The Aphids

Three little-known nearctic species with similar apically knobbed siphunculi to Epameibaphis, but without the stiletto-shaped R IV+V of anthemid-feeding aphids, their hosts being Solidago.

Cachryphora canadensis Hille Ris Lambers (Fig.49f) Apterae are “green with a black dorsal shield” (from slide label in BMNH collection); BL 1.0-1.3 mm. On Solidago spp. in eastern USA (North Carolina, Pennsylvania) and Canada (Ontario).

Cachryphora imbricaria Richards Apterae are dusky yellow with dark apices to antennae, legs and siphunculi BL 1.1-1.7 mm. On Solidago sp.in North Carolina, USA.

Cachryphora serotinae (Oestlund) Plate 15d (Fig.49e) Apterae are golden yellow to yellowish apple-green, siphunculi dusky with dark apices; BL 1.1-1.3 mm. On Solidago spp. in western USA (Colorado, Idaho, Utah). Monoecious holocyclic, with oviparae and apterous males in October (Palmer 1952, as Rhopalosiphum serotinae).

Calaphis Walsh

Calaphidinae: Calaphidini

Delicately built aphids with rather long thin legs, generally pale-coloured but sometimes with distinctive markings and/or dark wing veins. Other characters of the genus are the well-developed antennal tubercles and the absence or indistinctness of the radial sector in the forewing. Some species have all viviparae alate, others have both apterous and alate viviparae. As treated here, Calaphis is holarctic and includes Kallistaphis and Cepegillettea. There are about 19 species, 11 of which are in North America, three are palaearctic and three are East Asian. All live on Betulaceae except for one North American species on Myrica (myricae) and two East Asian species on Magnolia. The two Magnolia-feeding species were reviewed by Takahashi (1965b), Higuchi (1972) and Moritsu (1983) as Neocalaphis, and probably warrant separate generic or at least subgeneric status, but this genus name is currently listed as unavailable (Nieto Nafría & Favret 2011). The Betula-feeding species are parasitised by Aphidius aquilus, which also attacks other related aphids feeding on birches (Starý 1973).

Calaphis alni Baker (fig. 29D) Apterae and alatae of BL about 2.3-2.4 mm, yellowish with distinctive black markings; alatae usually have a large black dorsal abdominal patch and apterae have two bands across the thorax and one across the posterior abdomen. Feeding mainly on the stems and foliage of tender growing terminals of Alnus rugosa (Pepper 1950), in eastern USA and Canada (New Brunswick). The sexuales are undescribed.

Calaphis arctica Hille Ris Lambers Apterae of BL about 1.9-2.1 mm, green, with dusky dorsal hairs and spots (O. Heikinheimo, pers. comm.). Originally described from Betula nana in Greenland, and since recorded from Finland (BMNH collection; leg. Heikinheimo), alpine Germany (BMNH collection, including undescribed alate viviparae), Czech Republic (Holman 1996) and Kazakhstan (Kadyrbekov 2016b). Apterous males and oviparae were collected in late October in Germany (BMNH collection, leg. D. Hille Ris Lambers). Holman (1996) noted differences between the Greenland and European populations indicating that two taxa may be involved. 2n=18 (Blackman & Eastop 1994).

Calaphis betulaecolens (Fitch) Viviparae are all alate, bright lemon yellow without dorsal markings, with mainly blackish antennae and tibiae and dark wing veins (see influential points/gallery); BL 3.0-3.5 mm. Usually on leaves of Betula lutea and B. papyrifera, occasionally on other Betula spp. Widely distributed in North America. (Japanese records of this species apply to C. similis, q.v.). Quednau (1971) fully redescribed this species. Apterous oviparae and alate males occur in September-October. 2n=20.

Calaphis (Cepegillettea) betulaefoliae (Granovsky) Apterae are yellow, greenish-yellow or green, without dorsal markings, antennae and legs mainly pale to dusky with black apex to antennal segment 3, femoro-tibial joints and tarsi, and black tips to the siphunculi; BL 2.9-3.5 mm. Alatae similar, with yellow-orange head and thorax and dark wing veins. Occurring singly or in small groups on undersides of leaves or young growth of Betula spp. (glandulosa, occidentalis, papyrifera, pumila); recorded locally in USA (Illinois, Wisconsin, Colorado), common and widespread in Canada. Prefers shady positions; very active, dropping when disturbed. Apterous oviparae and alate males occur in September-October (Granovsky 1928b).

Calaphis betulella Walsh Plate 9e Viviparae are all alate, yellowish with orange and black markings, black stripes along the femora and wholly black tibiae and tarsi; BL 2.0-2.5 mm. Feeds only on Betula nigra, on undersides of leaves. Apterous oviparae and alate males occur in September-October. In eastern USA. 2n=18.

Calaphis betulicola (Kaltenbach) Apterae are pale yellowish to green, with black apices to antennal segments, femoro-tibial joints, tibial apices and tarsi, and dark tips to siphunculi (see influentialpoints.com/Gallery); BL 2.0-2.3 mm. Alatae have darker wing veins than the rather similar species C. flava. On undersides of leaves of Betula spp. (davurica, pendula, platyphylla, pubescens, ermannii, kirghisorum, mandshurica), usually on seedlings and small trees less than 1 m high (Stroyan 1977). Holarctic in distribution, from western Europe across Russia to China (Qiao et al. 2005b) and Japan, and also in USA (California; Quednau 1971) and Canada (New Brunswick, Quebec; BMNH collection). Apterous oviparae and alate males occur in September-November. (Pre-1957 records include C. flava.) 2n=18.

Calaphis coloradensis Granovsky Apterae are pale yellow to yellow-green, tinged with green, sometimes with a very light brownish cast, with dark apices to antennal segments, femoro-tibial joints, apices of tibiae, and tarsi; BL 1.5-1.7 mm. Alatae have both cubitus veins of forewing heavily bordered. Feeds only on Betula occidentalis, on lower sides of leaves and terminals of young growth. In western USA and Canada (Alberta, Colorado, Idaho, Montana, Washington, Utah). Apterous males and oviparae occur in Sepember-October (Granovsky 1939, Palmer 1952). 2n=18.

Calaphis flava (Mordvilko) (= basalis Stroyan 1957; = granovskyi Palmer 1952) Apterae are pale green or yellowish, with dark apices to antennal segments, femoro-tibial joints, tibial apices and tarsi (see influentialpoints.com/Gallery); BL 1.9-2.7 mm. Alatae have wing veins dark, but less dark than in betulicola, and the siphunculi are pale. Betula pubescens seems to be the preferred host in Europe (Heie 1982), but the young growth of many other Betula species may be colonised, and it has also been collected from Alnus firma and A. serrulata (BMNH collection). Widespread in Europe (where until 1957 it was confused with the less common betulicola), eastward to Siberia and Central Asia, and introduced into South Africa, Australia, New Zealand and North America (where it was described as granovskyi – Palmer 1952). DNA barcoding has revealed that there may be two or even three cryptic species present in North American populations identified as C. flava (Y. Lee et al. 2017), and presumably there is similar diversity yet to be discovered in Old World populations.. Alate males and apterous oviparae occur in September to November, and may be produced on numerous Betula spp. (BMNH collection). 2n=18.

Calaphis leonardi Quednau Viviparae are all alate, pale yellowish-green and orange with brown-black antennae, tibiae and tarsi, and dark wing veins (see influential points/gallery); BL c. 3.0-3.3 mm. Feeds only on Betula populifolia, in north-eastern USA and eastern Canada (Quednau 1971). Oviparae have dark dorsal markings and occur in September-October; males are not yet recorded. 2n=20*.

Calaphis magnoliae (Essig & Kuwana) All adult viviparae are alate, pale shiny straw yellow to yellow-green with antennae and bases of tibiae banded with black, and forewing veins conspicuously black-bordered (see influential points/gallery); BL 1.4-2.2 mm. On leaves of Magnolia kobus and M. liliflora in Japan and Korea. Sexual morphs are undescribed. 2n=20 (Blackman 1986, as Neocalaphis).

Calaphis magnolicolens (Takahashi) All adult viviparae are alate, almost translucent yellowish white to very pale green (see Moritsu 1983, as Neocalaphis); BL 2.3-3.3 mm. On undersides of leaves of Magnolia obovata in Japan. Immatures feed alongside main veins. Sexual morphs are undescribed. 2n=20 (Blackman 1986, as Neocalaphis).

Calaphis manitobensis Richards Apterae are bright yellow, with antennae and legs almost entirely pale; BL about 2 mm. Alatae and sexuales are undescribed. Known only from the original description, from Betula pumila in Manitoba, Canada (Richards 1968e).

Calaphis (Cepegillettea) myricae (Patch) Plate 4a Apterae and alatae are both green with dusky or dark distal antennal segments, tibiae and tarsi, and siphunculi either wholly or apically black; BL about 2.5 mm. On leaves of Myricacae (Myrica gale, Comptonia peregrina) in north-eastern USA and eastern Canada. Apterous males and oviparae occur in September (BMNH collection.; leg. E. MacGillivray).

Calaphis neobetulella Quednau All viviparae are alate, greenish yellow with black dorsal markings on head, thorax and abdominal tergites 4-5, and antennal flagellum, legs and siphunculi also mainly or entirely black; BL 2.0-2.5 mm. On undersides of leaves of Betula nigra in north-eastern USA (North Carolina, Virginia, New Jersey, Washington DC). Alate males and apterous oviparae occur in September (Quednau 1971).

Calaphis similis Quednau All viviparae are alate, yellow with reddish head and two red spots on abdomen, with antennal flagellum, tibiae and tarsi mainly black, siphunculi pale, wing veins bordered with black; BL 3.0-3.5 mm. On undersides of leaves of Betula spp. (costata, davurica, mandshurica, platyphylla var. japonica) in China, Korea, Siberia and Japan (Quednau 1979, Pashchenko 1988a, Qiao et al. 2005b). Oviparae and alate males were collected in October (BMNH collection, and Paik 1972; as Kallistaphis sp.) Higuchi’s (1972) account of C. betulaecolens applies to this species.

Calaphis ugamica Kadyrbekov Apterae and alatae are pale green with apices of antennal segments, femoro-tibial joints and tarsi brown; BL 2.0-2.3 mm. The siphunculi of the aptera (but not those of the alata) are dark-tipped. On upper sides of leaves of Betula turkestanica in southern Kazakhstan (Kadyrbekov 2014a). The description of the alata does not provide distinguishing characters from that of C. flava.

Calaphis viridipallida Palmer Apterae and alatae are pale greenish-yellow with apices of antennal segments, femoro-tibial joints, tibial apices and tarsi dark; BL 1.6-2.2 mm. On undersides of leaves of Betula spp. (glandulosa, lutea, papyrifera), widely distributed in North America (California, Colorado, Minnesota, North Carolina, Manitoba, Ontario, Quebec, Labrador). Oviparae and alate males occur in September-October. 2n=18.

Callipterinella van der Goot

Calaphidinae: Calaphidini

A genus of three species associated with Betula. Apterous viviparae occur commonly and have a brown vertex to the head and usually other dark dorsal markings. The siphunculi are dark and have rows of minute spinules. Stroyan (1977) and Heie (1982) both provided accounts of this genus.

Callipterinella calliptera (Hartig) (= annulatus Koch) Apterae are yellowish green to green, usually with dark transverse bands on all tergites (see influentialpoints.com/Gallery); BL 1.5-2.5 mm. Alatae have dorsal markings less well developed. In ant-attended groups on undersides of leaves of Betula spp., often inside leaves sewn up by lepidopterous larvae; on B. pendula, less commonly B. pubescens, in Europe, and on B. platyphylla, B. mandshurica and B. dahurica in east Asia (but see under C. tuberculata below). It occurs throughout Europe and across Asia to east Siberia, Korea, China and Japan); introduced and widespread on planted B. pendula in North America. Oviparae and alate males occur in September-November. Hajek & Dahlsten (1988) studied the population dynamics of this species in California. 2n=20.

Callipterinella minutissima (Stroyan) Apterae are very small, stout, oval -bodied, green to yellowish green with a dark transverse bar on abdominal tergite 8 only; BL 0.9-1.4 mm. On Betula spp. in Europe, Central Asia (Kadyrbekov 2017a), and introduced into western North America, where it is also recorded from B. papyrifera (BMNH collection). The size and shape of the apterae enable them to feed inside bud scales and developing leaves when the buds burst in spring, and in the female catkins in summer (Hajek 1985). Oviparae (which are larger and darker than apterous viviparae) and alate males occur on the leaves in September-November.

Callipterinella tuberculata (von Heyden) Plate 9c, d Apterae are yellowish with a brown head and dark brown dorsal abdominal markings comprising especially a dark quadratic patch on abdominal tergites 4-6 (see influentialpoints.com/Gallery); BL 1.7-2.2 mm. Alatae have irregular small sclerites on posterior abdominal tergites, but no transverse bands. Apterous males and oviparae are found in September. Apparently specific to Betula pendula in Europe, but Pashchenko (1988b) recorded apterous Callipterinella with a dorsal quadrate patch similar to that of C. tuberculata on various Betula spp. in east Siberia, as well as apterae without such a patch on the same hosts, indicating that the specific distinction between callipterus and tuberculata, which are closely related, may not be applicable in that part of the world. Also recorded from Kazakhstan (Kadyrbekov 2017a) and China (G. Zhang et al. 1986b). 2n=20.

Campanulaphis Kadyrbekov

Aphidinae: Macrosiphini

One species in Central Asia related to Uroleucon, but adapted to feeding on roots and attendance by ants.

Campanulaphis radicivora Kadyrbekov Apterae are dark brown to blackish; BL 2.1-4.1 mm. In ant-attended colonies on roots of Campanula spp. (glomerata, sibirica) in Kazakhstan (Kadyrbekov 2016c).

Capitophorus van der Goot

Aphidinae: Macrosiphini

About 30 species of pale, sometimes almost translucent, slender aphids with elongate appendages. Apterae bear long capitate hairs at least on the head and posterior abdominal segments. Alatae have only much shorter hairs, a dark dorsal abdominal often quadrate patch, and numerous rhinaria on antennal segments III, IV and usually V. They are widely distributed in palaearctic and oriental regions, and there are also five North American species. Six species are known to have host alternation, with primary hosts all in Elaeagnaceae. Secondary hosts are herbaceous Asteraceae and Polygonaceae. Some species are monoecious holocylic on plants in Elaeagnaceae, Asteraceaeae or Polygonaceae. The genus was onced used in a much broader sense to include most Macrosiphini with capitate hairs. A recent molecular phylogenetic study (Choi et al. 2017) places it in the Pterocomma group. Taxonomic accounts are available for North America (Corpuz-Raros & Cook 1974), north-west Europe (Heie 1994), UK and Ireland (Blackman 2010), central Europe (Heinze 1961a), India (Raychaudhuri et al. 1980b) and Japan (Miyazaki 1971).

Capitophorus archangelskii Nevsky Apterae are pale green to yellow-green with green markings, and pink eyes; BL 1.2-1.8 mm, on undersides of leaves of Elaeagnus spp. (angustifolia, occidentalis), and sometimes persisting on ends of shoots throughout the year (Nevsky 1928a). Alatae have a dark green central quadrate patch on the dorsal abdomen. In Caucasus, Iran, Kazakhstan, Uzbekistan, Afghanistan, Pakistan and India (West Bengal, Uttar Pradesh; BMNH collection). In Pakistan it is recorded from Calendula sp. (Naumann-Etienne & Remaudière 1995), so either there is a facultative migration to Calendula as secondary host, or two species of Capitophorus are being confused. Sexuales occur on Elaeagnus in October-November; males are alate (Nevsky 1928a).

Capitophorus bulgaricus Tashev Apterae are pale green; BL 1.3-1.75 mm. Alatae are undescribed. On undersides of leaves, usually along the veins, of Cirsium spp. In Bulgaria, Italy and Switzerland (BMNH collection, leg. D. Hille Ris Lambers). Monoecious holocyclic on Cirsium; Barbagallo & Patti (1998) described oviparae and males, collected in October.

Capitophorus carduinus (Walker) (Fig.22b) Apterae are. pale greenish white to yellowish green, almost translucent, often with two indistinct darker green longitudinal stripes (see influentialpoints.com/Gallery); BL 1.6-2.2 mm. Living all year round on Carduus and Cirsium spp., mainly on undersides of lower leaves. Oviparae and alate males occur in western Europe in September-October. In Europe, west Siberia, Iran, Iraq, western Himalayas and Korea. Records from North and South America may all be referable to C. elaeagni. Apterae of C. carduinus tend to have much smaller capitate spinal hairs on the anterior abdominal tergites than C. elaeagni, but in warm conditions (or when alatiform), C. elaeagni may also have smaller hairs. 2n=16.

Capitophorus cirsiiphagus Takahashi (Fig.22d) Apterae are pale green to white; BL c.1.5-1.6 mm. As noted by Miyazaki (1971), the spinal hairs on abdominal tergites 1-5 vary greatly in length and are often minute. On undersides of leaves of Cirsium spp. in Japan, Korea and east Siberia. Paik (1965) gives Aster tataricus var. hortensis as the host in Korea. The life cycle is unknown, probably monoecious holocyclic on Cirsium. 2n=16* (recorded as C. elaeagni in Blackman 1986).

