraceae. Robinson & Chen (1969) had previously
obtained transfers from Cornus and Helianthus to Sium suave, but not to other Umbelliferae/Apiaceae.] Aphis
astericola Tissot Apterae are dark reddish brown with
mainly dark appendages; BL c. 1.7 mm.
Alatae have 7-9 secondary rhinaria on ANT III only. Only known from Aster sp. in Florida, USA. Aphis
astragali Ossiannilsson (Fig.16i) Apterae are dull black, powdered with
whitish wax; BL 1.9-2.7 mm. On Astragalus
spp. in Sweden and Finland, where it is also recorded from Oxytropis campestris, and where oviparae and alate males are produced
(on both host genera) as early as July (Heikinheimo 1984). Also recorded from
Kazakhstan (Kadyrbekov 2017a). A.
masoni on islands in subarctic Canada is possibly a synonym. A record
from Algeria (Laamari et al. 2013)
needs further confirmation. Aphis
astragalicola Holman &
Szelegiewicz Apterae are black, with
middle parts of antennae and tibiae whitish; BL 0.9-1.2 mm. On Astragalus sp., “presumably living on
the basal parts of the stems” (from original description), and A. danicus (Stekolshchikov et al. 2008a). In Russia (west Siberia
and Transbaikalia), and also recorded from Turkey (Şenol et al. 2014c), Moldova and Kazakhstan
(Kadyrbekov 2017a). Alatae and sexual morphs are unknown. Aphis
astragalina Hille Ris
Lambers (Fig.16j) Colour of aptera in life unknown; BL c.2.4-2.5 mm. Described from a
white-flowered leguminous plant in Montana, USA and Astragalus sp. in Alberta, Canada, and there are later records from
Astragalus pectinatus in Alberta
and from Hedysarum alpinum in
Manitoba (Robinson 1991) and Alaska (BMNH collection, leg. A.G. Robinson). Aphis
aubletia Sanborn Apterae are undescribed, alatae have
black head and thorax and dusky to pale brown abdomen, siphunculi concolorous
with abdomen, and cauda “black on distal half, hirsute” (original
description); BL of alata c.1.3 mm. Alatae have secondary rhinaria
distributed III 7, IV 3, V 1. Colonising all the above-ground parts of Verbena aubletia (= Glandularia canadensis) in May in
Kansas, USA. Not recorded since original description. Aphis (Toxoptera) aurantii
Boyer de Fonscolombe Black Citrus Aphid Plate 13c, d Apterae are oval shiny, reddish-brown or
black, with black‑and‑white
banded antennae and black siphunculi and cauda (see
influentialpoints.com/Gallery);
BL 1.1-2.0 mm. Immatures are brownish. Alatae are brown-black with a black
pterostigma and, unusually for Aphidinae, a usually once-branched media; they
have secondary rhinaria distributed ANT III 2-14, IV 0(-6). In dense,
ant-attended colonies on shoots and undersides of young leaves of host
plants, causing slight rolling, twisting or bending of mid-ribs. Large
colonies produce an audible scraping sound when disturbed. Found on more than 120
plant species, particularly trees and shrubs, in numerous families,
especially Anacardiaceae, Anonaceae, Araliaceae, Euphorbiaceae, Lauraceae,
Moraceae, Rubiaceae, Rutaceae, Sterculiaceae, and Theaceae. It is an
important vector or viruses of economically-important plants (Blackman &
Eastop, 2000, p.354). Distributed throughout the tropics and subtropics
including the Pacific Islands, and in glasshouses in temperate climates. Apparently
it is almost completely anholocyclic everywhere; however two alate males were
collected on Citrus in December
1936 in Palestine (leg. F.S. Bodenheimer), and one ovipara was recently found
in a sample from Sambucus in Malta
(Pérez Hidalgo & Mifsud 2011). Stary et
al. (1988) reported on biocontrol measures in southern France, and
Hussein & Kawar (1985) studied natural enemies in Lebanon. Broughton
& Harris (1971) analysed the sound produced by A. (T.) aurantii, which is the only aphid with audible
stridulation. Populations in which alatae have several secondary rhinaria on
ANT IV have been recorded on Celtis
in Japan (as Aphis celtis Shinji
1922c) and on Ficus in China
(referred here to A. schlingeri).
Alatae of A. aurantii normally have
only 0-1 rhinaria on ANT IV. 2n=8. Aphis
austriaca Hille Ris Lambers Apterae are pale sea-green, darker
marginally, with head and prothorax blackish green, very faintly pruinose; BL
1.6-1.9 mm. Alatae have secondary rhinaria distributed (1 specimen) III
17-20, IV 4, V 0. Described from roots of
Silene sp., probably S. nutans, in Austria, and also
recorded from Poland, feeding quite deeply on roots of S. otites (Osiadacz 2009). Ant-attended. Very closely related to A. sambuci, and possibly just a
variant anholocyclic population of that species. Aphis
axyriphaga Pashchenko Apterae are dark dingy-green, BL c. 1.3
mm. The holotype aptera has a pair of dark irregular sclerotic patches on
dorsal abdomen, leaving broad unsclerotised spinal band. Alata has 4 large
secondary rhinaria on ANT III. On undersides of leaves of Axyris amaranthoides in east Siberia. Aphis
axyriradicis Pashchenko Apterae are dark emerald green, BL c. 1.7
mm. On roots of Axyris amaranthoides,
attended by ants. Alatae are undescribed. In east Siberia. Aphis
baccharicola Hille Ris
Lambers Colour of apterae in life
unknown, BL 1.4-1.7 mm. Alatae have 14-22 secondary rhinaria on ANT III only.
On Baccharis pilularis and Baccharis sp. in western USA
(California, Oregon). A member of the A.
carduella group, but with long, finely-pointed antennal hairs. Aphis
ballotae Passerini (= balloticola Szelegiewicz of Blackman & Eastop 2006; see García
Prieto et al. 2004) Apterae are dark grey-blue to mottled
green (see also influentialpoints.com/Gallery);
BL 1.0-2.0 mm. Alatae have secondary rhinaria distributed III 3-12, IV 0-3, V 0. On stems and undersides of
leaves of Ballota spp. causing
slight downward leaf-curl in early summer (Stroyan, 1984). Also recorded from
Dracocephalum, Leonurus and Marrubium spp.
Alatae have secondary rhinaria distributed III 3-12, IV 0-3. Monoecious
holocyclic, with alate males (Börner 1950; Tuatay & Remaudičre
1964). Throughout Europe (except
Scandinavia), and eastward to Crimea, Iran and Turkey. There is also a record
from Algeria (Laamari et al. 2013).
A member of the A. frangulae complex,
difficult to distinguish except by biology and host plant. Aphis
berberidorum Ortego &
Mier Durante Apterae are brown, with
antennae and legs partly pale; BL 1.5-2.4 mm.
Alata has 6-12 secondary rhinaria on ANT III only. On Berberis spp. in the southern Andes
(Chile and Argentina). Males are alate (Ortego & Mier Durante 1997). Apparently a member of the craccivora group. Aphis
berlinskii Huculak Apterae are matt brownish green to dark
green, siphunculi dark, cauda also dark but somewhat paler than siphunculi;
BL 1.1-1.4 mm. Alata has 6-8 secondary rhinaria on ANT III only. On Viburnum spp., forming colonies on
roots and underground runners, or tented over by ants. Only known from
Poland. Monoecious holocyclic, with oviparae and apterous males in late
August-September (original description). Aphis
berteroae Szelegiewicz Apterae have a dark green abdomen, rest
of body blackish green or blackish; BL 1.25-1.6 mm. Alata has 3-5 secondary
rhinaria on ANT III. In small, ant-attended colonies on roots of Berteroa incana, feeding on smaller
roots 30-50 cm below the surface. Only known from Poland. Monoecious
holocyclic, with apterous males (original description). Aphis
biobiensis Nieto
Nafría & Mier Durante Colour of apterae in life is unknown; BL
1.7-2.3 mm. Alatae have 6-9 rhinaria on ANT III and 0-1 on IV (0-3 on ANT III
in alatiform apterae). On Adesmia
viscida in Chile (Region VIII) (Nieto Nafría et al. 2016c). Aphis
blackmani Kadyrbekov Apterae are pale greenish with red eyes,
with only tips of appendages darkened; BL 1.7-2.2 mm. Feeding on the roots of
a species of Tragopogon in
Kazakhstan (Almaty region; Kadyrbekov 2019b). Other morphs and biology are
unknown. Aphis
boydstoni Pike Apterae are reddish brown to blackish,
with dark appendages; BL 1.6-2.9 mm. Alatae have secondary rhinaria
distributed III 22-31, IV 0-2. On Ceanothus
velitinus in open conifer forests in mountainous regions of north-west
USA (Washington, Oregon, Idaho, Nevada) and Canada (British Columbia).
Monoecious holocyclic with oviparae collected in September (Pike et al. 2004). Aphis
bozhkoae Eastop & Blackman (replacement
name for A. eupatorii Bozhko;
Eastop and Blackman 2005) Apterae are
yellowish with brown appendages; BL c. 1.9-2.0 mm. Alata has secondary
rhinaria distributed III 23-27, IV 12-15, V 4. On Eupatorium cannabinum in Ukraine (Bozhko 1976a, as A. eupatorii). Aphis
brachychaeta Holman Apterae are reddish brown; BL 1.5-2.2 mm.
Alata has secondary rhinaria distributed III 5-9, IV 0-2. On (probably) Stellera chamaejasme, colonising upper parts of roots and at bases of
stems, attended and sheltered by ants. In Mongolia. Presumed to be monoecious
holocyclic. Aphis
brachysiphon Narzikulov Apterae are dark brown; BL 1.3-1.7 mm. On
Clematis orientalis in Central Asia
(Iran, Tajikistan, Kazakhstan, Uzbekistan, Afghanistan; Remaudičre &
Remaudičre 1997). Generic placement is open to question; Kadyrbekov (2005b)
placed this species in Brachyunguis. Aphis breviseta
Holman Apterae are bright yellowish green; BL
1.1-1.9 mm. Alata has secondary rhinaria distributed III 3-6, IV 0(-1). On Potentilla spp., living on basal parts
of stems and rhizomes, under earthen sheaths built by ants. In steppe
localities in eastern Europe. Monoecious holocyclic, with apterous males
(original description). Aphis
brevitarsis Szelegiewicz Apterae are dark green with black
siphunculi and cauda; BL 1.8-2.1 mm. On stem of a Mentha sp. in Mongolia. Aphis
brohmeri Börner Apterae are dark purplish brown, BL
1.5-2.6 mm. Immatures have no discrete pleural wax spots (cf. fabae). Alata has secondary rhinaria distributed
III 9-18, IV 0-3. Colonies occur low down on stems and leaf sheaths of Anthriscus sylvestris, attended by ants.
There are also records from some other Umbelliferae/Apiaceae, e.g. Heracleum and Pastinaca (Chumak 2004), Carum
and Seseli (Ivanoskaya 1977),
and Aegopodium (Kadyrbekov 2012a),
although slide-mounted specimens may easily be confused with A. fabae, and it is possible that some
natural hybridisation occurs. Widely
distributed in northern and central Europe, and in Iran, western Siberia and
Kazakhstan. Monoecious holocyclic,
with apterous males (Heie 1986). Aphis
brotericola Mier
Durante Apterae are black, often
covered with whitish wax powder; BL 1.2-2.2 mm. Alata has secondary rhinaria
distributed III 5-16, IV 0-5. Forming dense colonies on leaves of Euphorbia spp. in Spain, Italy (Barbagallo & Patti 1998), France, Greece
(Papapanagiotou et al. 2012),
Turkey, Iran (Barjadze et al.