Capitophorus elaeagni (del Guercio) Plate 16e Apterae in spring populations are pale green, BL 1.9-2.5 mm, on undersides of leaves of Elaeagnus spp., and sometimes on Hippophae (see influentialpoints.com/Gallery). Fundatrices have brighter green dorsal spots in longitudinal rows, and feed on upper sides of young leaves. Alatae, produced in the second and third generations, have black head and thorax, black antennae and a blackish central abdominal sclerite; they fly to tubuliferous Asteraceae (Arctium, Carduus, Cirsium, Cynara, Gerbera, Silybum). Apterae on secondary hosts are pale greenish white to yellowish green with dark tips to siphunculi; BL 1.4-2.5 mm. They feed mainly on the undersides of the lower leaves. Males and gynoparae return to Elaeagnus in autumn. C. elaeagni is widely distributed in temperate and warm temperate regions of the world. C. evelaeagni Zhang, described from C. segetum in China, will come to elaeagni in the key to aphids on Cirsium, and is probably a short-haired or alatiform variant of this species. See also Hille Ris Lambers 1953a and Blackman & Eastop 2000, p.252. 2n=16.

Capitophorus eniwanus Miyazaki Apterae are white or pale yellow; BL 2.0-2.1 mm. On Polygonum and Persicaria spp. in Japan and east Siberia. A record from Turkey (Şenol et al. 2014b) needs further confirmation. The life cycle is unknown. [A sample of small (BL 1.3-1.7 mm) alate spring migrants with numerous secondary rhinaria (distributed III 46-68, IV 26-39, V 11-23) from Elaeagnaceae in China near Beijing (leg. V.F. Eastop) could possibly be this species. This sample has been karyotyped and has 2n=10.]

Capitophorus essigi Hille Ris Lambers Apterae are probably pale green; BL 2.1-2.4 mm. On Polygonaceae (Persicaria alpina, Aconogon phytolaccifolium) in western USA (California, Idaho, Oregon, Washington). Apparently heteroecious with Shepherdia as the primary host, as sexuales identified as C. essigi have been collected on Shepherdia in late August in Idaho (A.Jensen, pers. comm. and aphidtrek.org).

Capitophorus formosartemisiae (Takahashi) Apterae are white; BL 1.1-1.8 mm. On undersides of leaves of Artemisia spp., and occasionally recorded from other Asteraceae (Dendranthema, Erigeron, Saussurea, Senecio). In India, Pakistan, China, Szechuan, Taiwan, Japan and east Siberia. It has also been in eastern USA since at least 1978 (Skvarla et al. 2017)). Tao (1999) has recorded this species from Elaeagnus angustifolia, presumably its primary host, in Taiwan. 2n=16.

Capitophorus gnathalifoliae Shinji Apterae are green, BL not recorded. On Gnaphalium hypoleucum in Japan.

Capitophorus gynoxanthus Hille Ris Lambers Apterae are pale greenish, often with two rather indistinct green longitudinal dorsal stripes; BL 1.6-2.5 mm. Monoecious holocyclic on Cirsium spp., especially C. arvense, feeding between young leaves at stem apices and on undersides of radical leaves, and giving rise in autumn to oviparae and alate males. Widely distributed in northern and central Europe, Israel, and a sample from C. falconeri in Kashmir in BMNH collection may also be this species. Described as a subspecies of C. horni, but elevated to full species status by Blackman (2010).

Capitophorus himachali Chakrabarti & Maity Apterae are pale; BL 1.7-2.0 mm. On Polygonum sp. in Himachal Pradesh, India. Its distinctness from C. eniwanus in Japan needs to be confirmed.

Capitophorus himalayensis A.K. Ghosh, M.R. Ghosh & Raychaudhuri Apterae are slender, BL 1.4-1.6 mm, pale except for brownish frons and dusky tips to tarsi and siphunculi. Alatae and other morphs are undescribed, and the life cycle is unknown. On Hippophae sp. (Saha & Chakrabarti 1988d) in north-west India (Uttar Pradesh, West Bengal).

Capitophorus hippophaes (Walker) Apterae in spring colonies on Elaeagnaceae (Elaeagnus spp., Hippophae spp.) are pale green, slender, with a faint pattern of green spots; BL 1.5-2.1 mm. Fundatrices are very different; broadly oval, greenish with reddish spots, and their antennae are dark, 5-segmented and have a short processus terminalis. Alatae, produced in the second and third generations, are greyish-green with a black head and thorax, dark antennae, legs and siphunculi and a a large quadrate dark green patch on the dorsal abdomen; they migrate to various Polygonum and Persicaria spp. (Hille Ris Lambers 1953a). Apterae on Polygonaceae are pale greenish to yellowish white, sometimes with longitudinal rows of green spots (see influentialpoints.com/Gallery); BL 1.7-2.4 mm. They live on the undersides of the leaves. A complex of forms exists, including anholocyclic populations distributed to various parts of the world and distinguishable by differences in size and distribution of dorsal capitate hairs. C. hippophaes sensu stricto occurs in Europe, North Africa, south-west and central Asia, and introduced into North America. Populations on Elaeagnus spp. and Polygonaceae in east Asia differ in the absence of any conspicuous capitate hairs on ANT I and abdominal tergite 6, and are usually regarded as a subspecies, C. hippophaes ssp. javanicus Hille Ris Lambers; this form occurs on Polygonaceae in south-east Asia, Australia, New Zealand, Central and South America, and also in California, USA (Corpuz-Raros & Cook 1974). 2n=10 (for both hippophaes sensu stricto and ssp. javanicus).

Capitophorus horni Börner (Fig.22a) Apterae are very pale green aphids, translucent, often with two faint brighter green dorsal longitudinal stripes, or yellowish; BL 1.6-2.5 mm. On undersides of leaves of Cirsium spp., usually C. oleraceum, and also sometimes on Cynara scolymus (Robert 1969). Monoecious holocyclic, with alate males. In Europe (Denmark, Norway, Sweden, Germany, Austria), south-west Asia (Iran; Rezwani 1998) and North Africa (Tunisia; Boukhris-Bouhachem et al. 2007). [The alate vivipara of horni is undescribed; an immature alata in a sample from Vienna (BMNH collection, leg D. Hille Ris Lambers) has adult cuticle showing distribution of secondary rhinaria III 21-22, IV 5-10, V 4-5, differing from gynoxanthus (III 22-35, IV 12-19, V 2-8; Hille Ris Lambers, 1953)]. 2n=16.

Capitophorus hudsonicus Robinson Apterae are pale green; BL 1.3-1.6 mm. Apterae (including fundatrices), alatae, oviparae and males were all swept from mixed vegetation, but the host plant was “almost certainly Shepherdia canadensis” (Robinson 1979), and this has snce been confirmed (R. Foottit, pers. comm.). Only known from northern Manitoba (Churchill), Canada. Presumably monoecious holocyclic on Shepherdia, with an abbreviated life cycle; fundatrices were collected in mid-July, and oviparae and alate males (along with apterous and alate viviparae) in mid-August.

Capitophorus inulae (Passerini) (Fig.34c) Apterae are transparent yellowish white; BL c.1.5-1.6 mm. On Inula spp., especially I. conyza, in Europe, Turkey (Görür et al. 2011b), and around the Mediterranean (where it is found also on Dittrichia viscosa and Pulicaria spp.), occurring eastward to Central Asia. Introduced to Australia (A.C.T. ; BMNH, coll. V.F.Eastop). Sexual morphs and life cycle are unknown; possibly anholocyclic in southern England. 2n=16.

Capitophorus jopepperi Corpus-Raros & Cook (Fig.8b) Aptarae are narrowly spindle-shaped, pale to dark green, BL 1.3-1.5 mm. Described from Ambrosia artemisifolia in North Carolina, USA, and since found on the same host in Mexico (BMNH collection, leg. G. Remaudière).

Capitophorus litanensis Raychaudhuri, Singh & Raychaudhuri Colour of apterae in life is unrecorded; BL c.1.5 mm. On Bidens sp. in Manipur, India.

Capitophorus meghalayensis Basu & Raychaudhuri Apterae are pale, narrow-bodied; BL about 1.3-1.5 mm. Alatae with a faint brownish patch extending from abdominal tergites 2-6 (R.C. Basu & Raychaudhuri 1976c); apparently there is no clearly defined central sclerite. On Elaeagnus sp. in Meghalaya, India. Oviparae are described, but the life cycle and secondary hosts, if any, are unknown.

Capitophorus mitegoni Eastop Apterae are pale green, spotted with darker green; BL 1.6-2.2 mm. On undersides of leaves of Polygonum and Persicaria spp., widely distributed in Africa and also in Afghanistan, India, Bangladesh, Bhutan, Nepal, Hong Kong, Philippines, New Guinea and Australia; this form was described as a subspecies of C. hippophaes but given specific rank by David & Ghorpade (1974). Sexual morphs are unknown, but spring populations (BL of apterae 1.1-1.3 mm) on Hippophae in Afghanistan (BMNH, coll. R. van den Bosch) are possibly this species.

Capitophorus montanus Takahashi (Fig.22c) Apterae are pale green; BL c.1.0-1.3 mm. On leaves of Cirsium spp. in Taiwan, Japan, Korea and east Siberia.

Capitophorus pakansus Hottes & Frison (Fig.34b) Apterae on Elaeagnaceae in spring are pale, narrow-bodied; BL 1.6-2.0 mm. Fundatrices are oval and mottled dark green. Alatae have black head and thorax and a quadrate dorsal abdominal patch; they fly for the summer to Inula spp. Apterae on Inula are white or greenish yellow, BL 1.3-2.2 mm, living on undersides of leaves. There is also a record from Telekia speciosa (Chumak 2004). Primary hosts are Elaeagnus spp. and sometimes Hippophaes rhamnoides (Norway; BMNH collection), although gynoparae in Holland did not produce oviparae on Hippophae (Hille Ris Lambers 1953a, as vandergooti). Widely distributed in Europe, in Kazakhstan (Kadyrbekov 2013d), in north-west China (G. Zhang 1999, as vandergooti) and in eastern North America. 2n=16*.

Capitophorus rhamnoides Zhang, Chen, Zhong & Li Apterae are pale green to green; BL c.1.2 mm. On undersides of leaves of Hippophae rhamnoides in China (Gansu, Qinghai) (G. Zhang 1999). Aphids described as C. tanensis by the same authors seem to be the autumn form of this species (Eastop & Blackman 2005). Possibly a synonym of C. mitegoni.

Capitophorus rostratus Miyazaki Apterae are milky or creamy white; BL 1.0-1.4 mm. On the densely hairy undersides of leaves of Synurus pungens (= deltoides) in Japan. Monoecious holocyclic with oviparae and alate males in early October (original description).

Capitophorus shepherdiae Gillette & Bragg (Fig.8c) Apterae are pale green with darker apices to legs and siphunculi; BL 1.3-2.3 mm. In spring colonies on Shepherdia (= Lepargyrea) argentea in western USA. Apparently persisting on this host throughout the summer, but morphologically similar aphids have been found on Ambrosia spp. in California and Washington, so there may be a partial migration which needs to be confirmed by transference tests (Corpus-Raros & Cook 1974 and aphidtrek.org).

Capitophorus similis van der Goot (Fig. 54a) Apterae in spring colonies on Elaeagnaceae are pale green, spindle-shaped (see influentialpoints.com/Gallery), BL 1.7-2.3 mm. Fundatrices are more oval in shape with 5-segmented antennae. Alatae have a creamy-coloured abdomen with a dark quadrate dorsal patch, and migrate to found colonies on undersides of leaves of Tussilago, Petasites, Adenostyles and Homogyne. Apterae on secondary hosts are yellowish white to white; BL 1.5-2.6 mm. Males and gynoparae migrate back to Hippophae or Elaeagnus in September-October (Hille Ris Lambers 1953a). Loher & Lampel (1983) studied the factors controlling sexual morph production in this species. In Europe, and west and central Asia to eastern Himalayas. Anholocyclic overwintering populations may occur on secondary hosts in warmer climates (Patti 1983).

Capitophorus takahashii Strand Apterae are white; BL c. 1.5 mm. On Petasites tricholobus and Tussilago farfara in Taiwan (Takahashi 1925, p. 21, as Capitophorus sp; Tao 1999).

Capitophorus tricholepidis Chakrabarti Apterae are pale brown; BL 1.2-1.5 mm. On undersides of leaves of Tricholepis furcata, attended by black ants. In Uttar Pradesh, India.

Capitophorus wojciechowskii Wieczorek & Kanturski Colour of apterae in life unknown; BL 1.5-1.9 mm. On Hippophae rhamnoides and Hippophae sp. in northern Iran. Monoecious holocyclic, with fundatrices in April, and oviparae and apterous males in October-November (Wieczorek & Kanturski 2015).

Capitophorus xanthii (Oestlund) Apterae are pale greenish; BL 1.4-2.0 mm. On Xanthium spp. in North America. Heteroecious holocyclic, with Elaeagnus and Hippophae as primary hosts (Corpuz-Raros & Cook 1974). Males and oviparae were collected on Hippophae rhamnoides in Colorado in August-October (Palmer 1952), but the spring generations on the primary host have not been described.

Capraphis Mier Durante, Ortego & Nieto Nafría

Aphidinae: Macrosiphini

A genus for one species on Mulinum in South America with long, curved, rather rigid antennae “resembling the horns of a goat”, and without ocular tubercles (triommatidia) (Mier Durante et al. 2009).

Capraphis blackmani Mier Durante, Ortego & Nieto Nafría Apterae yellow or pale green, BL 1.7-2.2 mm. Apterae have 2-7 secondary rhinaria on ANT III, and alatae have 13-15. On leaves of Mulinum spinosum in Argentina (Mendoza). Monoecious holocyclic, with oviparae and apterous males collected in late March (Mier Durante et al. 2009).

Caricosipha Börner

Chaitophorinae: Siphini

One species in Europe characterised by its stalked eyes and long black pointed hairs.

Caricosipha paniculatae Börner Plate 5c Apterae are yellowish or reddish with brown sclerotic patches to completely black dorsally, with pale antennae, legs and siphunculi; BL 1.5-2.4 mm. On leaves of Carex spp., widely distributed in Europe. Monoecious holocyclic with apterous males. Active insects, running fast when disturbed (Heie 1982).

Carolinaia Wilson

Aphidinae: Macrosiphini

About 16 species typically associated with Rhus and Toxicodendron, or with host alternation between these genera and Cyperaceae, although the life cycles of most species are unclear. Remaudière & Muñoz Viveros (1993) expanded the concept of the genus to include Glabromyzus (four North American spp.; reviewed by Cook 1984c) and Juncomyzus (five east Asian spp.). Noordam (2004) reviewed the two species in Java.

Carolinaia caricis Wilson (Fig.18e) Apterae are greenish yellow, tinged with brown; BL c. 1.4-1.7 mm. Alatae have secondary rhinaria only on ANT III. On various Cyperaceae, where it feeds mainly on the inflorescences. Apparently heteroecious holocyclic in eastern USA, where Smith (1980) made successful transfers from Rhus radicans to Carex spp. It presumably lives continuously on Cyperaceae at tropical latitudes (Puerto Rico, Costa Rica,Venezuela). Very similar to C. tissoti (q.v.).

Carolinaia carolinensis Smith Apterae (fundatrices) are yellowish orange; BL c.2.2 mm. Alatae have a central black dorsal abdominal patch. On Rhus radicans in North Carolina, USA, causing slight cupping of leaves on ground; other species on this host have been found only on leaves of shoots climbing in trees (original description).

Carolinaia (Glabromyzus) corazonensis Remaudière & Muños Viveros Apterae are matt black with paler antennae and legs; BL 1.4-1.8 mm. On Cyperus incompletus (= hermaphroditus) in Mexico. Alatae have 8-13 secondary rhinaria on ANT III.

Carolinaia cyperi Ainslie Apterae are black or greyish black, BL 1.0-1.5 mm. Alatae have 7-11 secondary rhinaria on ANT III, and 0-1 on IV. On Cyperus spp., and far more rarely on other Cyperaceae. It forms large and dense colonies on the basal parts of the leaves, and is common throughout Central America (Florida to Surinam). It has economic importance as a vector of sugar cane mosaic potyvirus. Sexual morphs are unknown.

Carolinaia floridensis Remaudière & Muños Viveros Colour of apterae in life unknown; BL 1.2-1.5 mm. Alatae have a large dark dorsal abdominal patch. On Rhus radicans in Florida, USA. Apparently it is anholocyclic on Rhus.

Carolinaia (Juncomyzus) floris (Miyazaki) Apterae are yellowish brown to reddish brown, BL 1.4-1.7 mm. Alatae have 9-15 secondary rhinaria on ANT III. On Juncus and Carex in Japan, living in inflorescences. Described from Juncus and Carex (Miyazaki 1971), but fundatrices and alatae collected on Rhus ambigua (= Toxicodendron orientale) in Hokkaido, Japan (BMNH collection, leg. M. Miyazaki) were subsequently identified as this species. Its appearance in life on Rhus is not recorded. BL of fundatrices is 2.1-2.2 mm. An alata trapped in Korea also seems to be C. floris (BMNH collection, leg. W. Paik). Biology has not been studied, and the host alternation requires experimental confirmation.

Carolinaia (Juncomyzus) hillerislambersi (Calilung) Apterae are greenish black or black; BL 1.7-2.0 mm. Alatae have 10-16 secondary rhinaria on ANT III. In ant-attended colonies on stems of Scleria scrobiculata in the Philippines. An alata has subsequently been collected in Papua New Guinea (BMNH collection, leg. J.H. Martin).

Carolinaia (Glabromyzus) howardii (Wilson) Apterae are brown with dark brown siphunculi; BL 1.5-2.1 mm (on Rhus; 1.1- on grasses). Heteroecious holocyclic between Rhus radicans and various Poaceae (e.g. Elymus, Glyceria, Panicularia) (C.F. Smith, cited in Remaudière & Muños Viveros 1993). Records from cereal crops should probably be referred to C. rhois (Cook 1984c). In eastern USA.

Carolinaia (Juncomyzus) japonica (Takahashi) Apterae are shining yellowish brown, with black antennae, femora, tibial apices, siphunculi and cauda; BL c. 1.9 mm. On Toxicodendron trichocarpum and T. vernicifluum in Japan (Takahashi 1924a, as Myzus rhois; Takahashi 1963, as Sitomyzus japonicus – but the fundatrix described under this name is C. (J.) nigra, see Miyazaki 1971). Probably heteroecious, but secondary host unknown; oviparae were found on T. trichocarpum in September (Remaudière & Muñoz Viveros 1993).

Carolinaia (Juncomyzus) javanica Noordam Apterae are dirty green, with brown head, thorax and sides of abdomen, siphunculi black-tipped., cauda pale with wax at end; BL 1.22-1.67 mm. Alatae have 4-8 secondary rhinaria on ANT III. On spikelets of Cyperus spp. in Java (Noordam 2004).

Carolinaia justiciae Shinji Apterae are yellowish, BL unrecorded. Alatae have c.8 secondary rhinaria on ANT III. On Justicia procumbens var. leucantha in Japan (Kyushu, Saitama, Morioka; Shinji 1924). The generic position is uncertain, and it is regarded as a nomen dubium by Remaudière & Remaudière (1997).

Carolinaia (Juncomyzus) nigra (Miyazaki) Apterae are elongate oval, blackish brown; BL 1.8-2.0 mm. On Toxicodendron trichocarpum in Japan (Miyazaki 1971). Probably heteroecious, but the secondary host is unknown; oviparae, an alate gynopara and an alate male were collected on T. trichocarpum in September (Remaudière & Muñoz Viveros 1993).

Carolinaia (Juncomyzus) obscura (Hille Ris Lambers) Colour of apterae in life is unrecorded; BL 1.2-1.8 mm. On Juncus effusus, colonising leaves and stems near ground, often in ant shelters (Miyazaki 1971). In Japan, and (possibly) Thailand (Remaudière & Muños Viveros 1993).

Carolinaia (Glabromyzus) rhois (Monell) Plate 12a Apterae on Rhus are yellow-brown, pale rusty brown or greenish yellow with black clavate siphunculi (see influential points/gallery); BL 1.9-2.3 mm. In spring colonies occur on undersides of leaves of Rhus glabra and Rh. typhina (= hirta). Alatae have a brownish yellow or greenish yellow abdomen with dusky dorsal markings. Host alternation occurs to flowerheads of Poaceae (e.g. Avena, Dactylis, Elymus, Hordeum, Phleum, Triticum). Apterae on secondary hosts are small tomedium-sized , light brown to greenish yellow with black siphunculi. Widely distributed in North America. Records of G. howardii on cereals should probably be referred to C. rhois, following Cook (1984).

Carolinaia rhusifoliae (Richards) Apterae are shining black with brownish head and prothorax, antennae pale basally and dark distally, legs pale, siphunculi dark; BL c. 1.8 mm. On undersides of mature leaves of Rhus radicans in Ontario, Canada.

Carolinaia (Glabromyzus) schlingeri (Hille Ris Lambers) Colour of apterae in life unknown, probably rather dark; BL 1.4-1.7 mm. Alatae have a large dark dorsal abdominal patch. Described from Toxicodendron sp. (probably diversilobum) in California. The life cycle is unknown, possibly migrating to Poaceae or Cyperaceae; specimens from Luzula spp. in British Columbia, Canada (BMNH collection, leg. C.-k. Chan) may be the secondary host form of this species. Oviparae occur on Rhus in October (BMNH collection, leg, D. Hille Ris Lambers).

Carolinaia (Juncomyzus) scirpi (van der Goot) Apterae are shiny black, brownish black or greenish brown, BL c. 2 mm. Alatae have 10-19 secondary rhinaria on ANT III; the alata described by van der Goot (1917) under this name was probably that of C. javanica – see Noordam (2004). On Scirpus grossus in Java, feeding on the flower stalks and beneath the inflorescences. There are also records from Cyperus spp. and Carex baccans in Java (Noordam 2004). The life cycle is unknown. Apterae from Scleria elata in Assam, India, described under the name of Aulacorthum scirpi van der Goot (A.K. Ghosh & Raychaudhuri 1972), are a different species, probably correctly placed in Aulacorthum (q.v.).

Carolinaia (Glabromyzus) setariae Remaudière & Muños Viveros Apterae are dark brown, with black siphunculi (sometimes pale basally) and pale cauda; BL 1.4-1.9 mm. Alatae have 4-7 secondary rhinaria on ANT III. On a Setaria species in Mexico, and subsequently recorded from S. geniculata (= parviflora) in Brazil (BMNH collection, leg. C.L. Costa).

Carolinaia tissoti Remaudière & Muños Viveros Apterae are yellowish brown with antennae, legs, siphunculi and cauda mainly pale (Tissot 1933, as rhois); BL c. 1.8 mm. Alatae are very dark green to black and have 6-10 secondary rhinaria on ANT III (Tissot 1928, as rhois). Apparently it is heteroecious holocyclic with a similar life cycle to C. caricis, and the apterae of the two species on Carex are practically identical (Remaudière & Muñoz Viveros (1993). In eastern USA.

Casimira Eastop

Aphidinae: Aphidini

A genus for two or three species close to Aphis but with all first tarsal segments bearing two hairs (no medial sense peg), and abdominal segment 7 lacking the usual marginal tubercle. Alatae also differ in forewing venation from typical Aphis, having a once-branched media like Toxoptera. An undescribed Casimira sp. on Ozothamnus leptophyllus in New Zealand was included in molecular phylogenetic studies by von Dohlen & Teulon (2003); however their results placed this species within the genus Aphis. Further information about this species was provided by Teulon et al. (2013). The host of an Indian species (bhutanensis A.K. Ghosh, Basu & Raychaudhuri) also described in Casimira was unidentified.

Casimira canberrae (Eastop) Apterae are black, wax-powdered; BL 1.0-1.7 mm. Alatae have 1-4 secondary rhinaria on ANT III. On young leaves at growing points of Epilobium spp. in eastern Australia (Canberra, New South Wales; Eastop 1966). Hales et al. (2014) described the alate vivipara and ovipara. Monoecious holocyclic with oviparae in May; males are undescribed. 2n=8.

Catamergus Oestlund

Aphidinae: Macrosiphini

Two North American species related to Macrosiphum but with relatively small thin siphunculi.

Catamergus fulvae (Oestlund) Plate 26b (Fig.33g) Apterae are green, powdered with wax (see influential points/gallery); BL 2.3-2.8 mm. On lower parts of Impatiens spp. in north-eastern and midwestern USA, and eastern Canada (Ontario, Quebec, Newfoundland). Monoecious holocyclic with oviparae and alate males in September (BMNH collection, leg. J.P. Sijpkens).

Catamergus kickapoo (Hottes & Frison) Apterae are whitish green with dusky brown tips to appendages (see influential points/gallery); BL 1.9-2.2 mm. On undersides of leaves of Polygonatum spp. in eastern USA and Canada. Monoecious holocyclic with oviparae and alate males in August-September (Hottes & Frison 1931). 2n=10.

Cavariella del Guercio

Aphidinae: Macrosiphini

A holarctic genus of about 38 species, about half of them in Asia. Characteristic of the genus are the two hairs close together on abdominal tergite 8, placed on a prominent tubercle or supracaudal process. In alatae this tubercle is generally less conspicuous than in apterae, but its presence is indicated by the close proximity of the two hairs. Alatae have broad dark bars on the dorsal abdomen, often coalesced into a solid patch on abdominal tergites 3-5. Most species host-alternate between Salix and Umbelliferae/Apiaceae or the closely-related Araliaceae, one is known to be monoecious on Salix, and several others are recorded only from Salix with life cycle unknown. Molecular phylogenetic studies have now shown that Cavariella is allied to Pterocomma, despite the considerable differences in morphology. However species of subgenus Cavariellinepicauda have large abdominal marginal tubercles of similar size and distribution to those of Pterocomma. Parasitoids are Aphidius salicis (Starý 1973) and Ephedrus salicicola (D. Raychaudhuri 1990). Taxonomic revisions and/or keys are available for Europe (Hille Ris Lambers 1947b), Fennoscandia and Denmark (Heie 1992), UK and Ireland (Blackman 2010), Siberia (Ivanoskaya 1980), Siberia (Pashchenko 1987b), Japan (Miyazaki 1971), India (Raychaudhuri 1980) and North America (Stroyan 1969). See also Blackman & Eastop (2006).

Cavariella aegopodii (Scopoli) Willow-Carrot Aphid. Plate 15h (Fig.10b, (fig. 107H, fig. 108J) Apterae are green or yellowish green (see influentialpoints.com/Gallery); BL 1.0-2.6 mm. On leaves and umbels of numerous genera and species of Umbelliferae (incl. Aegopodium, Anethum, Angelica, Anthriscus, Apium, Carum, Chaerophyllum, Cicuta, Crithmum, Cryptotaenia, Daucus, Ferula, Foeniculum, Heracleum, Laserpitium, Lomatium, Oenanthe, Sium, Smyrnium). A major cosmopolitan pest of cultivated Umbelliferae/Apiaceae (Blackman & Eastop, 2000, p.253). Heteroecious holocyclic, with sexual phase on various Salix spp., or anholocyclic on Umbelliferae in warmer climates. Apterae in spring colonies on young leaves and catkins of Salix spp. are oval, somewhat dorsoventrally flattened, pale green, greenish-yellow or yellowish-white; BL 1.5-2.0 mm. Alatae produced from the second generation onwards have black head and thorax and a dark green dorsal abdominal patch, mostly migrating to secondary hosts in May-June, but sometimes recolonising willow, so that populations can persist on willow into July and August (Dunn 1965). The return migration from Umbelliferae/Apiaceae to willow occurs in late September to early November. Preferred primary hosts seem to be S. fragilis and S. alba; some Salix spp. seem only to be colonised in spring, by alatae from overwintered anholocyclic populations, and are thus acting as secondary hosts. Kundu & Dixon (1995) carried out experimental studies on this species that established the adaptive nature of aphid host alternation. [C. llhasana Zhang, described from apterae and alatae on Medicago sativa (unlikely to be the true host) in Tibet, shows no clear morphological differences from C. aegopodii, and may be a synonym. However, the absence of records from carrots in eastern Asia, which are a favoured host of C. aegopodii, suggests that a different species may indeed occur there.] 2n=10.

Cavariella angelicae (Matsumura) (Fig.10e) Apterae are yellowish green; BL c.2.1 mm. On various Umbelliferae/Apiaceae (Angelica, Heracleum, Pastinaca, Sium) in Japan, Korea and east Siberia, and recently recorded from Kazakhstan (Kadyrbekov 2012a; on Libanotus buchtormensis) and western Siberia (Altai Republic; Stekolshchikov & Novgorodova 2013). Alangium platanifolium (var. macrophyllum) is also recorded as a host in Korea (Paik 1965), but these were probably vagrants. Heteroecious holocyclic; the primary host generations are undescribed, but Lee et al. (2002) recorded a collection from Salix babylonica in North Korea.

Cavariella (Cavaraiellia) aquatica Gillette & Bragg Plate 15i (Fig 35a,b, fig. 108H) Apterae in spring colonies on Salix app. (especially lapponum, nigricans, purpurea, urbaniana) are narrow-bodied, green, BL 1.4-2.0 mm. Fundatrices are large (BL 2.0-2.7 mm) and remarkable for this genus in their complete lack of siphunculi. Alatae have a blackish head and thorax and dark dorsal abdominal bands sometimes partly fused into a solid patch; they migrate to found summer colonies on plants growing in water or marshy situations. Unusually for this genus, a wide variety of secondary hosts, mainly in Poaceae, Cyperaceae and Juncaceae, but also some dicots, may be colonised. Apterae on secondary hosts are pale yellowish green, wax-dusted underneath; BL 1.3-2.6 mm. The return migration to Salix occurs in September. C. aquatica is recorded, often at high altitudes, from Europe, Iran, Kazakhstan (Kadyrbekov 2017a), Afghanistan, northern India, Pakistan, east and west Siberia, China (as C. zhangi Zhang, Chen, Zhong & Li, in Zhang 1999), USA (Colorado) and Canada (Manitoba, North-West Territories). Remaudière (1970) gave a full account of this species. 2n=8.

Cavariella araliae Takahashi (fig. 107C) Apterae are pale yellow or pale green; BL 1.3-2.1 mm. On young stems and shoots of Araliacae (Aralia, Schleffera, Tetrapanax), and also recorded from Broussonetia kazinoki (Lee et al., 2002c). Presumably host-alternating between Salix spp. and Araliaceae, but this needs to be confirmed, as no fundatrices or sexual morphs have been described and the only Salix host identified to species is S. warburgii, in Taiwan, on which populations overwinter anholocyclically along the midribs of the leaves (Takahashi 1923; as neocapreae). Recorded from China, Taiwan, Vietnam (Yoshitomi et al. 2015), Japan, Korea, Okinawa and east Siberia. It seems possible that the aphids from Salix in India described as C. biswasi (q.v.) are the primary host forms of araliae. 2n=14.

Cavariella archangelicae (Scopoli) (Fig.10c,f) Apterae in spring colonies on Salix spp. are pale yellow-green to straw-coloured, those on secondary hosts are green or yellowish (see influentialpoints.com/Gallery), with siphuncular wax (discharged by prodding abdomen) colourless to slightly greenish (cf. konoi); BL 1.5-2.6 mm. Alatae have black head and thorax and broad dark green cross-bands on the dorsal abdomen, sometimes merged into a solid patch; migration occurs to Angelica spp. (including Archangelica) from late May to July. Certain other Umbelliferae/Apiaceae may sometimes be colonised (e.g. Apium, Heracleum, Myrrhis, Peucedanum, Pimpinella). Oviparae and alate, reddish males may be found back on Salix in October. Stroyan (1964a) discussed differences between archangelicae and the very similar C. konoi. In Europe (including Iceland), North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), Iran (Rezwani et al. 1994), Kazakhstan (Kadyrbekov 2009b), and introduced to western USA (Utah, Alaska; see Foottit et al. 2006). 2n=6.

Cavariella aspidaphoides Hille Ris Lambers (fig. 107I) Apterae on Salix in spring are green, elongate pear-shaped with pointed abdomen; BL 1.7-2.2 mm. Often they form mixed colonies with C. aegopodii. Alatae migrate in May to Umbelliferae/Apiaceae; the only recorded secondary host is Daucus carota ssp. maximus. Apterae on Daucus are probably green, BL c.1.3-1.5 mm. Only known from Israel and Iran (Hille Ris Lambers 1970c).

Cavariella bhutanensis Chakrabarti & Das Apterae on an unidentified Salix sp. in spring are broadly spindle-shaped, colour in life is unrecorded; BL 1.7-2.1 mm. Alatae have extensive dark dorsal sclerotisation with broad cross-bands coalesced into a patch on abdominal tergites 2-5. In Bhutan (Chakrabarti & Das 2009). The life cycle is unknown.

Cavariella biswasi Ghosh, Basu & Raychaudhuri Apterae in spring on Salix spp. (including babylonica, elegans) are pale (green?), spindle-shaped, BL 1.7-1.9 mm. Alatae, produced in July, have broad dark partially coalesced bands on abdominal tergites 3-5, and smaller sclerites on other tergites. In north-east India. The secondary host generations have not been described, although some apterae collected on Heracleum candicans in Kashmir were identified as this species (Bhagat 1981b). It could however be the primary host form of C. araliae (q.v.), from which it differs only in the size and form of the process on abdominal tergite 8 (A.K. Ghosh et al. 1969a).

Cavariella borealis Hille Ris Lambers (fig. 107F) Monoecious holocyclic on Salix arctica in north-west Greenland and northern Canada (North-West Territories), with an abbreviated cycle, producing oviparae and apterous and/or alate males in July. Colour of apterae in life not recorded; they are very variable in siz and form, with BL 0.7-2.3 mm. Alatae can be brachypterous, intermediate or fully-winged (Hille Ris Lambers 1960b). 2n=6*.

Cavariella bunii Narzikulov & Mukhamediev Apterae are green, BL 1.8-2.3 mm. On Bunium persicum and Bunium sp. in Tajikistan (Alayskiy Khrebet mountains), and also recorded (from Angelica decurrens) in Kazakhstan (Kadyrbekov 2018a). Not clearly distinct from C. aegopodii.

Cavariella (Cavaraiellia) cessana Zhang, Chen, Zhong & Li Apterae are green; BL c.1.6 mm. Alatae have 11-12 large round rhinaria on ANT III. On an unidentified species of Salix at 2500m altitude in Xinjiang province, China (G. Zhang 1999). Closely related to, and possibly synonymous with, C. aquatica.

Cavariella cicutae (Koch) Apterae are shining yellowish with two longitudinal green stripes; BL 1.5-2.1 mm. Living without host alternation in large colonies on upper parts of stems of aquatic plants in several genera of Umbelliferae/Apiaceae (Berula, Cicuta, Sium) in Europe (Hille Ris Lambers 1947b, 1952). Records of host alternation and a sexual phase on Salix in Belarus, Ukraine and Iran are probably referable to C. salicicola (q.v.), apterae of which on its secondary hosts are not clearly distinguishable from those of cicutae. Records from Salix in Italy (Barbagallo & Stroyan 1982; Barbagallo & Patti 1998; Barbagallo et al. 2011) are likely to be C. rutila Mamontova, which was erroneously synonymised with C. cicutae but has longer siphunculi similar to those of C. aegopodii, and no dorsal subapical hair on the cauda. [Secondary host forms of C. rutila have not been described, although Chumak (2004) records them from Anthriscus and Levisticum in Ukraine; however, these could be salicicola.] 2n=10

Cavariella (Cavariellinepicauda) cicutisucta Qiao Apterae pale, powdered with wax in life; BL 2.2-2.7 mm. Alatae have broad dorsal abdominal cross-bands. On undersides of leaves of Cicuta virosa in Shanxi Province, China (Qiao et al. 2005c). Life cycle is unknown.