2017b) and Morocco. Monoecious holocyclic in Spain, with alate males (García
Prieto et al. 2001). Aphis
brunellae Schouteden Apterae are yellowish or whitish,
sometimes with a mid-dorsal suffusion of dark green (see influentialpoints.com/Gallery);
BL 1.0-1.6 mm. Alatae have secondary rhinaria distributed III 6-9, IV 2-5, V 0-1. On stems, in bracts of
inflorescences and on undersides of leaves of Prunella spp., attended by ants. Widely distributed in Europe,
and in western Siberia (Altai; Stekolshchikov et al. 2008a). Monoecious holocyclic, with apterous males;
oviparae have markedly swollen hind tibiae bearing c.100 scent glands. In
spite of clear biological differences, the
mitochondrial COI and COII sequence of a sample identified as A. brunellae in Italy was
indistinguishable from that of A.
gossypii (Cocuzza et al. 2009). Aphis
brunnea Ferrari Apterae are brown-black; BL 1.7-2.5 mm. Alatae with secondary rhinaria distributed ANT III
4-13, V 0-3. On shoot apices of Ononis spp. (especially natrix) in the Mediterranean area and adjacent countries; France, Spain,
Italy, Austria, Slovakia, Bulgaria, Rumania, Israel, Jordan and Lebanon. In
Spain it has been found on Anthyllis
cytisoides (Nieto Nafría et al.
2005). The life cycle is unknown, perhaps anholocyclic in southern Italy
(Barbagallo & Stroyan 1982). 2n=8. Aphis
bupleuri (Börner) Apterae are brown-black covered with
greyish-white wax powder; BL 1.3-1.9 mm. Alatae have
secondary rhinaria distributed III 17-41,
IV 6-15, V 3-10. On stems, lateral shoots and inflorescences of Bupleurum
spp. in Europe (France, Spain, Germany, Poland, Italy, Hungary, Czech
Republic, Rumania, Ukraine). One of a
group of closely related wax-covered Aphis
species on Bupleurum, the
others being caroliboerneri, funitecta and talgarica. Nieto Nafría et
al. (1986) tabulated morphological differences, and Kadyrbekov (2001a)
provided a key. Monoecious holocyclic with alate males, and oviparae with
strongly swollen hind tibiae bearing more than 200 scent glands. A. bupleurisensoriata Bozhko is a
synonym (Kadyrbekov 2001a). Aphis
cacaliasteris Hille Ris
Lambers Apterae are plump-bodied,
matt grey-black with black appendages; BL 2.0-3.1 mm. Apterae have secondary rhinaria,
distributed III 0-20, IV 7-15, V 0-3, and alatae have them distributed III
23-40, IV 12-20, V 0-3. On Senecio spp.
and related genera of tubuliferous Compositae (Adenostyles, Doronicum)
in south and central Europe, and recently recorded from western Siberia (Altai Republic; (Stekolshchikov &
Novgorodova 2013). Monoecious
holocyclic, with oviparae and alate males in July-September. Lampel (1984)
gave a full account. Aphis
calaminthae (Börner) Apterae are pale to dark green with
brownish head; BL 1.0-1.1 mm. Alatae have secondary rhinaria distributed ANT
III 5-7, IV 0-2. In loosely rolled leaves and inflorescences of Clinopodium
spp. formerly placed in Calamintha
and Acinos (acinos, menthifolium, grandiflorum,
sylvaticum), and also recorded from
Hyssopus officinalis. Jörg & Lampel (1988) noted differences in the
morphology of samples from Acinos
arvensis (= Clinopodium acinos)
and Hyssopus officinalis. Ant-attended. In southern Germany,
Austria, Switzerland, Hungary and Czech Republic. Lampel & Meier (2007)
recorded it also from Italy and Poland. Life cycle and sexual morphs are
unknown. Aphis
caliginosa Hottes & Frison Small, brownish aphids with dark legs and
antennae, in ant-attended colonies on ends of branches of Cornus racemosa in eastern North
America (Illinois, Pennsylvania, North Carolina, New Jersey). Life cycle is unknown (Robinson & Chen
1969). Aphis
callunae Theobald Apterae are small, plump-bodied, blackish,
brownish violet or dark brown, with wax powder on dorsum arranged in
hexagonal fields, and dark brown to black appendages (see influentialpoints.com/Gallery);
BL 1.0-1.5 mm. Immatures are greenish. Alatae have secondary rhinaria
distributed ANT III 3-7, IV 0-3. On terminal shoots and inflorescences of
old, straggly plants of Calluna
vulgaris, well-camouflaged and easily overlooked. In western, northern
and central Europe, and also recorded from British Columbia, Canada.
Monoecious holocyclic on Calluna,
with oviparae and alate males in September in northern Germany (alate,
brachypterous. and apterous males were obtained in culture; Müller 1976b). Aphis (Zyxaphis) canae Williams (Fig.12v) Apterae are green with darker head and
dorsal abdominal patch or mottling, and white wax markings including a
transverse band on anterior abdomen; BL c.1.6 mm. On stems and leaves of Artemisia
and Seriphidium spp. in western USA
(see Palmer 1952). There is some confusion in the literature
with A. hermistonii and A. oregonensis, but the three species
seem separable by the key characters given. Aphis
carduicola Stekolshchikov Apterae are bluish green with end of
abdomen yellowish green, or pale yellowish green; BL 1.1-1.9 mm. Alatae have
secondary rhinaria distributed III 3-7, IV 0-1. At base of stem and on
petioles of basal leaves of Carduus
spp. in southern Russia (Astrakhan province) and Georgia (Stekolshchikov
2005), and also recorded from Turkey (Şenol et al. 2014c). Monoecious holocyclic; fundatrices were collected
in April-May on C. uncinatus. Aphis cari Essig
Apterae are transparent white, yellow or pale yellowish green with
distinctive dark green dorsal markings, including a transverse bar between
the siphunculi; BL c.1.7 mm. Alatae have secondary rhinaria distributed III
15-50, IV 0-4. On stems of Carum
kelloggii, and also recorded from Angelica
tomentosa, in California, USA. Only known from the original description,
and it seems possible that these are hot weather dwarfs of Aphis carduella (q.v.). Aphis caroliboerneri (Remaudičre) Apterae are grey covered with dense waxy
secretion; BL 1.0-1.3 mm. Alatae have secondary rhinaria distributed III
7-16, IV 1-5, V 0-3. In small colonies on flowerheads of Bupleurum
fruticosum. Only known from France
and Spain.
Aphis
carrilloi Ortego, Mier Durante &
Nieto Nafría Apterae are dark green,
with varable dorsal sclerotisation; BL 1.5-2.8 mm. Monoecious holocyclic on Gunnera magellanica in Chile (Maule).
Fundatrices, apterous fundatrigeniae, apterous viviparae and one alate
vivipara were collected in January-February (Ortego et al. 2013). Aphis
carverae Hales, Foottit & Maw Apterae are black with antennae and legs
partly whitish; BL 0.9-1.5 mm. Alatae are unknown. In small colonies on
growing tips of Epilobium billardierianum in Australia (New South Wales).
Monoecious holocyclic, with oviparae and apterous males in March-April (Hales
et al. 2014) Aphis
caryopteridis Holman Apterae are dark green with dark
siphunculi and cauda; BL 1.0-1.5 mm. Alatae have 3-5 secondary rhinaria on
ANT III. On stems of Caryopteris mongholica, ant-attended.
In Mongolia. Aphis
catalpae Mamontova
Apterae are rich yellow to yellow-green, with a dark green spot on
dorsal abdomen in life; BL c.1.6-1.7 mm.
On undersides of leaves, causing strong wrinkling, and along petioles,
of Catalpa spp. (bignonioides, speciosa), in Ukraine; also reported from Hungary, shrivelling
leaves of C. bignonioides and
producing much honeydew (Ripka 2001), Italy (Cocuzza et al. 2009), Turkey, Iran, Uzbekistan (Davletshina 1964),
Kazakhstan (Kadyrbekov 2017a) and east Siberia (Pashchenko 1997). The far
eastern population is dark green and has longer siphunculi. Holocyclic on Catalpa with alate males (Davletshina
1964), but reported to migrate facultatively to herbs and grasses in Ukraine
(Mamontova 1953). [The host association is a mystery as the two Catalpa species from which it is
recorded are native to North America, yet the aphid is only recorded from
Eurasia. Populations on Catalpa in
Italy identified as this species have mitochondrial COI and COII sequence
data very close to that of A. gossypii
(Cocuzza et al. 2009).] Aphis
ceanothi Clarke Apterae are dull reddish or amber-brown,
with black head and black markings posterior to siphunculi; BL 1.0-2.3 mm.
Alatae have secondary rhinaria distributed III 11-36, IV 0-5, V 0-1. On
flower stems and leaves of Ceanothus spp.
in western USA and Canada; also found (in California) on Noltea africana (BMNH collection, leg. E.O. Essig). Monoecious holocyclic; oviparae and
apterous males were collected in September in Manitoba, Canada (BMNH collection,
leg. A.G. Robinson). Aphis celastrii Matsumura (synonym of spiraecola acc. to Moritsu 1983). Apterae are “dark brown with a greenish
shade” according to the original description. There is doubt about the
identity of this species. The name has since been applied to yellow-green
aphids on Celastrum, Deutzia, Ilex and Polygonum in
Japan (Higuchi & Miyazaki 1969) and Korea (Lee et al. 2002c) which, according to the description
given by Takahashi (1966), and slides in the BMNH collection determined as celastrii by Takahashi, are not
clearly distinct from A. spiraecola.
Oviparae were collected on Deutzia
by Takahashi (1966). 2n=8. Aphis
cephalanthi Thomas Apterae are dark purplish brown to
almost black, wax-dusted (see influentialpoints.com/Gallery); BL 0.9-1.7 mm. Alatae have secondary rhinaria
distributed III 9-15, IV 0-5. On leaves, stems and flowerheads of Cephalanthus occidentalis, widely
distributed in North America, and also in Cuba (as Schizaphis sp.; Holman 1974). A record from Impatiens balsamina in South Africa (Millar 1990) perhaps needs
additional confirmation. Well-developed marginal tubercles are present on
most segments and, atypically for an Aphis,
the tubercles on abdominal tergite 7 are posterior to, and on a level with,
the spiracles. Oviparae and alate males occur in late September (in New York;
BMNH collection, leg. L.L. Pechuman).
Aphis cephalariae Barjadze
Apterae are black except for pale ANT III, IV and basal half of tibiae; BL 1.6-2.7 mm.
Alatae have secondary rhinaria distributed III 19-31, IV 0-8, V 0-2. Living
in dense, ant-attended colonies on stems of Cephalaria gigantea in Georgia (Barjadze 2011b), and
now also recorded from Dipsacus sp. in Turkey (Özdemir & Barjadze 2015). Life cycle is
unknown but probably monoecious and holocyclic. Aphis cerastii (Börner) Apterae are dark grey, BL 1.2-1.6 mm. On
subterranean parts of Cerastium arvense in Germany. Males are
apterous.