Cavariella digitata Hille Ris Lambers (fig. 107A, fig. 108A) The appearance in life is unknown, apterae are probably pale yellowish or greenish; spindle-shaped, rather flattened, BL 1.7-2.3 mm. Alatae have a dusky, somewhat perforated, dorsal abdominal sclerotic patch. On unidentified Salix sp(p)., described from Utah, USA (Hille Ris Lambers 1970c), and subsequently recorded from Oregon, Idaho, Wyoming and New Mexico, and from Alberta, Canada (aphidtrek.org). A record from Turkey (Görür et al. 2011a) is doubtful and needs confirmation. The life cycle is unclear; populations persist on Salix into August, suggesting that if host alternation occurs at all then the migration must be incomplete. Alate males occur in October.

Cavariella gilgiana Zhang, Chen, Zhong & Li Colour of aptera in life is unrecorded, probably pale green; BL c.1.0-1.1 mm. Apterae have 5-segmented antennae with PT/BASE less than 0.4. Alatae have 5-7 large round rhinaria on ANT III, and 0-1 on IV, and a very short antennal PT like the apterae. On an unidentified species of Salix in Ningxia province, China (G. Zhang 1999).

Cavariella gilibertiae Takahashi Apterae are milky white to yellowish brown; BL 1.8-2 mm. On Dendropanax spp. in Japan and Korea. Also collected in Korea on Miscanthus purpurascens (Lee et al. 2002c.)

Cavariella hendersoni Knowlton & Smith Appearance in life is unknown. On leaves of an unidentified Salix sp. in Utah, USA. The life cycle is unknown. The original description was of alatae only, which resemble those of C. salicis (Monell) in having secondary rhinaria on ANT IV and dorsal abdominal cross-bands separate or only partially coalesced. BL of alate paratypes is 1.7-1.9 mm (Palmer 1952). Differences from salicis include the usual presence of accessory hairs on R IV+V and more slender siphunculi. Stroyan (1969) described some characters of presumed apterae of this species from Logan Canyon, Utah, and its identification in the key is based on these specimens, now in the BMNH collection. There are records of hendersoni from seven other states of USA, and from Ontario, Canada (Smith & Parron 1978), but most of these are likely to be misidentifications (see Stroyan 1969).

Cavariella (Cavariellinepicauda) heraclei Takahashi Apterae are white; BL 2.3-3.0 mm. On Angelica and Heracleum spp. in Japan (Miyazaki 1971).

Cavariella (Cavariellinepicauda) hidaensis Takahashi Apterae are milky white or pale yellow-green, slightly pale brownish at apices of antennae and legs; BL c. 1.5 mm. On an unidentified Salix sp. in Japan (Takahashi 1961c), and also recorded from S. udensis in Kamschatica (Pashchenko 1988a). Another Salix feeder described in Elatobium, E. salicifoliae Zhang from Salix sp. in China, is said to resemble C. hidaensis but have a smooth tergum and shorter, broader R IV+V (Zhang & Zhong 1985).

Cavariella himachali L.K. Ghosh Apterae are greenish with two deep green streaks along sides of abdomen; BL c.2.1 mm. On apical shoots of Spiraea canescens and Indigofera gerardiana (= heterantha) in north-west India (Himachal Pradesh). The host plants are unlikely for a Cavariella, and require confirmation. (Note that Nudisiphon folisacculata with similar siphunculi is also recorded from Spiraea and ?Indigofera in this region.)

Cavariella indica Maity & Chakrabarti Appearance in life is unknown, probably pale green; BL of aptera c.1.9-2.1 mm. Alata is undescribed. In colonies on upper surfaces of leaves and on apical growth of Salix babylonica in Uttar Pradesh, India (Maity, Bhattacharya & Chakrabarti 1982), and also recorded (on Salix sp.) in Kazakhstan (Kadyrbekov 2017a). Monoecious holocyclic, with oviparae and males in October-December (Medda et al. 1990). On the basis of the published description, indica seems morphologically very similar to C. aegopodii, except that R IV+V of the aptera is reported to have two accessory hairs.

Cavariella intermedia Hille Ris Lambers Apterae are evenly light green, rather broadly spindle-shaped, BL 1.4-2.3 mm. Alatae differ from aegopodii by having only a rather indistinct smoky dorsal abdominal patch due to pale, partly coalescent sclerotic bars on abdominal tergites 3-6 (becoming somewhat darker in alatae developing at lower temperatures). Alatae also differ from aegopodii by having 2-7 secondary rhinaria on ANT IV. Recorded from Salix aurita, S. arbuscula and S. phylicifolia in Belgium, Netherlands, Switzerland and UK, and also from S. glauca and S. viminalis in Kazakhstan (Kadyrbekov 2012a, 2017a) and a Salix sp. in west Siberia (Stekolshchikov et al. 2008). Life cycle in not completely known; Hille Ris Lambers (1970a) found apterae producing oviparae and alatoid nymphs (presumed males) on Salix in September, which seems to rule out host alternation. 2n=6.

Cavariella japonica (Essig & Kuwana) (Fig.10h, fig. 107D, fig. 108E) Host-alternating between Salix spp. and various Umbelliferae/Apiaceae (Angelica, Anthriscus, Cryptotaenia, Dystaenia, Ledebouriella, Pastinaca, Peucedanum, Torilis). Apterae on Salix in spring are dusky yellowish, pinkish, purplish or greyish-brown, with dark distal sections to antennae, legs and siphunculi; BL 1.5-2.4 mm. Takahashi (1937b) also recorded a wholly black form of aptera from high altitudes in Taiwan, on S. fulvipubescens. Alatae have a dark orange-yellow abdomen with a black dorsal patch. Apterae on secondary hosts are variably coloured, yellowish to brownish. In Japan, Korea and east Siberia. In South Korea many samples have been collected also from Ranunuculus japonicus (= grandis) (Lee et al. 2002c). Sexuales are undescribed. 2n=8.

Cavariella konoi Takahashi (Fig.10d,g, fig. 107E) Host-alternating between Salix spp. and Umbelliferae/Apiaceae. Apterae in spring colonies on Salix are green or yellowish-green, often with two darker green longitudinal stripes (see influentialpoints.com/Gallery); BL c.2.3-2.5 mm. Alatae have a solid dark dorsal abdominal patch on abdominal tergites 3-5 and two characteristic small roundish spots close together on abdominal tergite 6, posterior to the level of the bases of the siphunculi. Other differences from the biologically and morphologically similar C. archangelicae were discussed by Stroyan (1964a), with descriptions also of males and oviparae. Apterae on secondary hosts are green, or yellow-green with faint darker green longitudinal stripes, with the siphuncular wax distinctly yellow (cf. archangelicae); BL 1.6-2.9 mm. On several genera of Umbelliferae (Angelica, Apium, Cicuta, Conioselinum, Foeniculum, Myrrhis, Peucedanum, Sium), and also recorded from Tetrapanax (Araliacae). Recorded from Europe (including Iceland), North America, Mexico, Hawaii, Japan, China, Korea and Siberia. Records from India (Himachal Pradesh; L.K. Ghosh 1986) perhaps need additional confirmation. In Europe the only secondary hosts seem to be Angelica spp., whereas in Japan Heracleum is also colonised (Miyazaki 1971), and in America secondary host populations are recorded on Angelica, Apium, Cicuta and Sium (Stroyan 1969). Stroyan (1969a) discussed North American records and geographical variation, and considered that records of C. hendersoni Knowlton & Smith from Cicuta maculata and Sium suave should be assigned to C. konoi (Stroyan 1969a). 2n=8.

Cavariella (Cavariellinepicauda) largispiracula Zhang, Chen, Zhong & Li Colour of apterae in life is unrecorded; BL 1.6-2.7 mm. On Angelica sinensis in Gansu province, China (G. Zhang 1999). Cavariella yuzhongensis seems to be a synonym (Eastop & Blackman 2005).

Cavariella nigra Basu Apterae in spring colonies on Salix are dusky black including antennae, legs, siphunculi and cauda, pear-shaped, somewhat flattened; BL 1.9-2.5 mm. Alatae have a dark brown central dorsal abdominal patch. The black coloration of the aptera is unusual for a Cavariella; the dark form of japonica reported from Taiwan is possibly this species, although japonica from Japan has a longer last rostral segment (R IV+V 1.4-1.6 × HT II, as opposed to R IV+V 1.0-1.2 × HT II in nigra) C. nigra is also unusual in having 0-1 secondary rhinaria near the middle of ANT V of the aptera (none on III or IV). On undersides of leaves of Salix elegans in West Bengal, India (A.N. Basu 1964). The life cycle is unknown.

Cavariella nigrocaudata Takahashi Apterae are dark brownish with appendages mainly dark; BL c.2 mm. On Tetrapanax papyriferus in Japan.

Cavariella nipponica Takahashi Apterae are whitish; BL c.1.6 mm. Alatae have dark dorsal abdominal cross-bands fused into a solid patch on abdominal tergites 3-5. On various Umbelliferae/Apiaceae (Angelica, Heracleum, Pachypleurum) in east Asia (China, Japan, Korea, east Siberia). In South Korea many samples have been collected also from Ranunuculus japonicus (= grandis) (Lee et al., 2002c). Apparently host-alternating with Salix as primary host, although the life cycle does not appear to have been studied and no sexuales have been described (Takahashi 1961c, Miyazaki 1971, Pashchenko 1988a).

Cavariella (Cavariellinepicauda) oenanthi (Shinji) (Fig.10i) Apterae are pale green with dark head and very variably developed dark dorsal markings; BL 1.5-2.1 mm. On several genera of Umbelliferae (Angelica, Cryptotaenia, Oenanthe), on stems at ground level, often in ant shelters (Miyazaki 1971). In Japan, Korea, east Siberia and Central Asia (Tajikistan; Narzikulov 1975). 2n=8.

Cavariella pastinacae (Linnaeus) (fig. 108C) Apterae in spring colonies on leaves and new growth of Salix spp. are rather shiny light green, with antennae mainly pale (darker on base VI), and legs pale except for tarsi; BL 1.6-2.2 mm. Alatae have very broad dark bands on abdominal tergites 3-5 coalescing to form an almost solid dark green to black trapezoid patch (see influentialpoints.com/Gallery). Host alternation occurs mainly to Heracleum, less commonly to Pastinaca, but there are also records from Angelica, Carum, Chaerophyllum, Cicuta, Foeniculum and Torilis. Apterae on secondary hosts are pale green, BL 1.8-2.9 mm. Oviparae and males are found back on Salix in October. C. pastinacae occurs throughout Europe and in south-west and Central Asia (Kazakhstan; Kadyrbekov 2005c), and is also widely distributed in North America, as well as being recorded from Argentina (Nieto Nafrìa et al. 1994). C. kamtshatica Ivanoskaya, recorded from Angelica and Heracleum in Kamchatka, is not clearly separable from its published description and could be large pastinacae, although specimens also collected by Ivanoskaya in west Siberia and identified by her as C. pastinacae are reported to be C. kamtshatica by Stekolshchikov & Khureva (2020). 2n=8.

Cavariella pustula Essig Apterae are pale greenish-yellow to orange, with siphunculi and distal extremities of antennae and legs dusky; BL about 1.6 mm. Alatae have black head and thorax and pale greenish-yellow abdomen with two light green dorsal longitudinal stripes (Essig 1937). On new growth and undersides of leaves of Salix argophylla (= S. exigua; BMNH collection; leg. R.C. Dickson) in California, and also recorded from unidentified Salix in California, Utah, Oregon and British Columbia (BMNH collection). Life cycle unknown.

Cavariella rutila Mamontova (fig. 107G) Apterae in spring colonies on leaves and new growth of Salix purpurea are reddish or bright yellowish with red markings; BL c.1.8-1.9 mm. Alatae are reddish-yellow with broad dark brown cross-bands on dorsal abdomen (Mamontova 1961). It is also recorded from S. cinerea and S. silesiaca, and from Estonia, Latvia, Ukraine and Greece. Records of C. cicutae from Salix spp. in Italy (Barbagallo & Stroyan 1982; Barbagallo & Patti 1998; Barbagallo et al. 2011) may also be referable to this species. The life cycle is unknown.

Cavariella salicicola Matsumura (Fig.10a, fig. 108F) Apterae in spring colonies on new growth of Salix are yellowish-green to green with bluish pruinosity (Moritsu 1983); BL c.2.0-2.2 mm. Immature alatae are bluish-grey with paired dark segmental markings, adult alatae have black head and thorax abd variably developed dark dorsal abdominal cross-bands or pleural markings. Recorded from numerous Salix spp. and also from Chosenia arbustifolia in east Asia (India, Japan, Korea, China, Tibet, Thailand and Siberia), and reports of C. cicutae host-alternating between Salix and Umbelliferae/Apiaceae in Belarus (Buga & Stekolshchikov 2012), Ukraine (Mamontova-Solukha 1961) and Iran (Hodjat 1984a) should probably be referred to C. salicicola. Osiadacz & Wojciechowski (2008) recorded it from Salix triandra in Poland, and Kadyrbekov (2014g, 2017a) recorded it from Salix spp. in Russia (Tyumen) and Kazakhstan. Migration occurs in April-May to various Umbelliferae/Apiaceae (incl. Apium, Cryptotaenia, Oenanthe, Levisticum, Sanicula, Sium), with a preference for plants growing in water or marshy situations. Apterae on secondary hosts are green with pale bluish wax markings; BL 1.7-2.2 mm. Sexuales are found on Salix in late October (Wang et al. 1988). The complete mitochondrial genome of this species (16,317 bp) has been ascertained (Wang et al. 2013). 2n=10 (Chen & Zhang 1985b).

Cavariella salicis (Monell) (fig. 108I) Apterae on Salix, according to Monell’s original notes, are green with two darker green longitudinal stripes on the abdomen (Stroyan 1969); BL 1.6-2.1 mm. Alatae have dusky head and thorax and green abdomen with irregular darker green markings. In east and central USA. A record from Turkey (Akyürek et al. 2019) is doubtful and requires confirmation. There is at least a partial migration to Umbelliferae/Apiaceae (Angelica, Oenanthe, Sium). Stroyan (1969a) clarified the status of salicis, which was for many years confused with aegopodii and other species.

Cavariella (Cavariellinepicauda) sapporoensis Takahashi Apterae are whitish; BL c. 3.0 mm. On Angelica and Heracleum spp. in Japan (Miyazaki 1971) and Korea, where many samples have been collected also from Arisaema amurense var. serratum (Lee et al. 2002c).

Cavariella saxifragae Remaudière Apterae are blackish; BL 1.4-1.8 mm. Alatae have secondary rhinaria distributed III 13-24, IV 0-2. Feeding is restricted to Saxifraga aizoides. In France (Pyrenees) and Scotland (Stroyan 1964a). Monoecious holocyclic with oviparae and apterous (alatiform) males (original description). The male genitalia were described and illustrated by Wieczorek et al. (2011).

Cavariella simlaensis Chowdhuri, R.C. Basu & Raychaudhuri Apterae are presumably dark brown or blackish in life; BL c.2.3 mm. Alatae have secondary rhinaria distributed III 48-65, IV 8-13, V 2-6. On Sanicula europaea in India (Himachal Pradesh). This is possibly the secondary host form of C. nigra (q.v.).

Cavariella takahashii Hille Ris Lambers As only the alata of this species is described, it could not be included in the key to Cavariella on Salix. BL of alata about 2.2-2.3 mm; dorsal abdominal markings are indistinct, and the femora have numerous long wavy hairs as well as a number of much shorter hairs on distal half similar to those on tibiae. The antennae have a short antennal terminal process and numerous secondary rhinaria; 51-52 on ANT III, 10-11 on IV and 3-5 on V. On Salix spp. in April in Japan (Hille Ris Lambers 1965). Appearance in life and biology are unknown.

Cavariella theobaldi (Gillette & Bragg) (fig. 107B, fig. 108B) Host-alternating between Salix spp. and Umbelliferae/Apiaceae. Apterae in spring colonies on leaves and new growth of Salix spp. are yellowish-green to green; BL 1.6-2.0 mm. Alatae have variably-developed dark dorsal abdominal cross-bands, often forming a central trapezoid patch. They fly in late April-June to secondary hosts (Heracleum, Pastinaca, or more rarely Aegopodium, Angelica, Chaerophyllum). Apterae on secondary hosts are rather bright green, with tapering siphunculi (see influentialpoints.com/Gallery); BL 1.8-2.8 mm. The return migration occurs in late September-October. In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), Turkey, Iran, east and west Siberia (Pashchenko 1988a), Kazakhstan (Kadyrbekov 2017a) and north-eastern North America. The male genitalia were described and illustrated by Wieczorek et al. (2011). 2n=8.

Cedoaphis Oestlund

Aphidinae: Macrosiphini

Two North American species with secondary rhinaria in apterae and a helmet-shaped cauda. The two have different biologies and different cuticular sculpture, and are unlikely to be truly congeneric.

Cedoaphis incognita Hottes & Frison Apterae are yellowish green with dusky brown head; BL 2.0-3.0 mm. Alatae have secondary rhinaria distributed III 22-28, IV 15-18, V 10-14. On Symphoricarpos spp., feeding on leaves in summer, and on roots (fundatrices) in spring. Widely distributed in USA. Monoecious holocyclic, although sexual morphs are apparently undescribed (Palmer 1952, as Aphis).