Aphis
cercocarpi Gillette
& Palmer Apterae are shining
greenish black with antennae and legs mainly pale, and siphunculi and cauda
black; BL 1.3-2.0 mm. On leaves and twigs of
Cercopus montanus in western
USA (California, Colorado). Aphis (Toxoptera) chaetosiphon
(Qiao) Colour of apterae in life is
unrecorded; BL 1.7-2.0 mm. On new shoots of
Camellia oleifera, attended
by ants. In Guangxi Province, China (Qiao et
al. 2008). Aphis
chetansapa Hottes & Frison Apterae are yellowish-brown to
yellowish-red with darker antennae and legs and blackish siphunculi, and dark
markings on posterior abdominal tergites.
Alatae have only 4-8 secondary rhinaria confined to the third antennal
segment, dark wing veins, and often have dark tranverse bands across last
three abdominal tergites (Hottes & Frison 1931). In tightly curled terminal leaves of Pyrus sp. and Prunus sp., recorded from Illinois, Missouri and Oregon,
USA. The description of this species
closely resembles that of A. longicauda. Life cycle is unknown. Aphis
chilopsidi Davletshina Apterae are pale green to dark greenish
brown, wax-dusted, with dark siphunculi and pale cauda; BL c.1.6-1.7 mm. In
small colonies on leaves and petioles of Chilopsis
linearis in Uzbekistan. Aphis
chloris Koch Apterae are pale green or lemon yellow,
or in at lower temperature olive green, with dark head, apices of antennae
and legs, and siphunculi (see also influentialpoints.com/Gallery);
BL 1.0-1.9 mm. Alatae have 5-9 secondary rhinaria on ANT III and 0-2 on IV.
On Hypericum spp., usually living
in ant-attended colonies at stem bases just below soil surface, occasionally
on aerial parts. Widespread in Europe, across Asia to China, and introduced
for control of Hypericum perforatum to South Africa (Durr 1983), Australia
(Briese & Jupp 1995) and Canada (Foottit et al. 2006). Monoecious holocyclic, with apterous males. Wilson
(1938) made one of the earliest studies of induction of sexual morphs using
this species. Aphis citrina Nevsky,
described from Central Asia as a subspecies of A. chloris and subsequently given full species status, seems
likely to be a synonym. 2n=8. Aphis (Zyxaphis) chrysothamni Wilson (Fig.21j) Apterae have wine-coloured head and pale
green to pinkish abdomen with (usually) a dark spot anteriorly, and some wax
at the end; BL 1.0 (summer dwarfs) - 2.4 mm. Alatae have
secondary rhinaria distributed III 4-10, IV 0-5, V 0.1. On Chrysothamnus
nauseosus in western North America. Oviparae and apterous males appear in
late September in Utah (BMNH collection, leg. G.F. Knowlton).
Aphis (Zyxaphis) chrysothamnicola (Gillette
& Palmer) (Fig.21k) Apterae are dusky reddish brown or dull
blackish green with dark bands on posterior abdomen; BL 1.7-2.1 mm. Alatae
have 3-7 secondary rhinaria on ANT III. On leaves and stems of Chrysothamnus nauseosus in western USA
(Rocky Mountain region). Oviparae and apterous males in October. Aphis ciceri Müller (Fig.16h) Apterae are dark green, appearing black,
shiny on dorsum; BL 2.2-2.6 mm. On lower part of stem of Astragalus cicer, just above the soil surface, attended by ants. Only known from
Germany (Jena), Iran (Rezwani et al. 1994) and Kazakhstan (Kadyrbekov 2017a).
Oviparae and apterous males were obtained in September-October (original
description).
Aphis
cimicifugae Holman Apterae are grass green with black
siphunculi and cauda; BL 1.8-2.3 mm. Alatae have 3-10 secondary rhinaria on
ANT III. On Actaea (= Cimicifuga) spp., on upper parts of
stems and in inflorescences, attended by ants (Pashchenko 1994b). In east
Siberia. Aphis
cinerea Nieto Nafría & Ortego Apterae are black, thickly coated with
whitish grey wax; BL 1.7-2.5 mm. Alatae have 3-8 secondary rhinaria on ANT
III. In dense colonies on Lathyrus
macropus in Argentina (Nieto Nafría & Ortego 2002). Life cycle and
sexual morphs are unknown. Aphis
cirsiioleracei (Börner) Apterae are brownish grey, lightly
wax-dusted; BL 1.2-1.6 mm. On basal
parts, and later in upwardly rolled leaves of Cirsium spp., in central and eastern Europe (Germany,
Switzerland, Czech Republic, Poland; also in northern Italy), and also
recorded from Kazakhstan (Kadyrbekov 2014f), where there is a record from Cousinia microcarpa. Monoecious holocyclic according to Börner
(1952). It has been suggested that A.
viridescens Bozhko, described from Cirsium elodes in Ukraine is a synonym, although Bozhko’s species
was pale green in life. Aphis
cirsiphila Pashchenko Colour of apterae in life unknown; BL c.2
mm. In dense colonies on leaves and stems of Cirsium spp. in east Siberia. A record from Turkey (Görür et al. 2017) needs further
confirmation. Monoecious holocyclic. Aphis
cisticola Leclant & Remaudičre Apterae are yellow to bluish green, with
light dusting of wax, dark apices to legs and antennae, and black siphunculi
and cauda; BL 1.0-1.8 mm. Alatae have
3-7 secondary rhinaria on ANT III, 0(-2) on IV. On young growth of Cistus
spp., especially C. salvifolius,
forming dense colonies on leaves shoots and flower stems, attended by ants.
In southern France, Corsica, Italy and Spain. Barbagallo & Stroyan (1982) tabulated morphometric
data for comparison with A. lupoi. Aphis (Toxoptera) citricidus
(Kirkaldy) Tropical Citrus Aphid Plate 8c Apterae are
shining, very dark brown to black, usually larger than A.(T.) aurantii, and with antennae less distinctly banded (see aphids
of Karnataka website); BL 1.5-2.4 mm. Immatures are brown. Alatae have a
shiny black abdomen, black ANT III, forewing with pale pterostigma and media
twice-branched; they have secondary rhinaria distributed ANT III 6-20, IV
0-4. (Specimens in alcohol colour the fluid deep red.) In ant-attended
colonies on young growth of host plants, rolling leaves and stunting shoots.
Mainly on Rutaceae, especially Citrus,
but occasionally large colonies develop on young growth of other trees and
shrubs (see also Blackman & Eastop 2000, p. 355). In southern Africa,
southern Asia, Australia, New Zealand, Pacific islands and subtropical and
warm temperate parts of South America (but records from Chile seem to be
unfounded – see Nieto Nafría et al.
2015a). More
recently it has spread to important citrus-growing areas in Central America,
the Caribbean and southern USA (Yokomi et
al. 1994, Halbert & Brown 1996), and to Madeira, Portugal and north-western
Spain (Hermoso de Mendoza 2008), but not yet to the Mediterranean region and
Middle East. It thrives in moist warm climates and seems able to tolerate
colder conditions than A.(T.) aurantii,
for example occurring at higher altitudes, but it is not found in regions
with long hot dry seasons. Apparently it is anholocyclic
everywhere except Japan, where there is a sexual phase (Komazaki 1988). Michaud (1998) reviewed
the literature on this aphid, which includes laboratory studies of
population parameters (Galatoire 1983, Takanashi 1989); field studies in
Japan (Komazaki 1988) and Venezuela (Rondón et al. 1983 - including natural control by fungus and predators);
and a review of transmission of citrus tristeza virus (Roistacher &
Bar-Joseph 1987). An exotic aphidiid parasitoid
Lipolexis oregmae has been used for
biological control in Florida (Persad et
al. 2007). Qureshi (2010) examined the potential impact of
climate change on populations on citrus in Florida. Nieto Nafría et al. (2005b) established that citricidus is the correct specific epithet for this aphid. 2n=8. Aphis
clematicola Pashchenko Apterae are yellow-green, wax-dusted; BL
c. 1.5 mm. On Clematis fusca in
east Siberia, forming large dense colonies on undersides of leaves, attended by ants. Aphis
clematidis Koch Apterae are dark olive green with a
blackish spinal stripe, marbled with white wax powder; BL 1.7-2.3 mm.
Smaller, paler specimens may occur in midsummer. Alatae have 5-9 secondary
rhinaria on ANT III. On young shoots, flower stems and leaves of Clematis spp. in central and southern
Europe. Pigmentation and length of siphunculi show great variation, and there
has been confusion with A. vitalbae;
reliable differences from that species are the longer pointed hairs on the
appendages and the longer hind tarsal segment II (Tashev 1964b). Monoecious
holocyclic (Börner 1952). 2n=8. Aphis
clematiphaga Pashchenko Apterae are bright green and dark green,
without wax markings; BL c. 2.2 mm. On
stems, flower shoots and undersides of leaves of Clematis
spp. in east Siberia. Monoecious holocyclic. Visited by ants. Aphis
clerodendri Matsumura Apterae are straw-coloured to bright
yellow green or olive-green; BL 1.1-1.9 mm. On Clerodendrum spp. in east Asia (Japan, Korea, Vietnam, Laos,
Taiwan, Philippines, Papua New Guinea) and Australia (Carver et al. 2003), forming loose leaf pseudogalls. Alatae have 3-8 secondary
rhinaria on ANT III. Very close to A.
gossypii, and the characters that separate the two species in east Asia
may not be reliable for other parts of the world. Takahashi (1966) described
a subspecies, A. clerodendri amamiana, specific to C. yakusimense (= C.
trichotomum var. fargesii) in Japan, but this seems
indistinguishable from A. gossypii.
2n=8. Aphis
cliftonensis Stroyan Apterae are rather bright pale green; BL
0.8-1.2 mm. On roots of Helianthemum
nummularium, in ant shelters. In England and Sweden (BMNH coll. VFE).
Monoecious holocyclic with apterous males (Stroyan 1984). Aphis
clinepetae Pashchenko Apterae are pale yellow-green or dark
green, usually wax-dusted; BL c. 1.5 mm. Alatae have secondary rhinaria
distributed III 9-18, IV 2-6, V 0-2. In small, dense colonies on stems and
flowers of Nepeta manchuriensis, and more rarely on Clinopodium chinense, in east Siberia. [Aphids from Calamintha grandiflora in Crimea (BMNH
collection, leg. J. Holman) have apterae intermediate in morphology between
this species and clinopodii, but
include an alata with secondary
rhinaria distributed III 8 and 11, IV 3 and 3, so are possibly this species.]
Aphis
clinopodii Passerini Apterae are blackish green; BL 0.9-1.2
mm. Alatae have 2-5 secondary rhinaria on ANT III. Curling and distorting
terminal leaves of Clinopodium spp. in Europe (England,
France, Spain, Italy, Germany, Poland, Austria, Czech Republic, Ukraine). One
of a closely-related group of European species on Lamiaceae, and very similar
to A. serpylli (but with a longer
antennal terminal process) and A.
origani (but with shorter siphunculi). Monoecious holocyclic. Several
colonies in south-east England were all attended by the ant Myrmica sabuleti (Wood-Baker 1979). 2n=8. Aphis
clydesmithi Stroyan Apterae are dull green; BL 1.1-1.3 mm.
Alatae have 2-3 secondary rhinaria on ANT III. On Sanicula canadensis, living at base of stem and on root collar,
attended by ants. In North Carolina, USA. Aphis
coffeata Mamontova Apterae are coffee brown with a pale
cauda; BL c. 1.8 mm. Secondary rhinaria in alata (one specimen) distributed
III 12-13, IV 0-2. In colonies on stems and in flowerheads of Melampyrum nemorosum and M. pratense in Ukraine, Byelorussia
and western Russia. It was redescribed
by Stekolshchikov et al.
(2007b). Aphis comari
Prior & Stroyan Apterae are blue-green, very dark green
or bluish black; BL 1.0-1.9 mm. Alatae have 4-8 secondary rhinaria on ANT
III. On shoots or leaves of Comarum palustre growing in sphagnum
bogs, sometimes in ant shelters built from plant debris. In northern and
north-west Europe (UK, Denmark, Finland, Kola Peninsula, Poland), and also
recorded from Kazakhstan on Sibbaldianthe
(= Potentilla) bifurca and Potentilla ?virgata (= dealbata) (Kadyrbekov 2009b, 2014d,
2017a). Monoecious holocyclic, with apterous and brachypterous males
(original description). Aphis
commensalis Stroyan Apterae 1.4-1.7 mm, dark grey-green to
blackish-green, heavily clothed in mealy greyish wax powder (see
influentialpoints.com/Gallery), living in late spring and summer within empty leaf-edge pocket-galls of
the psyllid Trichopsylla walkeri on
leaves of Rhamnus cathartica in
Europe, and also in Kazakhstan (Kadyrbekov 2017a). Earlier in the year fundatrices amd
fundatrigeniae cause rolling and twisting of leaves (see Pérez Hidalgo et al. 2010). Small apterous males and
oviparae mature within the gall in September-October, and eggs are laid
within the gall (Stroyan 1952, or for a fuller account, Böhm 1963). Alatae
are unknown. Aphis comosa
(Börner) Apterae are brown with shining black
dorsum, antennal flagellum and middle sections of tibiae pale; BL 1.5-2.4 mm.
Alatae have 4-9 secondary rhinaria on ANT III. On Melilotus albus and Lathyrus
pratensis in Europe (France,
Germany, Austria, Italy, Spain). Also recorded from Latvia, Poland and
Romania by García Prieto et al.
(2004), but their record of its occurrence in UK appears to be erroneous.
Some records may apply to A.
pseudocomosa (q.v.). Redescribed by Hoffmann (1972). Aphis
conflicta Nieto Nafría, Ortego & Mier
Durante Apterae are shing black or
dark reddish brown, with some whitish wax powder, antennae and tibiae being
predominantly yellow; BL 1.2-2.3 mm. Monoecious holocyclic on Ochetophila (= Discaria) trinervis in
Argentina, and also recorded from species in several other genera of
Rhamnaceae (Colletia, Condalia, Discaria, Trevoa), and
also on Tristerix (Loranthaceae),
in Chile and Argentina. Nieto Nafría et
al. (2008) described all morphs including fundatrices (collected in
November), oviparae and apterous males (early April). A sample collected in
January at Neuquén (Argentina) included small (aestivating?) apterae. López
Ciruelos et al. (2017b) reviewed
morphological variation, host plants and distribution. Aphis
confusa Walker Apterae are pale yellow, yellowish green,
green or dark green, depending on location on host; BL 1.0-2.3mm. Alatae have
secondary rhinaria distributed III 3-7, IV 0-1. On Dipsacaceae (Dipsacus, Knautia and Scabiosa,
occurring both as larger green or dark green aphids on upper parts of stems
and inflorescences, or in summer as smaller and paler yellow or yellower
green specimens on undersides of leaves, at base of stems or on roots (Heie,
1986). Muir (1959) recorded it from Succisa
pratensis. Ant-attended. Widely distributed in Europe, in Central Asia
(Kadyrbekov 2017a), and Remaudičre et
al. (2006) reported its occurrence in Turkey, Iran and Mongolia.
Monoecious holocyclic, with apterous males. Kanturski & Bezdĕk
(2019) provided a detailed redescription including sexual morphs. A very
variable species, and there is no reliable way of distinguishing it
morphologically from the equally variable polyphagous A. gossypii, which may sometimes occur on Dipsacaceae. A. gossypii is, however, unlikely to
occur on basal parts of the plant, and is less assiduously ant-attended. In A. gossypii there is also usually more
contrast between the pigmentation of siphunculi and cauda. Aphis
conspicua Nieto Nafría & Mier
Durante Apterae are matt yellowish
grey to matt black; BL 1.1-1.9 mm. In compact groups on stems of Baccharis spp. (linearis, salicifolius).
Alatae seem rare; the one known specimen has 6-7 secondary rhinaria on ANT
III and 0-1 on IV. In Argentina (Neuquén) and Chile (Maule). Sexual morphs
are unknown (Nieto Nafría et al.
2019c). Aphis
coprosmae Laing Apterae are in life covered in fluffy
white wax); BL c. 1.6 mm. Alatae are remarkable in having only 0-1 secondary
rhinaria on ANT III. On Coprosma spp.
in New Zealand (Cottier 1953, Teulon et
al. 2013). Monoecious holocyclic; eggs are laid on stem of plant and
hatch in September. DNA analysis has revealed an nnusually high level of
genetic diversity for a single species (Teulon et al. 2013). Aphis
coreopsidis (Thomas) Plate 6c Apterae 1.5-1.8 mm, yellow to green with
darker antennae, legs and siphunculi (see
influentialpoints.com/Gallery),
on stems and leaves of new shoots of Nyssa
sylvatica in spring. Apparently
host-alternating in North America (Illinois) between Nyssa and secondary hosts in Compositae/Asteraceae (Bidens, Clibadium, Eupatorium, Sonchus), Malvaceae (Hibiscus, Sida) and Lamiaceae (Blephilia,
Nepeta). Oviparae in autumn on Nyssa are small (1.1-1.2 mm), yellowish-green; males are alate,
about 1 mm, dark green and brown (Hottes & Frison 1931). The fundatrices on the primary host are
still undescribed. Presumed
anholocyclic populations occur on secondary hosts (especially Bidens pilosa) in central and South
America and Hawaii, and alatae identified as this species have been trapped
in Ghana, Uganda (BMNH collection) and Saudi Arabia (Hussain et al. 2015a). Aphis
coridifoliae Mier Durante
& Ortego Apterae are pale or
greenish yellow powdered with white wax, with brown-black siphunculi and
cauda, and frequently with dark brown intersegmental markings on dorsal
abdomen; BL 1.0-1.7 mm. Alatae have 4-8 secondary rhinaria on ANT III. In
small dense colonies on Baccharis
coridifolia, and perhaps on other
Baccharis spp., in Argentina (Mier Durante & Ortego 1999). Aphis
cornifoliae Fitch Apterae are small (1.0-1.4 mm), dark brown
to dull greenish-black, on both upper and undersides of leaves of Cornus spp., living without host
alternation; males are apterous (Palmer 1952). Widely distributed in USA and Canada. Robinson & Chen (1969) discussed
differences from A. carduella (as helianthi). A member of the asclepiadis group (Lagos et al. 2015). Aphis
cornuta Blackman & Brightwell Apterae are bright yellow to pale yellow-green with pale green head and
prothorax, black siphunculi and a dusky cauda (see
influentialpoints.com/Gallery); BL 0.9-1.8
mm. living without host alternation at stem bases of Picris (= Helminthotheca)
echioides, attended by ants.
Oviparae and apterous males were collected in September (Blackman et al. 2019a). Only known from
southern England. Aphis
coronillae Ferrari Apterae are dark brown to brownish green
with grey-black dorsal shield (not wax-powdered; see
influentialpoints.com/Gallery); BL 1.3-2.2 mm. Alatae have secondary
rhinaria distributed III 3-9, IV 0-2. Living on basal parts of certain
Leguminosae/Fabaceae (Coronilla, Glycyrrhiza, Securigera, Trifolium, Medicago), sheltered by ants. Widely distributed in
Europe, and also in west Siberia (Stekolshchikov et al. 2008a) and Kazakhstan (Kadyrbekov 2017a). Monoecious
holocyclic, with males (mostly apterous, occasionally brachypterous or alate)
and oviparae appearing in September.
Hoffmann (1968) studied host relationships and described populations
on Medicago lupulina as a
subspecies (A. coronillae arenaria Hoffmann, as scaliai ssp. arenaria).
Stroyan (1984) discussed morphological variation in relation to geography and
host plant. A. yangbajaingana Zhang,
in Tibet on Medicago, is very
similar but has shorter, thicker siphunculi. Aphis
coronopifoliae Bartholomew Apterae are green, BL 1.4-1.7 mm. Alatae
have secondary rhinaria distributed III 11-17, IV 2-7. On Cotula
coronopifolia in California, USA. It seems likely, as suggested by Cook
(1984a), that this species is A. mimuli (q.v.). Aphis (Bursaphis) costalis Cook Colour of aptera in life unknown,
probably pale green or yellowish green; BL 1.5-1.8 mm. On Mimulus spp. in California, USA (Hille
Ris Lambers 1974b, as mimuli Oestlund).
It is a member of the oenotherae group,
which typically host-alternate between Ribes
and Onagraceae. An aphid in central Washington identified as this species
is believed to migrate from Ribes
aureum to Mimulus and Veronica,
but there are possibly other similar species in that region, including one
that is monoecious on Ribes cereum (see
aphidtrek.org). Aphis
cottieri Carver Apterae are brown-black, shining; BL
2.2-2.7 mm. Alatae are without secondary rhinaria. On leaves and stems of Muehlenbeckia spp. in New Zealand.
Oviparae in April-June, males are unknown (Carver 2000, Teulon et al. 2013). Podmore et al. (2018) reported on genetic
variation in mitochondrial COI sequence. Aphis coweni
Palmer Apterae are lead-grey to greenish black;
BL 1.4-2.0 mm. Alatae have secondary rhinaria distributed III 15-22, IV 2-7.
On leaves of Veratrum spp. (californicum, viride) in western North America, and apparently also on various Veratrum spp. in east Siberia
(Pashchenko 1988a, as A. veratri). Aphis craccae L. Apterae are black, covered in grey wax
powder (see influentialpoints.com/Gallery); BL 1.9-2.8 mm. Alatae
have 5-13 secondary rhinaria on ANT III. Living in often dense, ant-attended
colonies on terminal growth, flowers and seed-pods of vetches (Vicia spp., especially V. cracca). Records from other hosts may be based on confusion with other
“black-backed” Aphis species. Widely distributed in Europe,
eastward to China, Japan and Korea, and also in north-eastern USA and Canada.
Oviparae and alate males appear in September (Heie 1986).