Cedoaphis maxsoni (Palmer) Plate 11f Apterae are pale amber with variably developed dark dorsal patches; BL 1.3-2.7 mm. Alatae have secondary rhinaria distributed III 30-46, IV 18-20, V 1-5. Described from small specimens (BL 1.3-1.7 mm) on roots of Eriogonum sp. in late summer in Colorado, USA. Subsequently much larger specimens (BL 2.3-2.7 mm) were collected in spring from Compositae/Asteraceae (Petasites, Senecio) in Oregon (BMNH collection, leg. D. Hille Ris lambers) and California (BMNH collection, leg. R.C. Dickson). Oviparae were found on Eriogonum in late August (Palmer 1952, as Amphicercidus).

Cerataphis Lichtenstein

Hormaphidinae: Cerataphidini

Eight south-east Asian species, at least three of which host-alternate between galls on Styrax and monocotyledonous secondary hosts (Araceae, bamboos, Orchidaceae, Palmaceae, Pandanaceae) in their native regions. Several species are more widely distributed by commerce on their palm and orchid secondary hosts. Most species (but not C. bambusifoliae) have unusual yeast-like extracellular symbionts, found also in Tuberaphis and Glyphinaphis (Fukatsu et al. 1994, Xu et al. 2018)). Apterae on secondary hosts are dark, round, scale-like, with a marginal ring of wax. Alatae have a once-branched media and 5-segmented antennae. Noordam (1991) provided a generic diagnosis and account of Javanese species. Qiao et al. (2018) reviewed and keyed the species in China. Blackman et al. (2019b) reviewed and keyed the species introduced to western Europe. Russell (1996) provided a key to the species on palms and orchids. Fukatsu et al. (1994) studied their phylogenetic relationships with other Cerataphidini.

Cerataphis bambusifoliae Takahashi (fig. 30A) Apterae on stems and undersides of leaves of bamboos (Arundinaria, Bambusa, Chimonobambusa, Fargesia, Pleioblastus, Sinarundinaria) are black, aleyrodiform, with a white wax fringe; BL 1.5-2.5 mm. Qiao et al. (2018) also record it from two species of Allium. Alatae have antennae with annular secondary rhinaria distributed III 24-34, IV 9-13, V 8-13. Alatae sexuparae are produced in large numbers in September-December (Takahashi 1939, Saha & Chakrabarti 1988b, Kurosu et al. 1996), but the primary host generations were unknown until recently, when galls found on Styrax suberifolius in a mountainous region of Taiwan were linked to C. bambusifolae by studying morphology of the first instars produced by emigrant alatae and by comparison of mitochondrial DNA sequences (Kurosu et al. 2008). The galls are elongated fig-shaped, up to 9 cm long, and thin-walled, at the tips of lignified twigs. Many second instar sterile soldiers were found in the galls, and alatae were present in June. The species is known from China, Taiwan and Uttar Pradesh, India.

Cerataphis brasiliensis (Hempel) (= fransseni Hille Ris Lambers, = variabilis Hille Ris Lambers, = palmae Ghesquière of many authors) The gall of this species on Styrax benzoin is bag-like, greyish-green, 1.0-4.7 cm long, with an apical slit-shaped opening. It arises from the stem, either from a terminal or an axillary bud (Stern et al. 1995, as fransseni). Apterae in the galls are small (BL about 1.3 mm) with spine-like frontal hairs including one pair of particularly large ones, short 5-segmented antennae and siphuncular pores on small cones with a ring of 3-5 hairs. Second instar sterile soldier larvae move in and around the galls (Noordam 1991, as fransseni). The dynamics of soldier production was investigated by Stern et al. (1994). Alatae (Plate 3a) emerging in June-August have antennal secondary rhinaria distributed III 15-23, IV 2-8, V 0-4, and BL 1.2-1.6 mm. They migrate to found colonies on palms, the apterae on palms (Plate 3b) being sedentary dark brown flattened, almost circular, fringed with white wax and often with one or more very narrow transverse white stripes across the middle region (see aphids of Karnataka website); BL 1.0-1.9 mm. Palm-feeding populations were previously known as C. variabilis and C. palmae (Stern et al. 1995, Blackman & Eastop 2000). Host alternation occurs in Java and Malaysia. Anholocyclic populations are widespread in the tropics and subtropics on coconut and some other palms including ornamental fan-palms, more rarely on oil palm, with records also from Musa and Cycas (Blackman & Eastop 2006). Josephrajkumar et al. (2011) reported on its preference among coconut varieties in India. In recent years it was reported from the Canary Isles (Pérez Hidalgo et al. 2000), and in a French nursery on ornamental palms (Butia capitata) imported from Brazil (Germain & Chapin 2004). In populations of apterae on palms there is a curious polymorphism first noted by van der Goot (1917; as lataniae), studied further by Noordam (1991; as palmae) and most recently in Brazil by Mews et al. (2008). See http://edis.ifas.ufl.edu/pdffiles/IN/IN92400.pdf for an illustrated on-line account of this aphid. 2n=18.

Cerataphis formosana Takahashi Apterae are black, BL c. 2.1 mm, with marginal wax, recorded as feeding on young leaves of Cocos sp. in Taiwan (Takahashi 1924). Only known from the original description. It apparently differs from other Cerataphis in the shape and position of the frontal horns. Its biology is unknown.

Cerataphis freycinetiae van der Goot Apterae are dull black, aleyrodiform with an encircling flat fringe of white; BL 1.3-1.6 mm. On young leaves of Freycinetia spp. It is also recorded from Pothos scandens, and aphids provisionally identified as this species have also been collected on Poikilospermum sp. (BMNH collection, VFE 16,237). There is also a Chinese record from Archontophoenix (Qiao et al. 2018). Widely distributed in south-east Asia, and also recorded from Australia (Queensland, BMNH collection, leg. J.H. Martin).

Cerataphis jamuritsu (Takahashi) Galls are large (12.0-14.5 cm long), pendulous, flask-shaped, greyish-brown dusted with white powder, broadest at the distal end where there is a very large, slit-like opening (fig. 124B and Aoki et al. 1998). They are found on Styrax suberifolia in Taiwan and China (Hong Kong; BMNH collection). Second instar soldiers guard the outside surface of the gall and may sting by piercing the skin of the intruder. Alatae (BL about 1.5 mm) are mature in the galls in November, flying to its secondary host, the rattan Calamus quinquesetinervius, where colonies have been found in ant shelters in southern Taiwan (Kurosu et al. 2004). The apterae on rattan are dark grey or reddish with an encircling fringe of white wax; BL 2.1-2.4 mm.

Cerataphis lataniae (Boisduval) Apterae are oval, flattened, shiny brown or orange-brown, with a flat white marginal fringe of wax (see aphids of Karnataka website); BL 0.9-1.6 mm. It resembles C. brasiliensis, and the two have been confused in the past, but it lacks the one or more pairs of dagger-like hairs on the underside of the head of C. brasiliensis. Eastop (1966) gave other distinguishing characters. It appears to be widespread on palms, especially Latania and other fan-palms, raffia palms and Cocos nucifera, through the tropics and in glasshouses, and recently introduced to the Canary Isles, where it also colonised Strelitzia alba (Pérez Hidalgo et al. 2000), although its exact distribution is uncertain because of confusion with C. brasiliensis.

Cerataphis orchidearum (Westwood) Orchid Aphid, Scale Aphid (Fig.25a) Apterae are dark reddish brown to black dusted with wax, with a fringe of radiating plates of white wax (see aphids of Karnataka website); BL 1.0‑1.9 mm. On various Orchidaceae, widely distributed in the tropics, and in European and North American glasshouses. Earlier records of C. lataniae from orchids probably all refer to this species. Anholocyclic; the sexual phase is not known. See Heie (1980b) or Zimmerman (1948; under C. lataniae) for further information. 2n = 16 (samples from Cymbidium, Dendrobium and Epidendrum) or 18 (samples from Angraecum, Sarcochilus and Butia).

Cerataphis pothophila Noordam Apterae are brownish black with a flat encircling fringe of white wax, and a fine white transverse stripe across the mid-dorsal region; BL 1.35-1.55 mm. On undersides of leaves of Pothos roxburghii in Java (Noordam 1991).

Cerataphis vandermeermohri Hille Ris Lambers Galls are extensive structures consisting of bunches of long, thin, dichotomously branched tubes attaining 20-35 cm in length, resembling some species of stagshorn fern (fig. 124H). The colony size in a gall can reach 94,000 individuals, of which c. 45% are sterile second instar soldiers (Kurosu & Aoki 1997). On Styrax subpaniculatus (= serrulatus var. mollissimus; Huang et al. 2003) in Sumatra, and galls are also recorded from S. paralleloneurus in China (Qiao et al. 2018). The emigrant alatae, BL about 2.1 mm, were collected in February-March, but the secondary host is still unknown. The generic position of this species, originally described in Astegopteryx, is uncertain, and possibly a new genus should be created for it (see Fukatsu et al. 1994). [N.B. Trichoregma vandermeermohri Takahashi 1935, described from bamboos, is a synonym of Astegopteryx basalis.]

Ceratoglyphina van der Goot

Hormaphidinae: Cerataphidini

Aphids similar to Astegopteryx but distinguishable from most other Hormaphidinae by the rounded cauda and anal plate. The primary host has recently been shown to be Styrax as in other Cerataphidini, but many populations are probably anholocyclic on bamboos. One species was described from Camellia. Noordam (1991) reviewed the genus, Qiao & Zhang (1999a) and Qiao et al. (2018) reviewed and keyed the species in China, and Stern et al. (1997) reviewed life cycles and taxonomic relationships. Chen & Qiao (2012c) studied variation in the wax glands.

Ceratoglyphina bambusae van der Goot Galls on Styrax in Malaya, similar to those of C. roepkei in Sumatra (see below) but less twisted, were identified as housing the primary host generations of C. bambusae by comparison of mitochondrial DNA sequences (Stern et al 1997). The host was determined as Styrax benzoin, but Kurosu (cited in Aoki et al. 2002) subsequently found many galls of this species on the Malay Peninsula on S. fraserensis, so the record from S. benzoin needs confirmation, as does a Chinese record from S. suberifolius (Qiao et al. 2018). Alatae migrate to bamboos (Bambusa, Dendrocalamus, Gigantochloa, Phyllostachys, Schizostachyum). Apterae on bamboos are oval-bodied, dark brown or dark marbled greenish black, with pale brown legs, and a flat fringe of white wax; BL 1.3-2.2 mm. Immatures are yellowish to green, with brown heads in later instars (Noordam 1991). Dense colonies form near bases of leaves, often attended by ants. Collected on bamboos in China, Indonesia and Malaysia, but the galls on Styrax are only recorded from the Malay Peninsula. 2n=12 (Chen & Zhang 1985).

Ceratoglyphina bengalensis L.K. Ghosh Apterae are pale brown with two rather indistinct longitudinal green stripes and a narrow, flat, wax fringe; BL 2.1-2.5 mm. On undetermined bamboos in India (West Bengal, Assam; M. R. Ghosh et al. 1974), and also recorded from Java, where the aphids were found living on the border of the sheath and at the leaf-bases of young shoots (Noordam 1991).

Ceratoglyphina brachyskeroae Qiao & Zhang Appearance in life is unrecorded; BL of aptera c.1.8 mm. On an unidentified bamboo in Fujian province, China (Qiao & Zhang 1999a).

Ceratoglyphina camellis Qiao & Zhang Apterae are black in life, and presumably with little or no wax; BL c.1.7 mm. On Camellia japonica in Guizhou province, China (Qiao & Zhang 1999e, as Ceratocallis). Alatae and life cycle are unknown. [Ceratocallis was synonymised with Ceratoglyphina by Chen et al. (2014).]

Ceratoglyphina phragmitidisucta Zhang Appearance in life is unrecorded; BL of aptera c.1.7 mm. On Phragmites communis (= australis) in Fujian province, China (Qiao & Zhang 1999a). Further collections are required for confirmation that this is the normal host.

Ceratoglyphina roepkei (Hille Ris Lambers) The gall of this species, found on Styrax paralleloneurus in Sumatra, is remarkable; it is composed of thin, spirally-twisted tubular subgalls, 15-20 cm long, their bases radiating from a circular attachment site on one side of the stem near an axil-bud (fig. 124E). Alate sexuparae were found within the gall in February, but adult alatae have also been found in September (Aoki & Kurosu 2010). This species lives without host alternation on S. paralleloneurus (Kurosu & Aoki 1995, as Astegopteryx roepkei). It was transferred to Ceratoglyphina following a phylogenetic analysis of mitochondrial DNA by Stern et al. (1997).

Ceratoglyphina styracicola (Takahashi) The gall generations on Styrax suberifolia were described in Taiwan (Takahashi 1936, as Astegopteryx). The galls are large, almost globular, somewhat like a compact cauliflower head, 55-80 mm in diameter, in clusters arising from the stem (fig. 124J and Kurosu & Aoki 1990a, as C. bambusae). The surface is covered with white wax powder, and the inside is a labyrinth of green plant tissue (Takahashi 1924a, Aoki et al. 1977). Gall inhabitants, which may number 200,000 per gall, include yellowish-brown adult apterae, immatures developing normally, and strongly sclerotised ‘biters’, which do not usually undergo development beyond the second instar (Aoki 1979a, Kurosu & Aoki 1991b, as C. bambusae). Aphids readily wander from these galls and may invade galls of Astegopteryx bambucifoliae on the same plant (Kurosu & Aoki 1990a). Alatae emerge from late December to March of the following year (Kurosu & Aoki 1998) and migrate to Pleioblastus (= Arundinaria) sp.; they have secondary rhinaria distributed III 17-23, IV 6-9, V 4-8. Apterae on bamboos are similar to those of C. bambusae. Sexuparae are produced on the bamboo from end of May to early June (Kurosu & Aoki 1998a). Only recorded from Taiwan.

Ceratopemphigus Zehntner

Eriosomatinae: Pemphigini

One species in Asia related to Prociphilus but with Hormaphidinae-like frontal prominences, very small in adult alatae but well-developed in fourth instars.

Ceratopemphigus zehntneri Schouteden Galls on Ligustrum robustum var walkeri in Sri Lanka are attached to the stem; bulbous, bright green, 4-5 cm in diameter when mature and becoming grey-brown with age. Immature alatae in the galls secrete copious flocculent wax, and adult alatae are wax-dusted. Apterae (BL 2.4-2.8 mm) were described from galls on L. sinense in Hong Kong, China (Qiao & Fang 2007). Cock et al. (2010) provided illustrations of the galls and of all known morphs. The life cycle in unclear; alatae and immatures were also collected in Sri Lanka on Brunfelsia uniflora (Doncaster, 1956), but this is a non-native plant unlikely to be a true host. Only known from Sri Lanka and China.

Ceratovacuna Zehntner

Hormaphidinae: Cerataphidini

A genus of about 20 or more species in east and south-east Asia, closely related to Astegopteryx, and with a similar alternation between Styrax and Poaceae, although this has only been confirmed for two species. Most species form dense colonies on the undersides of leaves of bamboos or large grasses. The apterae are small to medium-sized, yellow to brown in colour and secrete much wax. Some species have horned first instar soldiers as in Pseudoregma. The alatae of many of the bamboo feeders are rare or unknown. As in the case of Astegopteryx, characters used to separate species are frequently those known to vary according to the environment, such as the degree of development of the wax glands, and the genus is in need of further taxonomic revision. Accounts are available for Japan (Takahashi 1958), Java (Noordam 1991), Korea (Paik 1965), India (A.K. Ghosh 1988, Chakrabarti & Debnath 2011), China (Jiang et al. 2015b, 2021, Qiao et al. 2018) and Taiwan (Liao 1976). Stern (1998) analysed mitochondrial DNA sequences to determine the evolution of horned soldiers and relationships to other genera. Phylogenetic relationships of Ceratovacuna and the related genus Pseudoregma based on two mitochondrial and two nuclear genes were studied by R. Zhang et al. (2013). Aoki & Kurosu (2010) reviewed life cycles and biology, and Aoki et al. (2013) used a combination of molecular and morphological data to distinguish species of the C. nekoashi group. Chen & Qiao (2012c) studied variation in the wax glands.

Ceratovacuna angusticorna Qiao Apterae are reddish brown with tufts of white wax; BL 1.8-2.3 mm. On undersides of leaf bases of a plant identified as Indocalamus tessellatus (and also on other unidentified bamboos) in the original description (Jiang et al. 2015b), but as Indocalamus sp.in Qiao et al. (2018). In China (Fujian, Guizhou). Alatae and life cycle are unknown. Possibly this is a synonym of C. perglandulosa.

Ceratovacuna atrovirensa Qiao Apterae are dark green, covered dorsally with white wax powder, with flaky wax on marginal areas; BL 1.3-1.5 mm. In small populations on undersides of leaves of Indocalamus tesselatus in China (Tibetan plateau); Jiang et al. 2021b). Alatae and life cycle are unknown.

Ceratovacuna beijingensis Qiao Apterae are brown, covered in white wax powder; BL c. 2.4 mm. On leaf sheaves of an unidentified species of Poaceae (Jiang et al. 2015b), perhaps Imperata cylindrica (Qiao et al. 2018). In China (Beijing). Alatae and life cycle are unknown. Possibly this is a synonym of C. cynodonti.

Ceratovacuna cerbera Aoki, Kurosu, Shin & Choe Apterae covered in heavy filamentous wax; BL 2.1-2.5 mm. In colonies on undersides of leaves of Sasa borealis in Korea, and on Sasa sp. in Japan. This is Ceratovacuna “sp. C” of Takahashi (1958c) and Paik (1972). Large first-instar pseudoscorpion-like soldiers are produced along with the normal first instars (Aoki et al. 1999). Alatae produced in November founded new colonies on Sasa, suggesting that this species is without host alternation and anholocyclic in Japan.