Aphis
craccivora Koch Plate 6d (Fig.41k) Apterae are shining black, with immatures
lightly dusted with wax, usually ant-attended (see influentialpoints.com/Gallery); BL 1.4-2.2 mm. Alatae have
2-10 secondary rhinaria on ANT III. Colonising
young growth of numerous plants, particularly Leguminosae/Fabaceae; plants in
other families tend to be colonised more in the dry season. A major pest of
leguminous crops (Blackman & Eastop 2000, p. 227). Worldwide, but
particularly common in warm temperate and tropical regions. Anholocyclic
almost everywhere, but a sexual phase with alate males has been reported from
Germany, India and Argentina. Rhopalosiphum
esculentum Raychaudhuri & Raychaudhuri, described from Manihot esculenta in southern India,
is a possible synonym (see also Skvarla et
al. 2019). Wongsa et al. (2017)
have studied variation in COI sequence of populations of A. craccivora on leguminous crops in Thailand. A long-haired variant on Robinia in east Asia has been distinguished as a subspecies, A.
craccivora ssp. pseudoacaciae Takahashi (1966). Mehrparvar et al. (2012) found morphometric
differences between populations in Iran on different host plants, a
population from Robinia again being
the most distinct. The
discovery that populations on Medicago and
Robinia have different
endosymbionts (Brady & White 2013) provides further confirmation that two
distinct taxa are involved. Mitochondrial DNA (COI) analysis of numerous
samples of A. craccivora group by
J. Wang et al. (2011) also supports
the existence of a separate taxon on Robinia.
All 37 mitochondrial genes of the form on Robinia
have been sequenced (Song et al.
2016). [If populations on Robinia
need a new name then A. robiniae
Macchiati 1885 is available.] 2n=8. Aphis
crassicauda Smith &
Eckel Apterae are reddish tan to
light brown, dusted with wax; BL 1.1-2.1 mm.
Alatae
have 7-14 secondary rhinaria on ANT III. On
Viburnum spp. in North Carolina,
USA. Monoecious holocyclic, with oviparae and apterous males in
September-October (original description). Aphis
crepidis (Börner) Apterae are dark bluish green to yellow-green,
not wax-powdered (see influentialpoints.com/Gallery); BL 1.15-1.95 mm. Alatae have 3-7 secondary
rhinaria on ANT III. On Crepis
spp., living in ant shelters at base of plant, and also once found on Picris echioides (in Italy: BMNH
collection, leg. H.L.G. Stroyan and D. Hille Ris Lambers). In west, central and south Europe,
including Iberian peninsula, and also recorded from Iran (Mokhtari et al. 2012). Monoecious
holocyclic; oviparae in September, males apparently undescribed. 2n=8. Aphis
crinosa (Paik)
Apterae yellowish brown with white wax secretion (Shinji 1923, as Pterocomma ligustri); BL 2.3-2.7
mm. On trunks and branches of Ligustrum spp., and also recorded from
Lonicera and Viburnum. In Japan and Korea (Lee et al.
2002). Aphis
curtiseta Holman Apterae are green to dark green; BL
1.0-1.7 mm. On Hieracium spp.,
living mostly on basal parts of plants, and in early summer on
inflorescences. Ant-attended. In Austria, Czech Republic, Slovakia, Bulgaria,
Moldova and Ukraine. Monoecious holocyclic, with apterous males (Holman
1998). Aphis
cuscutae Davis Apterae pale green, thinly wax-dusted; BL
1.5-2.1 mm. Alatae have antennal segment III swollen with numerous
tuberculate secondary rhinaria. Described from Cuscuta epithymum growing on alfalfa in Utah, USA, and also found
on Cuscuta sp. growing on Rhus laurina in California (BMNH
colln, leg. R.C. Dickson). Aphis cuyana
López Ciruelos &
Ortego Apterae
are caramel to brown with “dorsal bright spots” (absent in midsummer dwarfs);
BL 1.3-2.3 mm. On Gymnophyton
polycephalum in Argentina (López
Ciruelos et al. 2017a). Alatae have
secondary rhinaria distributed ANT III 6-12, IV 0-4.
Aphis
cytisorum Hartig Apterae are black, dark brown or very
dark green, usually densely covered in grey wax powder, ant-attended (see influentialpoints.com/Gallery);
BL 1.5-2.8 mm. On young shoots and petioles, and later on
inflorescences and developing seed pods, of woody
Leguminosae/Fabaceae (Laburnum, Cytisus, Spartium, Sophora, Calicotome). In Europe eastward to Russia and Turkey,
Iran, North Africa, China (as A. sophoricola Zhang); also widespread in North America, and recorded
from Peru (Vilca Mallqui & Vergara Cobián 2011) and Argentina (see Mier Durante et al. 2012). Alatae have secondary
rhinaria distributed III 4-9, IV 0, V 0 in A. cytisorum s. str., and III 5-9, IV 0-2, V 0-1 in A.
cytisorum ssp. sarothamni Franssen,
which occurs on Cytisus scoparius in
Europe. Stroyan (1984) compared morphometric parameters of the two subspecies. Apterae are almost
impossible to separate, but oviparae of ssp. sarothamni have more swollen hind tibiae than cytisorum, with many more scent glands.
Males are alate. There is a complex of closely-related species on woody
Leguminosae/Fabaceae in Europe, including
others with very similar apterae on Cytisus
and Chamaecytisus (A. pseudocytisorum, A. zweigelti). 2n=8 (both subspecies). Aphis
danielae Remaudičre Apterae are black; BL 1.6-2.5 mm. On Lycium spp. in Chile and Argentina. In
Chile colonies also occur regularly on flower buds of a cactus, Echinopsis chilensis (Heie et al. 1996). Monoecious holocyclic,
with oviparae and apterous males in April (Remaudičre & Ortego 1998). Aphis
dasiphorae Holman Apterae are reddish brown, powdered with
greyish wax; BL 1.2-2.4 mm. Alatae have 4-10 secondary rhinaria on ANT III.
In dense colonies on terminal parts of shoots of Potentilla (as Dasiphora) spp. in eastern Russia, Kazakhstan
(Kadyrbekov 2012a) and Mongolia. Apparently monoecious holocyclic. Aphis
decepta Hottes & Frison Apterae are yellow with dark dorsal
markings posterior to siphunculi, dark brown siphunculi and a brown cauda; BL
1.4-2.2 mm. Alatae have secondary rhinaria distributed III 37-48, IV 0-4. On
undersides of leaves of Pastinaca sativa (a European plant) in
north-eastern USA (Illinois, New York, Pennsylvania). Native hosts are
probably Heracleum spp.; it is
recorded from H. lanatum in Manitoba, Canada
(Rojanavingse & Robinson 1976) and from H. maximum in Illinois (Lagos et
al. 2014). A member of the asclepiadis
group. Aphis
dianthiphaga Pashchenko Apterae are dark emerald green, BL
c.1.4-1.5 mm. In small to medium-sized colonies feeding at bases of stems of Dianthus repens in east Siberia. Aphis diluta
Pashchenko Apterae are pale or bright green; BL
c.1.8 mm. Alatae have secondary rhinaria distributed III 11-25, IV 9-17, V
5-12. On stems and under leaves of Potentilla
supina, forming large, loose colonies, and sometimes causing leaf
deformation. Possibly also on Geum
aleppicum. In east Siberia. Aphis
dlabolai Holman Colour of apterae in life unknown, probably
black dusted with whitish wax; BL 1.8-2.4 mm. Alatae have 6-13 secondary
rhinaria on ANT III only. On Euphorbia sp. in Iran. Aphis
droserae Takahashi Apterae are greenish black, with
appendages mainly dark; BL 1.0-1.2 mm. On flowers and undersides of leaves of
Drosera spp. in China and Taiwan
(Tao 1991), and now recognised to occur also in southern Germany (Barjadze et al. 2017b). Aphis
duckmountainensis Rojanavongse
& Robinson Apterae are black, BL
2.0-2.4 mm. Apterae have secondary rhinaria distributed III 3-26, IV 10-20, V
0-6, and alatae have them distributed III 44-53, IV 14-22, V 3-8. On terminal
growth of Senecio pauperculus (= Packera paupercula) in Manitoba, Canada. Aphis
ecballii Rusanova Apterae are yellow-brown or dirty yellow,
with dark siphunculi and yellow cauda; BL 1.9-2.1 mm. Alatae have secondary
rhinaria distributed III 8-13, IV 1-3. In colonies on leaves and shoots of Ecballium in Azerbaijan. Probably a
member of the frangulae group; the
description does not allow apterae to be differentiated from those of A. gossypii. Aphis egomae
Shinji Apterae are pale yellow to yellow-green
or orange, with black siphunculi; BL small, unrecorded. Alatae have 3-6
secondary rhinaria on ANT III. On Perilla
frutescens and Salvia splendens,
feeding on undersides of young leaves and causing severe leaf-curl. Moritsu
(1983) also recorded it from Physostegia
sp. In Japan, and also in South Korea (Lee et al. 2002c). Aphis
elatinoidei Nevsky Apterae are pale green, BL 1.2-1.6 mm. On
Elatinoides elatine in Uzbekistan
(see also Davletshina 1964). Aphis
elegantula Szelegiewicz Apterae are green mottled with darker
green, with black siphunculi and cauda; BL 1.75-2.1 mm. Alatae have secondary
rhinaria distributed III 3-6, IV 0-1, V 0-1. On Polygonum angustifolium in Mongolia (Szelegiewicz 1964a), and
also recorded from south-west Siberia (Kurgan province; see Novgorodova &
Stekolshchikov 2013). Aphis elena Lagos-Kutz & Voegtlin Apterae are dark green, dusted with white wax;
BL 1.3-1.6 mm. Alatae have secondary rhinaria distributed ANT III 5-6, IV
1-2. On Pycnantherum virginianum in
Illinois, USA (Lagos-Kutz et al. 2017). Very similar in morphology to A. gossypii, but DNA analysis indicates a closer relationship
with another North American Lamiaceae feeder, A. monardae. The life cycle is unknown. Aphis (Bursaphis) epilobiaria Theobald Apterae are reddish brown to very dark
blackish brown or blackish green, with a striking pattern of dense pleural
wax bands across the dorsum, except for a spindle-shaped mid-dorsal clear
area (see influentialpoints.com/Gallery);
BL (1.3)-2.1-2.7 mm. Alatae have secondary rhinaria distributed III 7-13, IV
2-8, V 0-2. On shoot apices and inflorescences of Epilobium hirsutum, and occasionally other Epilobium spp., in Europe (Stroyan 1984; Heie 1986), and also
reported to occur in Turkey, Iran, Kazakhstan and Pakistan. Monoecious holocyclic, with oviparae and
alate males in September. Rakauskas (2008) found an Oenothera sp. colonised by A.
epilobiaria including sexual morphs and eggs in autumn in the Czech
Republic, but fundatrices hatching the following spring did not complete
their development when reared on Oe.
biennis. 2n=8. Aphis (Bursaphis) epilobii Kaltenbach Apterae are brown or greenish black,
rather more uniformly wax-powdered than A.
epilobiaria (see influentialpoints.com/Gallery), and smaller; BL 1.3-2.1
mm. Alatae have secondary rhinaria distributed III (5-)9-20, IV 2-10, V 0-4. On upper
parts of stems, flowerheads and undersides of upper leaves of Epilobium montanum, and more rarely other species such as E.
lanceolatum. Usually without ant
attendance. Widespread in Europe, also reported from Iran, Kazakhstan
(Kadyrbekov 2017a), and introduced to North America (Foottit et al. 2006). Monoecious
holocyclic, with oviparae and alate males in September. 2n=8.
Aphis epipactis Theobald Apterae are very dark green, almost sooty black; BL c.1.3-1.7 mm. On Epipactis spp. in Europe. There are no satisfactory distinguishing characters, and possible synonymy has been suggested with A. fabae (Börner 1952) or, more likely because of the long hairs and frequent presence of marginal tubercles on abdominal tergites 2-4, A. ilicis (Eastop & Hille Ris Lambers 1976) or A. viburni (Stroyan 1984); Epipactis possibly acts as a reserve host for various members of the A. fabae group. Aphis equiseticola Ossiannilsson Apterae are light green or dark green, BL 1.4-2.0mm. Alatae have secondary rhinaria distributed III 8-11, IV 0-5, V 0. On Equisetum silvaticum in northern Europe (Sweden, Germany). Presumably monoecious holocyclic, with alate males in early August (Heie 1986). Aphis
(Pseudoprotaphis) erigerontis Holman Apterae are dark green dusted with bluish
grey wax; BL 1.5-1.9 mm. Alatae have secondary rhinaria distributed III 27-32,
IV 10-15, V 7-13, VI base 3-4. In ant shelters on root collar and roots of Erigeron acer in Czech Republic, and
found on Conyza canadensis in
Poland (Osiadacz & Hałaj 2017); also reported to occur in Finland,
Poland, Slovakia, Spain (García Prieto et
al. 2004), Iran (Rezwani 2010) and Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic, with apterous males
(original description). Aphis eryngiiglomerata Bozhko Apterae are greenish black, dusted with wax powder; BL 1.6-2.2 mm. Alatae have secondary rhinaria distributed III 5-15, IV 0-3. In ant-attended colonies on Eryngium spp. in eastern Europe (Czech Republic, Poland, Ukraine, Turkey), Kazakhstan (Kadyrbekov 2017a) and south-west Siberia. Aphis esulae (Börner) Apterae are pale brown, sometimes blackish dorsally especially behind and between siphunculi; BL 1.8-2.3 mm. Alatae have 9-12 secondary rhinaria on ANT III. On stems and inflorescences of Euphorbia esula, and possibly on other Euphorbia spp., in Netherlands (Barjadze et al. 2017b), Austria, Bulgaria, Hungary, west Siberia (Stekolshchikov et al. 2008a), Turkey (Akyürek et al. 2019) and Kazakhstan (Kadyrbekov 2011b). A single aptera from northern England (BMNH collection, leg. Airy Shaw) also appears to be this species. A member of the difficult A. euphorbiae group, distinguished in Börner’s very brief original description by its short hairs and association with E. esula. Szelegiewicz (1962b) provided a redescription on the basis of specimens from Bulgaria, and his interpretation is followed here. However, populations of A. euphorbiae group aphids with short hairs are not restricted to E. esula, and this plant may also be colonised in western Europe by aphids with typical A. euphorbiae morphology, so further work is needed. Aphis etiolata Stroyan Apterae are pale yellowish , whitish or bluish green; BL 1.4-2.0 mm. Alatae have 3-5 secondary rhinaria on ANT III. On roots of Rumex acetosella in ants’ nests, rather deep in soil. In Europe (England, Netherlands, Sweden, Finland, Poland, Czech Republic, Slovakia). Monoecious holocyclic, with apterous males. Aphis eucollinae López Ciruelos &
Ortego Apterae are yellow, sometimes
with brown dorsal markings, and with appendages and siphunculi brownish; BL
1.3-2.3 mm. Alatae have 5-9 secondary sensoria on ANT III only. On Euphorbia collina (= portulacoides ssp. collina) in Argentina.
Monoecious holocyclic, with oviparae and apterous males in April (López Ciruelos et al.
2016).
There is a close morphological similarity to the palaearctic species A. esulae. Aphis eugeniae van der Goot Apterae are orange-yellow to brownish-orange with black siphunculi and cauda; 1.3-2.1 mm. Alatae have 5-10 secondary rhinaria on ANT III, and 0-1 on IV. Resembling A. spiraecola, but may be distinguished by the characters given in the key to polyphagous aphids, and also by the presence of modified, peg-like hairs on the hind tibiae. Most commonly on Euphorbiaceae, with a preference for woody species (Glochidion, Breynia, Phyllanthus), but plants in other families may be colonised, e.g. Compositae/Asteraceeae (Chromolaena, Clibadium), Convolvulaceae (Convolvulus), Dipsacaceae (Dipsacus), Malvaceae (Hibiscus), Myrtaceae (Eugenia) and Rosaceae (Pyrus, Eriobotrya). Throughout east and south-east Asia (westward as far as Pakistan; Naumann-Etienne & Remaudičre 1995), in Australia (Eastop 1966, as A. hardyi). It is also now in Hawaii (Foottit et al. 2012) and New Caledonia (Mille et al. 2020), and (since 2011) in Florida, USA, where large damaging colonies are reported on Eriobotrya japonica (Skvarla et al. 2017). A record from Tunisia (Ben Halima-Karmel 2012) needs further confirmation. Calilung (1976) gave an account of this species in the Philippines. Pseudotoxoptera pyrisucta Qiao & Zhang, described from Pyrus pyrifolia in China (G. Zhang et al. 1999d), is likely to be a synonym. 2n=8. Aphis
evonymi Fabricius
Apterae are various shades of chocolate brown, often with transverse
bars of wax dust on anterior abdominal tergites (see
influentialpoints.com/Gallery);
BL 1.7-2.9 mm. Alatae have secondary rhinaria distributed ANT III
18-28, IV 0-7, V 0. Monoecious and holocyclic in Europe on
Euonymus europaeus, curling the
leaves. It is also recorded from east Siberia, on E. maximowiczianus (Pashchenko 1997). Males, and usually the gynoparae also, are
apterous. Hybridisation with A. fabae occurs readily in the
laboratory, and may occur in the field, so subspecies status might be more
appropriate for this aphid (Müller & Steiner 1986). Heie (1986) provided a general account, and
discussed the confused application of this name (as A. euonymi; note that correct spelling is evonymi) in the literature. 2n=8. Aphis eupatorii Passerini Apterae are dark green, or whitish yellow in midsummer dwarfs; BL 0.9-2.0 mm. Alatae have secondary rhinaria distributed III 4-10, IV 0-4. In ant-attended colonies on young growth and undersides of leaves of Eupatorium cannabinum in Italy. Monoecious holocyclic, fundatrices having been found in early June (Barbagallo & Stroyan 1982). A member of the frangulae group, close to frangulae s. str. (q.v.) which, according to Börner (1952), has Eupatorium cannabinum as a secondary host in Germany. No reliable morphological discriminants are available. Aphis euphorbiae Kaltenbach Apterae are blackish brown, shiny or wax-dusted, with tibiae pale brown except at apices, and siphunculi and cauda black (see influentialpoints.com/Gallery); BL 1.3-2.4 mm. Alatae have secondary rhinaria distributed III 7-18, IV 0-5. On upper parts of stems of Euphorbia spp., especially (in Europe) E. cyparissia. Sexual morphs have not been described, and it seems to be mainly or entirely anholocyclic. In Europe (but rare in north, and apparently not in Iberian peninsula – see García Prieto et al. 2004); in western Siberia, south-west and central Asia, and introduced to USA (California, but an early record from Nebraska is not this species; see Foottit et al. 2006). A member of a taxonomically difficult group of closely-related species on Euphorbia. Records of A. euphorbiae from Africa and Australia, as well as many of those from southern Europe and the Middle East, seem to be referable to A. tirucallis. Aphis euphorbicola Rezwani & Lampel Apterae are dark brown or greyish brown, lightly wax-dusted; BL 1.0-1.7 mm. Alatae have secondary rhinaria distributed III 6-12, IV 1-2. On undersides of leaves and shoots of Euphorbia denticulata in Iran. Aphis exploratus Rusanova Apterae are greenish yellow or dark orange, siphunculi yellowish with darker apices, cauda yellow; BL 1.5-1.9 mm. Alatae have secondary rhinaria distributed III 6-8, IV 0-3. On leaves, flowers and fruits of Paliurus in Azerbaijan. Possibly this was A. nasturtii. Aphis exsors Rusanova Apterae are brown or green, with dark siphunculi and cauda; BL 1.6-2.3 mm. Alatae have secondary rhinaria distributed III 9, IV 3-4, V 2. Described as occurring on various Compositae/Asteraceae (e.g. Conyza canadensis, Dahlia, Sonchus) in Azerbaijan. This species could not be included in keys on the basis of the published description. Aphis
fabae Scopoli Fig. 4 Plate 6e Apterae are dull black,
sometimes with white wax markings, usually ant-attended (see influentialpoints.com/Gallery);
BL 1.5-3.1 mm. Alatae have secondary rhinaria
distributed III (3-)7-33, IV 0-14, V 0-3. Curling leaves
of Euonymus europaeus in spring
(also Philadelphus coronarius
and/or Viburnum opulus, depending
on subspecies – see below), and migrating to a wide range of secondary hosts,
including young growth of some trees, and many crops. In Europe there seems to be a complex of
sibling species or partially interfertile subspecies, of which four are known
to overwinter on Euonymus; they are
A. fabae
sensu stricto (which in host
transfers will colonise Vicia faba),
A.
fabae cirsiiacanthoidis Scopoli (which will transfer to Cirsium arvense), and two that are
dealt with here as separate taxa; A.
solanella (which will transfer to Solanum
nigrum) and A. evonymi, which
stays through the summer on Euonymus.
A. fabae sensu stricto, A. fabae cirsiiacanthoides and A. solanella will also sometimes use Viburnum opulus as their primary host.
Another
closely related form, A. fabae mordwilkoi Börner &
Janich, has a sexual phase on Viburnum opulus, or sometimes Philadelphus
coronarius, but
not on Euonymus, and migrates particularly to Arctium spp. and Tropaeolum majus. This subspecies also occurs in Canada, where
it was studied (as barberae
Robinson) by Robinson (1980) and Barber & Robinson (1980), and has a
sexual phase on Viburnum trilobum.
Oviparae on primary hosts in autumn are small, with strongly swollen hind
tibiae, and males are alate.
Anholocyclic populations of aphids of the A. fabae group (especially its short-haired close relative A. solanella) occur on secondary hosts
in southern Europe, south-west Asia, Africa, Indian subcontinent, Korea (Kim et al. 2006b), South America, Hawaii
and Auckland Isles. Accounts of this
group in Europe were given by Stroyan (1984), Heie (1986) and Müller (1988),
and Blackman & Eastop (2007) discussed the taxonomic problems.