Ceratovacuna cynodonti Chakrabarti & Debnath Apterae are blackish, covered with wax; BL 1.4-1.6 mm.In compact colonies on the lower surfaces of young leaves of a Cynodon sp. in northwest Himalaya, India (Chakrabarti & Debnath 2011). Other morphs and life cycle are unknown.

Ceratovacuna doipuiensis Sirikajornjaru, Rojanavongse & Upatham Apterae covered in filamentous wax; BL 1.8-2.2 mm. In colonies on undersides of leaves of Thyrsostachys siamensis in northern Thailand (Sirikajornjaru et al. 2002). Alatae and life cycle unknown.

Ceratovacuna floccifera Noordam Appearance in life unknown, but apterae presumably with flocculent wax; BL 1.4-1.6 mm. First instar soldier larvae occur with enlarged forelegs and very long frontal horns. On undetermined bamboos in Java (Noordam 1991). Life cycle is unknown. Closely related to and possibly synonymous with C. japonica and/or C. longifila.

Ceratovacuna graminum (van der Goot) Apterae are black or greyish black, with dark appendages, body covered with a thin dense layer of wax; BL 1.8-2.1 mm. Alatae have secondary rhinaria distributed III 25-35, IV 7-9, V BASE 9-13. On undersides of leaves of a grass (presumed to be a Panicum sp.) in Java, on an unidentified grass in Vietnam (Szelegiewicz 1968, as ?orientalis), and now also recorded from Imperata cylindrica in China (Fujian; Jiang et al. 2015b). A member of the difficult C. nekoashi group, and possibly an earlier name for C. oplismeni.

Ceratovacuna hoffmanni (Takahashi) Apterae are of unknown appearance in life but probably darkish and with much wax secretion; BL 1.8-2.7 mm. On bamboos (Bambusa, Pleioblastus, Schizostachyum) in China and Sumatra. C. longifila and C. japonica (see below) are also very similar aphids, differing mainly in degrees of wax gland development, and further work is needed to show whether any or all of them are distinct species.

Ceratovacuna imperata Qiao Apterae are red-brown with white wax powder; BL 1.5-1.7 mm. On undersides of leaves of Imperata cylindrica in China (Tibet, Yunnan; Jiang et al. 2015b). Similar to and possibly synonymous with C. perglandulosa.

Ceratovacuna indica M.R. Ghosh, Pal & Raychaudhuri Apterae are pear-shaped, yellowish in life and covered with white wax; BL 1.4-1.7 mm. In colonies on the undersides of older leaves of unidentified bamboos in West Bengal (M.R. Ghosh et al. 1977). Possibly this species belongs in Astegopteryx (Chakrabarti & Debnath 2011).

Ceratovacuna japonica (Takahashi) Apterae on bamboos are pinkish brown to dark brown in life, secreting white wax tufts or filaments; BL 1.5-2.0 mm. On undersides of leaves of small bamboos (Arundinaria, Bambusa, Sasa) in Taiwan, Japan and Korea. Anholocyclic populations live throughout the year on bamboos, but at least in Japan there is also host alternation to Styrax japonica, where banana bundle-shaped galls are produced, similar to those of C. nekoashi but with a longitudinal suture (Kurosu et al. 1990, as C. pseudostyracophila; Aoki & Kurosu 1991). Kurosu & Aoki (1994) studied the process of gall formation, and the roles of first instar “outsiders” left outside the gall and of sterile second instar soldiers. Two other species described from bamboos in Taiwan, brevicornis Takahashi and longifila Takahashi (the latter secreting very long wax filaments), may be synonyms of japonica (but see Liao 1976). 2n=12.

Ceratovacuna keduensis Noordam Apterae are dull black, with paired spinal and marginal rows of short, conical wax tufts; BL 1.6-1.8 mm. On undersides of leaves of undetermined bamboos, often forming large colonies, in Java (Noordam 1991), and now also recorded from an unidentified bamboo in China (Jiang et al. 2015b). Life cycle is unknown. Possibly not distinct from C. hoffmanni.

Ceratovacuna lanigera Zehntner Plate 3c Apterae are pale green or brownish yellow or greyish brown, densely covered with white wax that forms thick columns at margins of body (see aphids of Karnataka website), and may become filamentous (Noordam 1991; Rueda & Calilung 1975); BL 1.4-2.3 mm. Alatae are brown‑black and have secondary rhinaria distributed III 16-25, IV 5-10, V base 2-9. Immature alatae cluster together and produce abundant long filamentous wax. On lower sides of leaves of Poaceae (Imperata, Miscanthus, Oplismenus, Pseudechinolaena, Saccharum, Themeda), in dense colonies, often attended by ants. In India, Nepal, Bangladesh; east and south-east Asia, Fiji and Solomon Islands. It is apparently anholocyclic everywhere; Kurosu & Aoki (1986) found many alate sexuparae in a colony on sugar cane in Japan, and obtained some first instar sexual morphs, but these were possibly non‑functional. Alate sexuparae have also been recorded from October to January and from April to June in Okinawa (Aoki & Kurosu 2010). C. lanigera is a serious pest of sugar cane in south-east Asia, and more recently in western and southern India (Joshi & Viraktamath 2004). However in Japan and Fiji it has only been found on Miscanthus, suggesting geographical variation in its host plant preferences; possibly there is also some confusion with the very similar C. panici. See also Blackman & Eastop (2000). 2n=12.

Ceratovacuna multiglandula Qiao Apterae are dark brown, covered with white wax powder; BL 1.9-2.2 mm. On undersides of leaves of Phyllostachys angusta in China (Zhejiang). Originally described also from Pseudoasasa japonica (Jiang et al. 2015b), but this is not confirmed by Qiao et al. (2018). Alatae and life cycle are unknown.

Ceratovacuna nekoashi (Sasaki) The gall on Styrax spp. in spring in Japan, Korea and Taiwan resembles a bunch of bananas (or a cat’s paw, which is “nekoashi” in Japanese), and is developed from an axillary bud (Kurosu & Aoki 1990). Alate emigrants (BL c.1.7-1.8 mm, with dark cross-bands on abdominal tergites 6-8) emerge in June-July and migrate to Poaceae. C. nekoashi is recorded from several grass genera (Apluda, Arthraxon, Microstegium, Miscanthus, Oplismenus, Panicum), but Aoki et al. (2013) suggest that at least some of these records are based on misidentifications, as despite extensive searching in Japan and Taiwan they only found C. nekoashi and other members of the nekoashi group (oplismeni, orientalis, subtropicana) on grasses of the genus Microstegium. Apterae on undersides of leaves of Microstegium are dark reddish brown, densely covered in white wax wool, and with a marginal fringe of wax tufts; BL 1.5-1.9 mm. The sexuparae returning to Styrax in October differ from the spring emigrants by having frontal horns and less sclerotisation of abdominal tergite 6 than alatae from galls (Takahashi 1958). They deposit first instar sexuales on the leaves; these move to the branches without moulting and develop to maturity without feeding, the eggs being laid on bark (Kurosu & Aoki 1990). Secondary host populations are recorded from Japan, Taiwan, Korea and Uttar Pradesh, India (Chakrabarti & Maity 1982). 2n=12.

Ceratovacuna oplismeni (Takahashi) “Cat’s paw” galls of this species occur on Styrax japonicus in Okinawa (Japan), and on S. formosanus in Taiwan. Aoki et al. (2013) distinguished C. oplismeni from C. nekoashi and its other close relatives (orientalis, subtropicana) by analysis of mitochondrial rRNA sequences. Immature galls typically have radiating sub-galls resembling a sunflower. Mature and open galls were found in June-July. Secondary host generations occur on Microstegium vimineum in Okinawa and on Microstegium sp. in Taiwan; apterae on Microstegium are generally blackish purple (cf. nekoashi), sometimes tinged with brown, coated with wax powder and with a fringe of long white wax tufts. Colonies probably persist on the secondary host during winter. Sexuparae are undescribed.

Ceratovacuna orientalis (Takahashi) “Cat’s paw” galls of this species occur on Styrax formosanus in Taiwan. Aoki et al. (2013) distinguished C. orientalis from C. nekoashi and its other close relatives (oplismeni, subtropicana) by analysis of mitochondrial rRNA sequences. Mature or nearly mature galls were found in July and August, and colonies on grasses (Microstegium sp.) were found in early August; apterae on Microstegium are reddish brown with a fringe of long white wax tufts. Sexuparae are undescribed.

Ceratovacuna panici (van der Goot) Apterae are yellowish or brownish, with white wax forming tufts or columns marginally, shorter than is sometimes the case in C. lanigera (Noordam 1991); BL 1.5-1.9 mm. Alatae have secondary rhinaria distributed III 17-30, IV 7-15, V base 6-14. On undersides of leaves of tropical grasses (Cynodon, Microstegium, Panicum, Paspalum, Setaria) in Java, and there is also now a record from Imperata sp. in China (Fujian; Huang et al. 2012). The available morphs were fully redescribed by Noordam (1991), but the life cycle is unknown. C. panici is closely related to C. lanigera, and possibly some records of C. lanigera from grasses other than Saccharum outside Java are referable to this species.

Ceratovacuna perglandulosa R.C. Basu, A.K.Ghosh & Raychaudhuri Apterae are reddish brown or black, covered in white wax (see aphids of Karnataka website); BL 1.8-2.2 mm. In India, on Saccharum officinarum and “Grassum prothama”, an untraceable plant name (R.C. Basu et al. 1975). Chakrabarti & Debnath (2011) record it from Apluda mutica.

Ceratovacuna silvestrii (Takahashi) Apterae are yellow to reddish brown or almost black in life, with numerous white wax tufts over dorsum and sides of body; BL 1.6-2.3 mm. On undersides of leaves, near their bases, of both Arundinaria spp. and Bambusa spp. There are now also records from China on Phyllostachys heterocycla (= edulis) and Indosasa shibataeaoides (Jiang et al. 2015b). Recorded from China, Taiwan, and India (as subglandulosa Hille Ris Lambers & Basu 1966). Agarwala et al. (1987) described a parasitoid. 2n=12 (Khuda-Bukhsh & Kar 1987).

Ceratovacuna spinulosa (A.K. Ghosh & Raychaudhuri) Apterae are “deep brown”, presumably secreting wax; BL c.1.2-1.3 mm. On Ischaemum albens in West Bengal, India. Possibly this is a member of the Ceratovacuna nekoashi group (see Aoki et al. 2013)

Ceratovacuna subtropicana Aoki & Kurosu “Cat’s paw” galls of this species occur on Styrax formosanus in Taiwan. Aoki et al. (2013) distinguished C. subtropicana as a new species distinct from C. nekoashi and its other close relatives (oplismeni, orientalis) by analysis of mitochondrial rRNA sequences, and described and illustrated the galls, apterous fundatrigeniae, soldiers, emigrant alatae, as well as first instar and adult apterae and sexuparae collected on the secondary host (Microstegium sp.). Mature, open galls were found in late May-June. Apterae on Microstegium grasses are yellowish orange (cf. nekoashi), wax-covered, with long white wax marginal tufts. Sexuparae were found on Microstegium in November, and differ from those of C. nekoashi in having vestigeal frontal horns. Colonies survive the winter and persist throughout the year in Taiwan on Microstegium (Aoki et al. 2013).

Ceriferella Carver & Martyn

Lizeriinae

Two Australian species producing thick strands of wax from numerous dorsal wax glands. Apterae have eyes reduced to three facets, and 5-segmented antennae. The host plant of one species described from Western Australia (dossuaria; Carver & Martyn 1965) is still unknown. Quednau (2010) reviewed and keyed the species and illustrated all available morphs.

Ceriferella leucopogonis Carver and Martyn Apterae are dark reddish brown with long thick strands of white wax dorsally and laterally; BL 0.8-1.1 mm. Living singly or in small colonies on foliage of Epacridaceae (Leucopogon spp. and Lissanthe strigosa) in south-east Australia.

Ceruraphis Börner

Aphidinae: Macrosiphini

Three nominal species, one nearctic and two palaearctic, that host-alternate between Viburnum and Cyperaceae. Although the three species share some features and are clearly related in biology, each has some unique morphological peculiarities which could justify separate generic or subgeneric status. The genus was reviewed by Chakrabarti & Medda (1985), and MacGillivray (1960) compared the two species in North America.

Ceruraphis eastopi Hille Ris Lambers (Fig.58a,c) Colour of apterae (fundatrices?) is unrecorded, but presumably pale grey, green or brown with blackish head and siphunculi; BL 3.4-3.8 mm. On Viburnum cotinifolium, in pseudogalls formed by folding leaf margins ventrally (Bhattacharya et al. 1980). Heteroecious holocyclic; secondary host generations are undescribed, although a sample collected in Iran on Carex sp. is ascribed to this species (Naumann-Etienne & Remaudière 1995). In Iran(?), Pakistan and northern India. [The original description of C. eastopi from Pakistan is of alatae and immatures only, collected on Viburnum cotinifolium in Pakistan in September. The apterae described by Bhattacharya et al. (1980) from Viburnum sp. in Garhwal Himalaya (Uttar Pradesh) in June, are keyed by us as eastopi, but are possibly the fundatrices of a different species.]

Ceruraphis eriophori (Walker) Plate 11e Apterae (including fundatrices) blackish (see influentialpoints.com/Gallery); BL (fundatrices on Viburnum) 2.5-3.0 mm, BL (apterae on Cyperaceae) 2.0-2.8 mm. Curling leaves of Viburnum spp. in spring, migrating from second generation to Cyperaceae (Carex, Cyperus, Eriophorum, Luzula, Typha). Apterae on secondary hosts have a tuft of wax at end of abdomen. In Europe, Iran (Hodjat 1993), northern India (BMNH collection, leg. S. Chakrabarti) and introduced to North America. Ant-attended. 2n=14.

Ceruraphis viburnicola (Gillette) (Fig.58b,d) Apterae (fundatrices) dark green, wax-dusted, with black antennae, legs, siphunculi and cauda; BL 2.5-3.0 mm. In curled leaves of Viburnum spp. in spring. Heteroecious holocyclic, migrating to Cyperaceae on basis of greenhouse transfers (MacGillivray 1960), although not yet collected in the field on secondary hosts. Apterae reared on Scirpus were brown dorsally and pink ventrally with pink head and no white wax tuft at the end of the abdomen, and had BL 1.9-2.4 mm (cf. eriophori; MacGillivray 1960). Sexuales occur on Viburnum in September-November (Palmer 1952). In North America; a record from Turkey (Kaygin et al. 2010) requires further confirmation. 2n=14.

Cervaphis van der Goot

Greenideinae: Cervaphidini

A genus with four or five species in India and south-east Asia, in which the apterae have remarkable branched marginal hair-bearing processes (fig. 92A). In alatae these processes are reduced to low, flat hair-bearing tubercles, except on the head. The siphunculi are long, cylindrical, slightly curved outwards and a little swollen subapically, where there is a ring of small hairs. For revisions of Cervaphis see Hille Ris Lambers (1956b), A.K. Ghosh (1982a), A.K. Ghosh & Agarwala (1993), Noordam (1994), Qiao & Zhang (2000b) and D. Zhang & Qiao (2008b).

Cervaphis echinata Hille Ris Lambers Apterae are probably greenish yellow, elongate oval, BL 1.2-1.6 mm. On undersides of young leaves of Schoutenia ovata (= Actinophora fragrans) in Java. It is also reported from Paulownia sp. in China (Hainan; Q. Liu et al. 2013). The aptera was redescribed by Noordam (1994). Other morphs are undescribed.

Cervaphis quercus Takahashi Apterae are yellow with pale siphunculi, BL about 1.7 mm, feeding on undersides of leaves and new growth of Quercus spp. and Castanea spp. in east Asia (recorded from Japan, Taiwan, Thailand, Laos, China, Korea, and Assam, India). Sometimes ant-attended. Alatae have brownish head, thorax and dorsal abdominal markings including a large central patch, and siphunculi almost black (Takahashi 1924). Biology is largely unstudied; alate oviparae have been found in Japan from July onwards, produced by apterous sexuparae, but males have not been recorded (Takahashi 1923). However, the complete mitochondrial genome has been published (Wang et al. 2014). 2n=8.

Cervaphis rappardi Hille Ris Lambers Apterae are yellow or greenish yellow (see aphids of Karnataka website) , BL 1.2-1.7 mm. Alatae have olive green abdomen with head and thorax slightly darker. Noordam (1994) provided a detailed redescription. On flowers, flowerstalks, and sometimes on leaves, young shoots, or young fruit of Theobroma cacao, attended by ants (Dolichoderus). Feeding may cause flower heads to shrivel and fall off. Colonies also occur on Nephelium lappaceum which may be the native host plant, Aglaia sp. and Erycibe grandiflora (in Java). Sexual morphs are unknown. In Indonesia, Malaysia, New Guinea, Philippines, and India, where a subspecies, C. rappardi indica A.N. Basu, was described from Cajanus cajan in Assam and West Bengal (see A.K. Ghosh & Agarwala 1993 and aphids of Karnataka website for further information). 2n=8 for ssp. indica (Kar et al. 1990).

Cervaphis schouteniae van der Goot Plate 5h Apterae are whitish-green to yellowish-white with pale yellowish-brown siphunculi; BL 1.5-1.9 mm. Alatae have brownish-yellow head and thorax, pale yellow abdomen, dark antennae and black siphunculi. Noordam (1994) provided a detailed redescription. On undersides of young leaves, feeding along main veins near petioles. Host plants include trees and shrubs mainly in Tiliaceae (Grewia, Microcos, Schoutenia), but also recorded from plants in other families, such as Pterospermum (Sterculiaceae), Dipterocarpus (Dipterocarpaceae) and Zizyphus (Rhamnaceae). Recorded from Malaysia, Indonesia, Philippines, Thailand, Cambodia and India. Sexuales are unrecorded. Population trends in north-east India were studied by Agarwala & Dixon (1986), and developmental stages are described by Agarwala et al. (1985). A parasitoid, Trioxys peniculatus, was recorded by Agarwala et al. (1987).