Hybridisation and reproductive isolation between members of the complex have
been studied by Thieme & Dixon (1996), Raymond et al. (2001) and Tosh et
al. (2004), and Görür et al. (2005) studied the role of phenotypic
plasticity in host race formation and speciation. Béji et al. (2015) studied
mitochondrial DNA (COI) variation in A.
fabae-group populations in Tunisia. Sandrock et al. (2011) conducted microsatellite DNA analyses of A. fabae s. str. collected at
different latitudes in Europe and demonstrated a relationship between
climate, population structure and life cycle. See also Blackman & Eastop
(2000, 2006). 2n=8 (all subspecies). Aphis
farinosa Gmelin
Apterae 1.6-2.5 mm, dull pale to mid green with cauda distinctly
darker than the long pale siphunculi, in dense colonies on young shoots of Salix spp. in spring and early summer,
attended by ants (see influentialpoints.com/Gallery). Alatae
have secondary rhinaria distributed III 5-11, IV 0-2. Monoecious,
with green oviparae and orange males appearing from July onwards, although
occasionally populations of viviparae persist until August or September (Stroyan
1984, Heie 1986). It occurs throughout
northern temperate parts of the world, and in Argentina. A nominal subspecies, A. farinosa yanagicola
Matsumura, occurs commonly on Salix
spp. (daphoides, rorida) in Japan, and is also recorded
from China (Qiao & Jiang 2005); it is distinguished by having long, fine
hairs on the third antennal segment (about twice the middle diameter of the
segment), and it produces sexuales in the autumn (Takahashi 1966). Populations on Salix gracilistyla in Siberia have short antennal hairs like
European farinosa, but a longer
last rostral segment; R IV+V 1.27-1.45 × HT II, as compared with 1.07-1.25 ×
HT II (Holman 1987). 2n=6. Aphis
feminea Hottes
Apterae red with black head, pro- and mesothorax, antennae, legs,
siphunculi and cauda, lightly dusted with white wax. It forms dense colonies on young twigs and
small branches of Prunus serotina
in north-eastern USA (Hottes & Frison 1931). Aphis (Zyxaphis) filifoliae (Gillette
& Palmer) (Fig.12s) Apterae are brownish amber to dark brown
or blackish, with lighter brown markings and a reticulate pattern of white
wax; BL c.1.1 mm. On leaves and stems of Artemisia
and Seriphidium spp., and also
reported to occur on Chrysothamnus nauseosus, in western USA. Monoecious
holocyclic; oviparae and apterous males were collected in early October (see
Palmer 1952). Aphis
filipendulae Matsumura Apterae are dark greenish brown,
yellow-green or green, powdered with wax; BL c. 1.0-1.3 mm. On flower stems
and lower leaf surfaces of Filipendula spp.,
causing wrinkling of leaves. In Japan (Hokkaido), east Siberia (Takahashi
1966; Pashchenko 1988a) and Kazakhstan (Kadyrbekov 2012a). Aphis (Bursaphis) fluvialis Martin Apterae are bluish green with a slight
wax bloom: BL 1.0-1.7 mm. On shoot apices and flower stalks of Epilobium hirsutum in Africa (west
Sudan), attended by ants. Probably anholocyclic. Martin (1982) discussed the
possible origins of the aphid and its host from Europe. 2n=9. Aphis
foeniculivora Zhang Apterae are green, BL c. 1.4 mm. On Foeniculum vulgare in China. Apart
from a longer antennal terminal process, this species could not be
distinguished by its published description from A. gossypii. Aphis
folsomii Davis Apterae are reddish brown (see influentialpoints.com/Gallery); BL 1.6-1.8 mm. On Parthenocissus spp., forming dense clusters along midribs of
leaflets and petioles, attended by ants (Williams 1911, as Aphis parthenocissi). Widely
distributed in USA east of Rocky Mountains. Aphis
forbesi Weed Apterae are dark bluish green, sometimes
with yellowish mottling (see influentialpoints.com/Gallery); BL 1.0-1.9 mm. Immatures are yellowish green. Alatae have black
dorsal abdominal cross-bands and only 1-4 secondary rhinaria on ANT III. In
ant-attended colonies on shoots and at bases of leaf petioles of Fragaria spp., and also sometimes on
the roots. Native to North America, and introduced into Europe about 1928;
also in Kazakhstan, west
Siberia, Japan, Australia (Brumley 2020)
and South America. Monoecious holocyclic with apterous males; Marcovitch
(1925) discovered the photoperiodic induction of sexual morphs in aphids
while working with this species. 2n=8. Aphis frangulae Kaltenbach Apterae are various shades of yellow,
green, blue-green, brown or greenish black, sometimes mottled (see influentialpoints.com/Gallery);
BL 0.9-2.4 mm. Alatae have secondary rhinaria distributed III 3-16, IV 0-8, V
0-3. A complex of species/subspecies utilising a wide range of herbaceous
plants, some of which have a sexual phase in Europe on Rhamnus frangula, curling its leaves in spring, and the name is
also applied to aphids utilising other Rhamnus spp. (davurica,
ussuriensis) as primary hosts in
east Asia (Pashchenko 1997). [A.
rhamnicola (q.v.) described from R
davurica in Korea is also not clearly distinct.] Parthenogenetic
overwintering of members of the group also occurs on the evergreen R. alaternus in the Mediterranean
region (Swirski & Amitai 2001), and an aphid on Sageretia in Pakistan has also been identified as A. frangulae (Naumann-Etienne & Remaudičre 1995). Stroyan (1984) reviewed the group and discussed
the taxonomic problems. In Europe, A. frangulae sensu stricto migrates
from R. frangula to Epilobium angustifolium, also
occurring frequently on Capsella
bursa-pastoris and Lysimachia
vulgaris, and A. frangulae beccabungae Koch migrates from the same primary
host but preferentially colonises Veronica
beccabunga, also occurring on Solanum
tuberosum and various Lamiaceae. A. f. beccabungae has
particularly been reported to occur on various labiates (Dracocephalum,
Scutellaria, Ziziphora) in the Kazakhstan part of West Tien-Shan
(Kadyrbekov 2005c). However, summer populations on labiates are
practically indistinguishable from those of other species in the complex
(e.g. balloticola, lamiorum, nepetae) that are monoecious holocyclic on the
same plant genera, and populations outside Europe (e.g. in south-west and
central Asia), can usually only be identified as frangulae (sensu latore). A
third European form only known from Germany, A. frangulae testacea Thomas,
may be monoecious on R. frangula.
Populations of these three subspecies possibly show colour differences in
life (see Heie 1986). There are no
fully reliable morphological discriminants, perhaps because natural
hybridisation may tend to confuse secondary host associations and life cycle
categories, and there are also geographical differences, which may change
with time. For example, Stroyan (1984,
p.124) noted a frangulae-like aphid that had recently been collected in England
on Hypericum; this seems to be the
same as an Aphis sp. collected in
Syria and Lebanon prior to 1954 (BMNH collection, leg. A.L. Talhouk). Alatae
in populations on Nepeta from Spain
(leg. H.L.G. Stroyan) and USA (Idaho, Utah; leg. G.F. Knowlton) all have a
large number of secondary rhinaria distributed over ANT III, IV and V, and
may be a distinct species. The cosmopolitan pest A. gossypii (q.v.) is
closely related, and introduces a further complication to the taxonomic
treatment of the group, as it can be conveniently
treated as a subspecies of frangulae in
Europe (Stroyan 1984, Heie 1986). However, this becomes untenable when
world-wide populations are taken into account (see below and Blackman &
Eastop 2017), and recent work comparing mitochondrial DNA sequences (Cocuzza et al. 2009, Carletto et al. 2009a) has shown clear
differences between the frangulae
and gossypii groups. Cocuzza &
Cavalieri (2014) have gone on to compare a mitochondrial DNA (COI) sequence
in numerous samples of frangulae-group
aphids collected from Lamiaceae. They found generally very low levels of
variation, and in particular suggested that three nominal species (lamiorum, stachydis, symphyti)
should be regarded as synonyms of frangulae,
but this does not take into account the different life cycles of these
species. Further molecular work is needed using different genetic markers.
2n=8. Aphis franzi Holman Colour of apterae in life unknown; BL
1.1-1.7 mm. Alatae are undescribed. On Seseli
elatum ssp. austriacum in
Austria, and reported also to occur in Kazakhstan on several other Seseli spp., with records also from Peucedanum morisonii and Silaum alpestre (Kadyrbekov 2009a,
2012a, 2017a); also in west Siberia (Kadyrbekov 2014g – but there
on Linaria, unlikely to be a true
host).
Aphis fraserae Gillette
& Palmer Apterae are yellowish green,
mottled with darker green, often blackish, with dark siphunculi and cauda; BL
c.2 mm. Apterae have 0-15 secondary rhinaria on ANT III, and alatae have
18-28 on III and 1-7 on IV. On flower stalks of Frasera speciosa in western USA (Palmer 1952). Distinguished from
polyphagous Aphis spp. by the long R IV+V and short ANT PT. Aphis fuentesi Nieto Nafría
& Ortego Apterae are matt dark
green to matt black; BL 1.0-1.7 mm. Alatae have secondary rhinaria
distributed III 4-13, IV 0-3. In compact groups on stems of several Baccharis spp. in Argentina (Neuquén,
Chubut, Rio Negro) and Chile (Maule). Oviparae and males were collected in
April (Nieto
Nafría et al. 2019c). Aphis fukii Shinji Apterae are dark grey-brown or black,
with black siphunculi and cauda; BL 2.1-2.5 mm. Alatae have secondary
rhinaria distributed III 20-32, IV 5-11, V 1-7. Curling and twisting leaves
of Petasites spp. in east Asia
(Japan, Korea, Sakhalin, Taiwan). Also recorded (from Japan) on Senecio kaempferi (= Farfugium japonicum). 2n=8. Aphis fumanae Remaudičre
& Leclant Apterae are very dark
green to black, shining; BL 0.9-1.3 mm. Alatae have 3-8 secondary rhinaria on
ANT III. In small colonies on shoots and flower peduncles of Fumana
spp. in southern France. Monoecious holocyclic with apterous males
(original description). Colonies were attacked by the aphidiid parasitod Lysiphlebus fabarum Marsh. Aphis funitecta (Börner) Apterae are densely wax-covered, BL
1.6-2.4 mm. On Bupleurum spp. in Germany, Poland, Kazakhstan
(Kadyrbekov 2017a) and Russia (Altai Krai). Monoecious holocyclic with
apterous males, and oviparae having unswollen hind tibiae (Nieto Nafría et al. 1986). Aphis
galiae Ivanoskaya
Apterae are broadly oval, greenish brown with black antennae, legs,
siphunculi and cauda; BL c. 2.3-2.4 mm. Described from Populus in western Siberia, and also recorded from P. nigra in Kazakhstan (Kadyrbekov
2017a), but there are also records from Betula
nana (in west Siberia), and from Betula
sp. in the Kuril Islands (Ivanoskaya 1977, Krivoluckaya & Ivanoskaya
1966), indicating a host range that warrants further investigation (see also Stekolshchikov & Khureva 2020).
The life cycle is unknown. Aphis galiiscabri Schrank (Fig.29a) Apterae are greenish black, heavily
wax-powdered, BL 1.4-2.2 mm. Alatae have
secondary rhinaria distributed III 5-14, IV 0-3, V 0-1. On
upper parts of Galium spp., causing
shortening of stems so that leaves and flowers of young shoots become close-set.
It is also reported from some species of Asperula
and Rubia. In Europe, west
Siberia, Turkey, Iran and Central Asia (Kazakhstan); Nieto Nafría et al. (2005a) reported it also in
Mongolia and Canada. Populations in southern Italy show some morphological
differences (Stroyan 1984). Fundatrices have been collected in May (BMNH
collection), sexual morphs are apparently unknown. Aphis gallowayi Robinson Apterae are dark brown to black, BL
1.6-2.2 mm. On Astragalus pectinatus in
Alberta, Canada. Very similar to A.
astragalina, and which has been found on the same plant. Aphis gaultheriae López Ciruelos,
Brown & Nieto Nafría Colour in
life unknown, BL of aptera 1.8-2.1 mm. On Gaultheria
mucronata in southern Chile (Chiloé). Other morphs and life cycle are
unknown (López Ciruelos et al.