Chaetogeoica Tao

Eriosomatinae: Fordini

A poorly-known genus for five or six species related to Geoica, but with emigrant alatae having ciliated secondary rhinaria and embryos with fewer dorsal hairs. Complete life cycles presumably involve alternation between Pistacia and grass roots but have not been confirmed for any species. The root-feeding forms are known for only two species in India (Raychaudhuri, Pal & M.R. Ghosh 1978). Accounts are available for India (A.K. Ghosh 1984) and China (W. Zhang et al. 1995a; but see Remaudière & Remaudière 1997, p. 296 and 299).

Chaetogeoica foliodentata (Tao) The gall on Pistacia sinensis is a green and pinkish elongate sac about 4 cm long with longitudinal ridges and a terminal opening, arising from a leaflet (fig. 126D). Emigrant alatae, BL 1.4-1.9 mm (fig. 81E), are mature in May (Remaudière & Tao 1957). The secondary hosts are unknown, but presumably roots of Poaceae. Recorded from Pistacia in China and Taiwan, and single alatae of a Chaetogeoica that is possibly this species have also been trapped or collected in Korea (Paik 1972), the Philippines, Nepal (BMNH collection) and Sikkim, India (A.K. Ghosh & Raychaudhuri 1968a).

Chaetogeoica graminiphaga Raychaudhuri, Pal & M.R. Ghosh Colour of apterae in life is unrecorded; BL c.1.9 mm. On roots of various grass species (Capillipedium, Eleusine, Ischaemum, Polypogon) in West Bengal and Manipur, India. Life cycle is unknown.

Chaetogeoica ovagalla (Zhang) The gall on Pistacia sinensisis is an elongate cucumber-shaped sac 4-5 cm long arising from the midrib of a leaflet (G. Zhang 1995, as Baizongia ovagalla). Emigrant alatae, BL c. 1.6 mm, were collected in July in China (Shandong). Remaudière & Remaudière (1997, p. 299) drew attention to the different form of the dorsal hairs of embryos of this species compared with those of C. foliodentata, and suggested that C. sensucopia (Zhang), described in the same paper (as Baizongia sensucopia), may be a synonym. It seems likely that C. ulmidrupa Zhang, described with exactly the same collection data but with the host identified as Pistacia aculeata (G. Zhang & Qiao 1998a), may also be a synonym of C. ovagalla.

Chaetogeoica polychaeta Raychaudhuri, Pal & M.R. Ghosh Apterae are creamy white to pink or pale brown, with conspicuous pubescence; BL 1.5-2.3 mm. On roots of Capillipedium sp., Oryza sativa and other Poaceae in West Bengal and Manipur, India. Alatae and life cycle are unknown.

Chaetogeoica yunlongensis (Zhang & Zhong) The gall on Pistacia sinensis is an elongate cucumber-shaped sac 5-6 cm long with longitudinal ridges very like that of C. foliodentata (W. Zhang et al. 1995a). Emigrant alatae, BL c. 1.9 mm, were collected in mid-June in China (Yunnan; G. Zhang & Zhong 1985b).

Chaetomyzus Ghosh & Raychaudhuri

Aphidinae: Macrosiphini

Two Asian species on Rhododendron, perhaps related to Indomasonaphis but smaller aphids with scabrous dorsal processes and few caudal hairs.

Chaetomyzus rhododendri A.K. Ghosh & Raychaudhuri Colour of apterae in life is unrecorded; BL 1.8-2.4 mm. On undersides of young leaves of Rhododendron arboreum and Rhododendron sp. in northern India (Assam, Himachal Pradesh, Uttar Pradesh)and Nepal. Chakrabarti, A.K. Ghosh & Chowdhuri (1971) provided a redescription. A small sample of one alata and two immatures from Sonchus in Nepal (BMNH collection) suggests that the secondary host is a composite, as in the related Indomasonaphis.

Chaetomyzus rhododendrophila (Zhang, Zhang & Zhong) Apterae are undescribed; BL of alata c 2.8 mm. On Rhododendron simsii in Yunnan Province, China.

Chaetosiphella Hille Ris Lambers

Chaitophorinae: Siphini

Five or six species of dark, elongate-bodied aphids on grasses closely related to Atheroides but, unusually for a grass-feeder, they have a stiletto-shaped R IV+V. Kadyrbekov (2005b) provided a key to species, and Wieczorek (2008b, 2010) fully revised the genus.

Chaetosiphella berlesei (del Guercio) Apterae are dull dark grey to black, slightly wax-dusted; BL 1.7-2.0 mm. On upper sides of leaves of grasses (Aira, Corynephorus, Deschampsia, Festuca), mainly in dry sandy habitats, dropping off plant and running when disturbed. In Europe (not Iberian Peninsula) and west Siberia. Monoecious holocyclic with apterous males (Heie 1982).

Chaetosiphella longirostris Wieczorek Colour of apterae in life was not observed; BL 2.4-2.6 mm. Described from Ammophila arenaria at a coastal site in Portugal (Wieczorek 2008b), and there is also now a record from Stipa arabica in Iran (Mosapour et al. 2019). Biology is unknown.

Chaetosiphella massagetica Kadyrbekov Apterae are greyish green; BL 1.4-1.9 mm. On upper sides of leaves of Stipa spp. in Kazakhstan (Kadyrbekov 2005b, 2017a). A redescription of a paratype aptera was provided by Wieczorek (2010). Life cycle is unknown.

Chaetosiphella stipae Hille Ris Lambers (= Chaetosiphella pamirica Narzikulov) Apterae are dark grey or blackish brown to dull black; BL 1.5-2.1 mm. On Stipa spp. in Europe, and across Asia to Mongolia and China. In Central Asia it was also found on Acantholimon pamiricum (Plumbaginaceae; Narzikulov 1970, as Ch. pamirica). Geographical variation in abdominal chaetotaxy and other morphological features was studied by Wieczorek et al. (2017a). A form with a much shorter (but still stiletto-shaped) R IV+V, collected on Calamagrostis argentea (= Stipa calamagrostis) in southern France, and also found in former Yugoslavia (BMNH collection, leg. D. Hille Ris Lambers), was described as a subspecies, Ch. stipae ssp. setosa by Wieczorek (2008b), but should surely have full species status. Both taxa are monoecious holocyclic with apterous males (Nieto Nafría & Mier Durante 1998, Wieczorek 2008b). The male genitalia were described and illustrated by Wieczorek et al. (2011). Niu et al. (2016) compared its mitochondrial genome with that of Periphyllus acerihabitans.

Chaetosiphella tshernavini (Mordvilko) Apterae are blackish grey; BL 1.3-1.8 mm. On leaves of Corynephorus canescens, Festuca spp. and Stipa capillata in eastern Europe and west Siberia (Wieczorek 2010). Sexual morphs are unknown.

Chaetosiphon Mordvilko

Aphidinae: Macrosiphini

About 20 species widely distributed in the northern hemisphere and mostly living without host alternation on Rosaceae of the Rosa‑Fragaria‑Potentilla group. They are characterized by having 5 hairs on all first tarsal segments, and apterae with conspicuous capitate hairs, cylindrical siphunculi and a short cauda. Alatae have inconspicuous dorsal hairs and dorsal abdominal pigmentation. Fundatrices have shorter hairs than later generations of apterae, and parasitised apterae may also have short dorsal hairs. The economically important species, placed in the subgenus Pentatrichopus, were formerly included in Capitophorus. They are not attended by ants. Accounts are available for Europe (Hille Ris Lambers 1953, Heie 1994, Blackman 2010), Japan (Miyazaki 1971), China (Yang et al. 2005b), India (David et al. 1971a), and North America (Schaefers 1960, Richards 1963b). Remaudière (1991) provided a key to species.

Chaetosiphon alpestre Hille Ris Lambers Apterae are shiny pale green with a slightly brownish tinge; BL 1.5-2.1 mm. Alatae have a dark central dorsal abdominal patch. On flower stems, petioles and young stems of Potentilla spp. in Europe (France, Switzerland, Italy, Sweden, Poland, Czech Republic) and Kazakhstan (Kadyrbekov 2004b). Aphids collected on Artemisia sp. in Mongolia were also identified as this species (Szelegiewicz 1963). Two subspecies have been described, Ch. alpestre ssp. airolense Hille Ris Lambers found in Switzerland on P. puberula and also in Sweden on P. megalantha (Ossiannilsson 1959), and Ch. alpestre ssp. orientale Shaposhnikov, found in western Kazakhstan on undersides of leaves of P. supina (but subsequently reported from other Potentilla spp.; Kadyrbekov 2017a). Monoecious holocyclic, with alate males (ssp. airolense).

Chaetosiphon (Pentatrichopus) alpinum (Börner) Apterae are probably very pale whitish or yellowish; BL 1.4-1.8 mm. In leaf axils of Potentilla spp. in Austria.

Chaetosiphon chaetosiphon (Nevsky) Apterae are pale orange (original description), or pale green to mid-green, shiny (Hille Ris Lambers 1953); BL 2.0-2.6 mm. Alatae have a large black dorsal abdominal patch. On wild Rosa spp. in mountainous regions of southern Europe, south-west and central Asia and north-west Himalayas. Hille Ris Lambers (1953) suggested that the Swiss populations should be regarded as a subspecies, Ch. chaetosiphon ssp. montanum. The life cycle is unknown; fundatrices were found in May in Switzerland.

Chaetosiphon (Pentatrichopus) coreanum (Paik) Apterae are pale yellow to almost white; BL 1.7-1.9 mm (spring), 0.9-1.3 mm (summer). Alatae have a dark dorsal abdominal patch. On young growth in spring, and later on undersides of leaves, of Rosa spp. In Japan and Korea. Monoecious holocyclic, with oviparae in October (Miyazaki 1968a). 2n=8.

Chaetosiphon (Pentatrichopus) fragaefolii (Cockerell) Strawberry Aphid Plate 16c Apterae are translucent yellowish white to pale greenish yellow (see influential points/gallery); BL 0.9-1.8 mm. Alatae have a brown-black dorsal abdominal patch. On Fragaria spp., especially cultivated varieties; it occurs frequently on F. chiloensis in America, but rarely on F. vesca in Europe. Also sometimes found on Potentilla anserina. An important cosmopolitan vector of strawberry viruses (Blackman & Eastop 2000). Mainly anholocyclic; oviparae and both apterous and alate males are produced in certain laboratory cultures, but are rare in field. There has been some confusion in the North American literature with Ch. thomasi (Blackman et al. 1987). 2n=13, 14 or 15.

Chaetosiphon (Pentatrichopus) glabrum David, Rajasingh & Narayanan Apterae are pinkish brown or green, with dark dorsal abdominal cross-bands and brown appendages; BL 1.3-1.9 mm. Alatae also have dark abdominal cross-bands. On Rosa spp. in Himachal Pradesh, India. Very similar apterae, but with most dorsal body hairs very small, have been collected on Rosa sp. in Kashmir (BMNH collection, leg. N.D. Rishi), and could be a hot-weather form of glabrum, or possibly parasitised individuals.

Chaetosiphon (Pentatrichopus) gracilicorne David, Rajasingh & Narayanan Apterae are pale greenish yellow to dark green with pale appendages; BL 1.4-1.65 mm. On Rosa spp. in India (Himachal Pradesh). Chakrabarti (1976b) described alate viviparae and oviparae, both collected (as well as apterous viviparae) in December. 2n=16 (Dutta & Gautam 1993)

Chaetosiphon (Chaitomyzus) heterotrichum Chakrabarti, A.K. Ghosh & Raychaudhuri (Fig.48b) Apterae are dull white; BL 1.8-2.1 mm. Alatae lack dark dorsal abdominal markings. On undersides of leaves of Salvia glutinosa and Salvia sp. in northern India (Uttar Pradesh).

Chaetosiphon (Chaitomyzus) hirticorne (Takahashi) Apterae are yellow, with dusky to blackish ANT III-VI, but pale legs and siphunculi; BL c. 1.5 mm. The alata was briefly described by Moritsu (1983). On Clinopodium gracile in Japan, and from undersides of leaves of an unidentified plant of the Lamiaceae in China (Yang et al. 2005b; originally recorded as Micromyzodium clinopodii ssp. minensis by Zhang et al. 1999). In Japan it is also recorded from Salvia japonica, although this record could possibly be C. heterotrichum.

Chaetosiphon (Pentatrichopus) hottesi Stroyan Apterae are pale whitish yellow to yellowish green; BL 1.9-2.1 mm. Alatae have dorsal abdomen with dusky sclerotic bars, not fused into a solid patch. On Potentilla spp. and Rosa spp. in western North America.

Chaetosiphon (Pentatrichopus) jacobi Hille Ris Lambers Apterae are similar to fragaefolii but darker, with tergum brownish; BL 1.2-1.8 mm. On thin-leafed wild Fragaria spp. in western USA, colonising cultivated strawberries only in glasshouse or laboratory conditions. Apparently entirely anholocyclic (Blackman et al. 1987). 2n=17.

Chaetosiphon janetscheki (Börner) Colour of apterae in life unknown; BL 1.6-1.9 mm. On Geum sp. (either montanum or reptans) in Austria, and also on Geum plants imported to UK from Switzerland (Hille Ris Lambers 1953). Life cycle is unknown.

Chaetosiphon (Pentatrichopus) minor (Forbes) Apterae are yellow‑green; BL 1.0‑1.4 mm. On stems and undersides of leaves of Fragaria spp. in eastern North America, Honduras (Evans & Halbert 2007), Venezuela, Japan (Miyazaki 1971), Korea and the Philippines. Monoecious holocyclic, with alate males. See also Blackman & Eastop (2000). 2n=12.

Chaetosiphon (Pentatrichopus) muelleri Remaudière Apterae are whitish cream or yellowish; BL 1.0-1.4 mm. Dorsal abdomen of alata with a dark central patch, sometimes subdivided. On undersides of leaves of Potentilla speciosa in Turkey.

Chaetosiphon (Pentatrichopus) potentillae (Walker) Apterae are yellowish or whitish; BL 1.4-2.1 mm. On Potentilla spp., living singly or in small numbers on young leaves. Europe and North America. Monoecios holocyclic, with oviparae and apterous males in September-October (Heie 1994).

Chaetosiphon (Pentatrichopus) tetrarhodum (Walker) Apterae are pale green to yellow-green, occasionally reddish (see influential points/gallery); BL 1.0-2.6 mm. Alatae have a black central abdominal patch. On both wild and cultivated Rosa spp., colonising shoot tips in spring, and later much smaller individuals live dispersed on undersides of mature leaves. World-wide, except the Asian Far East. Monoecious holocyclic, with dark olive-green oviparae and small dark apterous males in October (Heie 1994). 2n=14 (two UK samples) or 16* (one sample from A.C.T., Australia).

Chaetosiphon (Pentatrichopus) thomasi Hille Ris Lambers Apterae are yellowish green (see influential points/gallery); BL 1.0-2.6 mm. On both wild and cultivated Rosa spp. in North America, and reported also to occur in South America (Chile, Argentina). Certain Potentilla spp., e.g. P. monspeliensis, may also be colonised (but it has not so far been recorded from P. anserina). Monoecious holocyclic on Rosa rugosa in British Columbia, with both apterous and alate males. Populations on Rosa multiflora in New Zealand appear to be morphologically distinct from both thomasi and fragaefolii, and a form with distinctive alatae on R. fendleri in western USA (Utah, Colorado) may also be an undescribed taxon (Blackman et al. 1987). 2n=12.

Chaitaphis Nevsky

Aphidinae: Macrosiphini

Five (or more) species on Amaranthaceae in eastern Europe and Asia, reviewed and keyed by Kadyrbekov (2002c).

Chaitaphis camphorosmae (Hille Ris Lambers) Apterae are pale green, with apices of appendages brownish; BL 1.1-1.6 mm. On Camphorosma spp. Austria and western Kazakhstan. Monoecious holocyclic; oviparae and alate males in September (original description, as Hayhurstia). Redescribed by Kadyrbekov (2002c).

Chaitaphis kazakhstanica Kadyrbekov Apterae are pale green with greyish tinge, with front of head and apices of appendages darker; BL 1.1-1.5 mm. In clumped and reddened apical leaves of Bassia sedoides (Kadyrbekov 2002c). Only known from Kazakhstan.

Chaitaphis safavii Remaudière (Fig.36b) Apterae are very pale green, with apices of appendages darker; BL 1.4-1.6 mm. On Kochia cana in Iran, and also recorded from that country on Suaeda fruticosa (Hodjat 1993). Monoecious holocyclic, with oviparae and alate males in October (Remaudière 1989b).

Chaitaphis shaposhnikovi Kadyrbekov Colour of apterae in life unknown, probably green; BL 1.4-1.7 mm. On Salsola sp. in west Kazakshtan (Kadyrbekov 2002c). Closely related to Ch. tenuicauda.

Chaitaphis tenuicauda Nevsky (Fig.36a,c) Apterae are yellow-green, with dark apices to antennae, legs and siphunculi; BL 1.1-1.6 mm. In dense colonies on stems of Kochia and Bassia spp., and also recorded from certain other Amaranthaceae (Salsola, Suaeda), although according to Kadyrbekov (2002c) this species does not occur on Salsola lanata. In Iran, Uzbekistan, Kazakhstan, Pakistan and India (New Delhi; BMNH collection, leg. Menon). Monoecious holocyclic, with oviparae and alate males in November-December (Remaudière 1989b). [Kadyrbekov (2002c) redescribed Ch. tenuicauda from Kochia (= Bassia) prostrata in Kazakhstan, but his specimens have a larger ratio of R IV+V to HT II than specimens from Bassia and Kochia spp. (prostrata, childsii) in India (BMNH collection) and from Amaranthaceae in Pakistan (Remaudière 1989b).]