2018). Aphis genistae Scopoli Apterae are black, coated with wax meal (see influentialpoints.com/Gallery);
BL 1.4-2.6 mm. Alatae have 4-8 secondary rhinaria on ANT III. On Genista
spp. in Europe, eastward to Ukraine and Turkey, and introduced to North
America (Foottit et al. 2006).
Sometimes with ants. Monoecious holocyclic, with oviparae and alate males in
September 2n=8. Aphis gentianae (Börner) Apterae are dark green, BL 1.2-1.6 mm, at
stem bases in ant shelters and on subterranean parts of Gentianaceae (Blackstonia, Centaurium, Gentiana) in
central, southern and eastern Europe, south west
Siberia (Novgorodova & Stekolshchikov 2013), and recently
found in Wales (Baker 2009a). A redescription of apterae and alatae was
provided by Barbagallo & Massimino Cocuzza (2014). Sexual morphs were
found in Poland in September (Szelegiewicz 1975a). Infestation of Gentiana cruciata stems by A. gentianae significantly deterred
oviposition by the butterfly Maculinea
alcon (Árnyas et al. 2009). Aphis gerardiae (Thomas) Apterae are blackish (see influentialpoints.com/Gallery),
BL 1.9-2.9 mm. On Agalinis and Aureolaria spp. in eastern USA
(Florida, Illinois, Indiana, Kansas, Kentucky, New York, Pennsylvania). Life
cycle seems not to have been studied. Aphis gerardianae Mordvilko Apterae are greyish black to black,
wax-dusted; BL 1.0-1.7 mm. Alatae have 8-11 secondary rhinaria on ANT III. On
stems and flower stalks of Euphorbia
sequieriana (incl. gerardiana) in central and eastern Europe (Jörg
& Lampel 1988), and eastward to Central Asia, where Kadyrbekov (2005b)
recorded it from several other Euphorbia spp. in Kazakhstan. In
Hungary it is attended by the ant Plagiolepis
pygmaea (Szelegiewicz 1966a). Aphis glareosae
Bozhko Apterae are dark brown,
wax-dusted; BL c. 2 mm. Alatae have secondary rhinaria distributed III 10-12,
IV 1-3. On Euphorbia glareosa, and
also on E. pannonica (BMNH
collection, leg. J. Holman), in
eastern Europe (Hungary, Czech Republic, Bulgaria, Rumania, Ukraine,
Turkey). Oviparae have been collected
on E. glareosa in October (BMNH
collection, leg. J. Holman). Aphis globosa Pashchenko Apterae are pale green with some dark
abdominal markings, BL c. 1.8 mm. On underground stems of Artemisia
gmelinii in east Siberia. Biology is unknown. Aphis glycines Matsumura Apterae are pale yellow or greenish
yellow with pale head and basal antennal segments, and siphunculi dark except
at their bases (see
influentialpoints.com/Gallery); BL 1.2-1.7 mm. Alatae have 4-9 secondary rhinaria on ANT
III. On stems and undersides of leaves of various Leguminosae/Fabaceae,
particularly Glycine spp., and a
major pest of soybean (Blackman & Eastop 2000, p.229); also recorded from
Amphicarpaea, Desmodium, Phaseolus
and Pueraria. In east and
south-east Asia, and introduced to USA in about the year 2000 (Ragsdale et al. 2004), and then to Canada (Hunt
et al. 2003). It has also been present in eastern Australia
since at least 2000 (Fletcher & Desborough 2000), and is also in New
Caledonia (Mille et al. 2020).
Heteroecious holocyclic in China and Japan with Rhamnus spp. (davurica,
japonica) as primary hosts (Wang et al. 1962, S. Takahashi et al. 1993, Kim et al.
2010b). In North America Rhamnus cathartica
and R. alnifolia are accepted as
primary hosts (Voegtlin et al. 2004, 2005; Yoo et al.
2005). Its introduction into USA stimulated the setting-up of a soybean aphid
suction trap network (reviewed by Lagos-Kutz et al. 2020), and led to intensive studies of all aspects of its
biology; see Tilmon et al. (2011)
for a general review, and Orantes et al.
(2012), Jun et al. (2013) and
Giordano et al. (2020) for
subsequent studies of genetic diversity in US and Asian populations. Naaum et al. (2014) have developed a
specific PCR assay with potential for field identification.
H. Li et al. (2018) developed
methods of instar identification. The complete genome sequence has been
published (Wenger et al. 2020), as
well as that of the mitochondrial genome (Song et al. 2019). 2n=8 (Mandrioli et
al. 2019b). Aphis
gnetuma Qiao Apterae are reddish brown to dark brown,
covered with white wax powder; BL 1.7-1.9 mm. On undersides of leaves of Gnetum parvifolium in Guangxi, China
(Y. Xu et al. 2021). Other morphs and life cycle are unknown. Aphis
gossypii Glover Plate 7c Apterae vary in colour from dark blackish
green or green mottled with dark green (larger specimens in favourable
conditions; see
influentialpoints.com/Gallery) to very pale whitish yellow (small specimens in
crowded colonies or hot conditions); BL 0.9-1.8 mm. Alatae have
secondary rhinaria distributed III
(3-)6-12(-15), IV 0(-2). On a very wide range of host plants, its polyphagy
being particularly evident during the dry season in hot countries. A major
pest of cotton and cucurbits, and in glasshouses in cold temperate regions
(Blackman & Eastop 2000, p.230). Distributed almost worldwide, and
particularly abundant and well-distributed in the tropics, including many
Pacific islands. Anholocyclic in warm climates, and mainly so in Europe.
Molecular studies indicated that occasional (“cryptic”) sex may be occurring
in southern France (Thomas et al.
2009), and subsequent work (Thomas et
al. 2012) found evidence of recent sexual reproduction in spring migrant
alatae alighting on melon plants in three French regions, although the
primary hosts of these alatae were unknown and some confusion with members of
the A. frangulae group seems
possible. Host alternation and a sexual phase occur more
regularly in parts of east Asia (e.g. Japan, Takada 1988; China, Zhang &
Zhong 1982a) and also in North America (Kring 1959), with several unrelated
plants utilised as primary hosts (incl. Catalpa
bignonioides, Hibiscus syriacus,
Celastrus orbiculatus, Rhamnus spp. and Punica granatum). Apterae of spring populations on
primary hosts are usually greenish. Males are always alate. However, Komazaki & Toda (2008) obtained wingless gynoparae from some
holocyclic clones in Japan, and Zhang
& Zhong (1982a) noted monoecious holocyclic populations on cotton and Hibiscus in China. Likewise there are aphids identified as A. gossypii that overwinter as eggs on
twigs of Lycium barbatum in China
(Yan et al. 2017). Molecular studies indicate the occurrence of
host-associated populations or races, some of which may have a world-wide
distribution (Charaabi et al. 2008,
Carletto et al. 2009b, L.Wang et al. 2016, Luo et al. 2016, R.H. Miller et
al. 2019); but see also Lokeshwari et
al. (2015). Genetic
diversity of A,. gossypii
populations in a cotton agroecosystem in northern Cameroon has been studied
by Brévault et al. (2008), and Nam et al. (2019) investigated the genetic
structure of populations on peppers in Korea using microsatellites. Najar-Rodríguez et al. (2009) demonstrated clear differences in both behavioural and
physiological mechanisms of host detection and acceptance of Malvaceae and
Cucurbitaceae by alatae of clones originating respectively from cotton and
pumpkin in Australia. Y.-C. Zhang et al.
(2017) found differences in gene expression profiles between cotton-specialised
and cucurbit-specialised lineages of A.
gossypii in China. Some populations regarded as A. gossypii certainly seem to be functioning as distinct species;
for example, populations
with a sexual phase on Rubia cordifolia
in Japan that seem to be isolated from those on other primary hosts
(Inaizumi 1981), populations reproducing sexually on Hibiscus in Korea that would not colonise vegetable crops
including cucumber (Y.-H. Kim et al.
2008), and a form feeding on Compositae/ Asteraceae in Europe and north
Africa that can produce sexual morphs under certain conditions (A. parvus Theobald? - see
Margaritopoulos et al. 2006). However, F. Zhang & Liu
(2012) demonstrated interbreeding between populations with different life
cycles and host associations in China. Populations producing eggs on roots of various plants (Ixeris, Lactuca, Marrubium, Viola) in China identified as A.
gossypii (e.g. Chu 1949) may be other closely-related monoecious
holocyclic species, such as A.
violaeradicis (q.v.). Taxonomic
implications of the genetics, biology and distribution of A. gossypii were reviewed by Blackman
& Eastop (2007, 2017). A. gossypii
has been often been treated as a subspecies of A. frangulae in the European aphid taxonomic literature, but
recent molecular work has shown that these two taxa are phylogenetically
distinct (see under A. frangulae).
Favret & Miller (2011) have designated a neotype for A. gossypii from material freshly collected in South Carolina,
and published its DNA barcode. The complete mitochondrial genome has been
published (S. Zhang et al. 2016),
and a high quality draft of the whole nuclear genome is now available (Quan et al. 2019) . 2n=8. Aphis
grandis Juchnevitch Apterae have dark brown head and prothorax,
rest of thorax and abdomen yellow with broad brownish cross-bands and
marginal spots, appendages mainly dark brown; BL 2.8-3.1 mm. Alatae have
23-27 secondary rhinaria on ANT III. In ant-attended colonies on green shoots
of Spiraea spp. in Kazakhstan.
Apparently migrating in June to an unknown secondary host. Aphis grata Pashchenko
Apterae are brown, pulverulent; BL 1.3-1.9 mm. Alatae have 8-9
secondary rhinaria on ANT III. In small dense colonies on undersides of
leaves and flower stems of Spiraea spp.
in east Siberia. Aphis
gratiolae Bozhko Apterae are yellowish with greenish
mottling; BL c.1.2 mm. Alatae have 4-5 secondary rhinaria on ANT III. On Gratiola officinalis in Ukraine. Aphis
gregalis Knowlton Apterae are bluish green to yellowish
green with darker head and pronotum, and variably-developed dorsal abdominal
dark markings; BL 1.2-1.9 mm. Secondary rhinaria of alata ditributed III 2-7,
IV 1-2. On Chrysothamnus spp. in
western USA (western slopes of Rocky Mountains), often in large colonies on
shoot tips, leaves and flower-stems, causing stunting and distortion of
growth. Oviparae and apterous males in October (original description).
Closely related to A. ornata, and without any clear distinguishing
characters in slide-mounted specimens. Aphis
grosmannae Börner Colour of apterae in life unknown,
probably dark green; BL 1.8-2.5 mm. Alatae have 6-12 secondary rhinaria on
ANT III. On stems and umbels of Peucedanum
spp. in central and eastern
Europe (Germany, Poland, Czech Republic), Kazakhstan and west Siberia. There
are also records from other Umbelliferae/Apiaceae (Angelica, Cicuta, Heracleum, Libanotis, Seseli), and
from Parasenecio (= Cacalia) hastatus (Ivanoskaja 1977; probably not a normal host).
Szelegiewicz (1964b) provided a redescription. Aphis (Bursaphis) grossulariae Kaltenbach Apterae are dull greyish green to dark green or yellow-green, with pale siphunculi (see influentialpoints.com/Gallery); BL 1.2-2.1 mm. Secondary rhinaria of alata ditributed III 7-19, IV 3-11, V 0-5. Host-alternating between Ribes spp., mainly |