Chaitogenophorus Zhang, Qiao & Chen

Chaitophorinae: Chaitophorini

A genus for one species in north-west China.

Chaitogenophorus yuzhongensis Zhang, Qiao & Chen Apterae elongate oval; BL c.1.3 mm. On Populus davidiana in north-west China (G. Zhang 1999).

Chaitophorus Koch

Chaitophorinae: Chaitophorini

A large holarctic genus (about 90 species) associated almost exclusively with Salicaceae, with individual species feeding on either Populus or Salix but never both, and often with a high degree of host specificity. They are very hairy, small to medium-sized aphids with short, stump-shaped siphunculi having at least some trace of reticulate ornamentation. The dorsal cuticle of the aptera is usually more-or-less completely sclerotic and variably sculptured over at least abdominal tergites 2-6. Alatae often have dark segmental cross-bands an marginal sclerites. The cauda is usually knobbed, but (unlike most Drepanosiphinae with knobbed caudas) the anal plate is entire. Colour photographs of some common European species can be found at: http://influentialpoints.com/Gallery/Chaitophorus_aphids.htm#popule. Pintera (1987) fully revised the Palaearctic fauna (58 species), Wieczorek & Osiadacz (2007) reviewed species in Poland, Nieto Nafrìa & Mier Durante (1998) provided a detailed account of the Iberian species, and there are accounts for India by Chakrabarti (1977) and A.K. Ghosh (1980), for China by Tao (1964), for Japan by Higuchi (1972), and for North America there are revisions by Hille Ris Lambers (1960a) and Richards (1972a – Canada only). Shingleton & Stern (2003) constructed a molecular phylogeny of 15 Chaitophorus species based on mtDNA sequences, which indicated that ant attendance has been gained or lost several times in the evolution of the genus. X. Yang et al. (2016) studied the molecular coevolution of Chaitophorus and its primary symbiont Buchnera. T. Liu et al. ( 2022) studied macroevolution of Chaitophorus and other Chaitophorinae in relation to changes in host use. Molecular studies have indicated that Chaitophorus may have a closer relationship to the monocot-feeding Siphini than to the Acer-feeding genus Periphyllus (Wieczorek & Kajtoch 2011); but the results of T. Liu et al. (2022) suggest an earlier transition to grasses in Siphini. The parasitoids Lysiphlebus salicaphis (refs: Smith 1944, Stary 1976) and Aphelinus aureus (Gahan 1924) specialise on Chaitophorus. Aphelinus fulvus also parasitises Pterocomma spp. on the same hosts (Kalina & Starý 1976).

Chaitophorus capreae (Mosley) Apterae are white to yellowish-white (see influentialpoints.com/Gallery); BL 0.8-1.9 mm. Alatae without distinct dark dorsal abdominal markings. Mostly on broad-leafed Salix spp., scattered on undersides of leaves, not attended by ants (see Heie 1982; Pintera 1987). Apterous males (yellow) and oviparae occur in October-November. Widespread in Europe, and eastward to Central Asia. 2n=30. C. flavissimus Mamontova, described from Salix caprea in Ukraine (Mamontova 1979), is very similar to capreae but has a less constricted cauda and possibly longer antennal hairs.

Chaitophorus clarus Tseng & Tao Appearance of apterae in life is unknown, probably pale-coloured; BL about 1.3-1.4 mm. Alatae are without any distinct dorsal abdominal markings. On Populus simonii in China (see Pintera 1987). Indian records under this name from Acer spp. are probably misidentified Trichaitophorus aceris. Biology is unknown.

Chaitophorus crinitus Ivanoskaya Apterae greenish-brown; BL 1.8-2.0 mm. Described from leaves of an unidentified Salix sp., attended by ants, in Siberia (see Pintera 1987), and since recorded from Salix alba (Novgorodova & Stekolshchikov 2013), and from S. cinerea (in Kazakhstan; Kadyrbekov 2017a).

Chaitophorus crucis (Essig) Apterae according to the original description (Essig 1912a) have distinctive pigmentation in life; rich green with broad light yellow-green spinal stripe from the base of the head to the cauda, and a transverse band of similar width and colour across the anterior abdominal tergites, making a large cross. BL about 1.3 mm. Alatae have dark dorsal abdominal markings. Found in dense colonies on undersides of leaves of Salix macrostachyae in California, but not recorded since the original description.

Chaitophorus diversisetosus Szelegiewicz & Czylok Apterae according to the original description (Szelegiewicz & Czylok 1981) are blackish green with a paler central patch, and have dark legs and tips to the siphunculi; BL about 1.8 mm. Alatae have dark dorsal abdominal cross-bands. In ant-attended colonies on undersides of leaves of Salix acutifolia in Poland. A related form, C. diversisetosus ssp. austriacus was described from Salix purpurea in Austria, France and Switzerland (Pintera 1987), and this subspecies is also reported to occur on S. caspica in Kazakhstan (Kadyrbekov 2017a). Apterae of this subspecies are “very pale green with a bright green lyre pattern and short median stripe sometimes darkened,” with legs also very dark; BL 1.3-1.6 mm. Colonies, on upper as well as undersides of leaves, were not attended by ants. An apterous male was collected in August.

Chaitophorus dorocola Matsumura Apterae are elongate-bodied, presumably pale-coloured and, according to original description, covered with a woolly wax secretion (an unusual feature for a Chaitophorus); BL about 1.3 mm. Alatae identified as this species by Pintera (1987) have broad dark dorsal abdominal cross-bands, and BL 1.8-2.1 mm. On undersides of leaves of Populus sieboldii in Hokkaido, Japan, and also reported to occur on P. davidiana in the Siberia (Pashchenko 1988a) and Populus sp. in West Bengal, India (Maity & Chakrabarti 1981a). Specimens from P. alba in China were described as a subspecies, Ch. dorocola ssp. wuweiensis Zhang, Qiao & Chen (G. Zhang 1999). Higuchi’s (1972) account of dorocola, and probably most records from P. maximowiczii, refer to C. inouyei. 2n=14.

Chaitophorus eoessigi Hille Ris Lambers Apterae are variable in colour and markings even within colonies, often with a dark dorsal lyre-shaped pattern,or almost wholly dark except for a pale spinal stripe; BL 1.1-1.9 mm. Alatae have broad dark dorsal abdominal cross-bands. Mostly on the apical parts of leaves, on both upper and undersides, of unidentified Salix sp(p). in southern California and Mexico (Hille Ris Lambers 1966a).

Chaitophorus eugeniae Basu & Raychaudhuri Apterae are pale green; BL c. 1.5 mm. Alatae are without dorsal cross-bands. In a dense colony, including immatures, attended by red and black ants. clustered along the veins on both sides of the leaves of a tree recorded as Eugenia heyneana (= Syzygium salicifolium), which was presumably a misidentified Salix sp. In Orissa, India (M. Basu & Raychaudhuri 1983).

Chaitophorus euphraticus Hodjat Apterae are pale green, BL 1.1-2.1 mm. Alatae are without dorsal cross-bands. On undersides of leaves of Populus euphraticus, attended by ants, in Iran, Jordan and Iraq; occasionally found on P. × euroamericana and P. nigra var. italica. This aphid possibly overwinters as parthenogenetic forms in southern Iran, possibly surviving in December-January in fallen leaves stuck together by lepidopterous larvae (Hodjat 1981). Sexuales (an ovipara and an alate male) were collected on P. × euroamericana in November in northern Iran. 2n=22.

Chaitophorus furcatus Quednau Apterae are bright or dull yellow, BL 1.2-1.6 mm. Alatae are without distinct dorsal abdominal markings. In ant-attended colonies on Salix caprea in Finland, Germany, Czechoslovakia and European Russia. Apterous males and oviparae were collected in Finland in late September (Pintera 1987). 2n=16*.

Chaitophorus gomesi Ilharco Apterae are pale green mottled with dark green, BL 1.1-1.4 mm. Alatae have dorsal abdominal cross-bands and smoky-bordered forewing veins (Nieto Nafrìa & Mier Durante 1998) . In large colonies on undersides of leaves of Salix spp., attended by ants. Only known from Portugal and Spain. Apterous males and oviparae were collected in November (Ilharco 1968).

Chaitophorus hillerislambersi Pintera Colour of apterae in life is unknown, probably pale with dark legs; BL 2.0-2.4 mm. Other morphs are unknown. On Populus sp. in Beijing, China (Pintera 1987), with a record also from Iran (Mortazavi 2015), which posssibly needs confirmation.

Chaitophorus himalayensis (Das) Apterae according to the original description (Das 1918) are light green to yellowish, distinctively marked with a dark green patch on mesothorax, and two dark green crescentic bands leaving paler areas around bases of siphunculi, meeting two elongate spots on abdominal tergite 1 (see also aphids of Karnataka website); BL 1.1-1.9 mm. The siphunculi are short and brown. Alatae have similar dark green dorsal markings. On Salix spp., including S., tetrasperma, feeding along midribs or leaves or on young growth, attended by ants. In northern India, Pakistan, and possibly Thailand and China; Hille Ris Lambers (1966c) pointed out that several records of this species from Thailand and China do not conform to the original description. Some of these authors may have had C. pakistanicus (q.v.). Sexuales are unknown. 2n=18 (Dutta & Gautam 1993).

Chaitophorus hokkaidensis Higuchi Apterae are elongate oval, black in life (Lee & Seo 1992), with legs paler; BL 1.3-1.8 mm. Alatae have broad dorsal abdominal cross-bands. Described from an unidentified Salix species in Hokkaido, Japan (Higuchi 1972), and recorded from undersides of leaves of Salix gracilistyla in Korea (Lee & Seo 1992).

Chaitophorus horii Takahashi Apterae are whitish to pale yellow, elongate oval; BL 1.0-1.8 mm. Alatae have black dorsal abdominal cross-bands. On Salix, especially narrow-leafed species, in Japan, Thailand and Siberia (Pintera 1987, Pashchenko 1988a). In Europe, narrow-leafed Salix have very similar aphids, often regarded as a subspecies C. horii ssp. beuthani (Börner) (see influentialpoints.com/Gallery), and this form is also reported to occur in Kazakhstan (Kadyrbekov 2017a) and west Siberia (Stekolshchikov & Khureva 2020). However, Higuchi (1972) may have been correct in synonymising this with horii s. str., as the differences in form of the dorsal hairs, etc, could be ascribed to geographical variation. Apterous males and oviparae of the European form occur in October. Colonies are not ant-attended (see also Heie 1982).

Chaitophorus hypogaeus Hille Ris Lambers Apterae are broadly oval, dark dirty green, with paler greyish central area and pale antennae and legs; BL 1.4-2.0 mm. Alatae have narrow, usually interrupted, dusky to dark dorsal abdominal cross bands. In often large, ant-attended colonies on the subterranean parts of one-year-old branches of Salix repens (= S. rosmarinifolia, apparently restricted to plants growing in dry sandy soil (Hille Ris Lambers 1947b, Pintera 1982). Recorded from Belgium, Netherlands and Poland. A record from Turkey (Şenol et al. 2017) requires confirmation because of discrepancies in colour in life and feeding site. Apterous males and oviparae occur in October.

Chaitophorus indicus Ghosh, Ghosh & Raychaudhuri Apterae are elongate oval, pale green; BL 1.3-2.0 mm. Alatae have dark dorsal abdominal cross-bands. In colonies on undersides of both young and older leaves of Populus spp. (original record from Litsea is presumed to be an error). In India (Himachal Pradesh, West Bengal), Pakistan (Nasir & Yousuf 1994) and Bhutan (Chakrabarti & Das 2014, as manaliensis). A record from Turkey (Şenol et al. 2014b) requires further verification. Oviparae and alate males occur in Himachal Pradesh in late October (S.K. Das et al. 1981). Pintera (1987) regarded C. manaliensis Chakrabarti 1977, described from a ?Salix sp., but with oviparae described from P. alba (Agarwala & Mahapatra 1990), as a possible synonym, and this is confirmed by examination of paratype apterae of manaliensis in the BMNH collection. 2n=18. [A chromosome number of 2n=18 was recorded for an aphid identified as C. manaliensis forming pseudogalls on leaf borders of a Salix sp. (N.B. Pal & Khuda-Bukhsh 1983), but this was probably a misidentification (Pintera 1987). However, Dutta & Gautam (1993) recorded 2n=18 for a population from Populus ciliata in Himachal Pradesh which they identified as C. manaliensis.]

Chaitophorus inouyei Pintera Apterae are pale yellow; BL 1.4-1.9 mm. Alatae have broad black dorsal abdominal cross-bands. On leaves of Populus spp. (koreana, maximowiczii) in Japan, and also recorded from P. davidiana and P. suaveolens in Siberia (Kamchatka, Sakhalin, Primorskiya Kray; Pashchenko 1988a). Higuchi’s (1972) description of C. dorocola applies to this species. Oviparae were collected on the trunk of P. maximowiczii in October (Pintera 1987). Ch. xizangensis Zhang (in G. Zhang & Zhong 1981b) and Ch. wuduensis Zhang, Qiao & Chen (in G. Zhang 1999) from China are possibly inouyei. 2n=26.

Chaitophorus israeliticus Hille Ris Lambers Apterae are oval, colour in life not known, probably pale; BL 1.0-1.2 mm. Alatae and other morphs are unknown. Originally described from an undentified Salix sp. in Israel (see Pintera 1987); collected from Salix alba in Turkey (BMNH collection), and also recorded from Iran (Hodjat 1984a).

Chaitophorus kapuri Hille Ris Lambers Apterae are elongate oval, presumably blackish with pale legs and a paler spinal stripe from metanotum to about abdominal tergite 3; BL 1.4-2.0 mm. Alatae have broad dark dorsal abdominal cross-bands and wing veins conspicuously bordered with fuscous. On upper and undersides of leaves of Populus ciliata in Murree Hills of Pakistan (Hille Ris Lambers 1966). A record from Turkey (Görür et al. 2011a) requires confirmation. Oviparae and both apterous and alate males are recorded by Saha & Chakrabarti (1988c), who also record predators and attendant ants.

Chaitophorus lapponum Ossiannilsson Apterae are broadly oval, probably blackish with legs paler; BL 1.7-2.3 mm. Alatae have narrow dorsal abdominal cross-bands. On leaves, petioles an shoot apices of Salix spp. in Finland and Sweden, and there are also records from several Salix spp. in eastern Kazakhstan (Kadyrbekov 2009a,b, 2017a). Records from Salix purpurea in Austria and Germany refer to C. mordvilkoi (see Heie 1982, Pintera 1987), and records from Turkey (Kök & Özdemir 2021) require further confirmation. Sexual morphs are undescribed.

Chaitophorus leucomelas Koch (= versicolor Koch) Plate 12a,b Apterae are rather elongate oval, pale green or yellow, typically with black or dark green longitudinal pleural stripes which may be segmentally divided, and dark siphunculi (see influentialpoints.com/Gallery); BL 1.2-2.4 mm. Alatae have dark brown dorsal abdominal cross-bands and separate marginal sclerites. On young shoots of Populus spp. in spring, and later under leaves, in leaves stuck together by moth larvae, or leaf galls vacated by other insects; always ant-attended. Host plants in Europe are mainly P. nigra and related species and hybrids, but in North America a wider range of species is colonised. Development at different temperatures was studied by Hintze-Podufal & Thorns (1979). Oviparae and alate males occur in October-November; sexual morph production was studied by Thorns & Hintze-Podufal (1979). Common and widely distributed in Europe, and also in North Africa (Boukhris-Bouhachem et al. 2007; Laamari et al. 2013), Asia including Kazakhstan (Kadyrbekov 2017a), Mongolia, east Siberia (Pashchenko 1988a), and probably China, as Ch. populiyunnanensis Zhang (Zhang & Zhong 1985b) does not seem to be specifically distinct. Introduced into South Africa (Transvaal; BMNH collection, leg. I.M. Millar) and North and South America. Ramirez et al. (2004) compared susceptibility of various poplar hybrids to this aphid in Chile. 2n=36 (Israel) or 40 (UK, South Africa); the different numbers may indicate sibling species. [A report of 2n=4 for a population in Peru (Rubín de Celis & Ortiz 1993) needs additonal confirmation.]

Chaitophorus longipes Tissot Apterae are mainly brown, darker on head and thorax than on abdomen (Tissot 1932b); BL 1.7-2.1 mm. Alatae have pale brown abdomen with dark brown dorsal cross-bands and marginal sclerites. On young tender stems of Salix caroliniana in eastern USA.

Chaitophorus longisetosus Szelegiewicz Apterae are elongate oval, blackish-brown including siphunculi, with antennae and legs also mainly dark; BL 1.4-1.9 mm. Alatae have not been found. In small colonies mainly on older trees of Populus alba, on undersides of leaves, especially those spun together by other insects, not attended by ants. Oviparae, which are larger (BL 1.8-2.5 mm), occur in in the colonies in October, but males are unrecorded. In central and eastern Europe (Szelegiewicz 1961a, Pintera 1982, Wieczorek & Osiadacz 2007), Turkey (Kök & Özdemir 2021) and eastern Kazakhstan (Kadyrbekov 2009a).

Chaitophorus longiunguis Robinson Apterae are green; BL 1.6-1.9 mm. Alatae are undescribed, and biology is unknown. On an unidentified Salix sp. in Utah, USA (Robinson 1974). It is regarded as a synonym of C. eoessigi by Smith & Parron (1978).

Chaitophorus macgillivrayae Richards Apterae are yellow; BL c.1.9 mm. Alatae have abdomen greenish with a dark central patch. On an unidentified Salix sp. in eastern Canada (Quebec, New Brunswick). Very similar to C. pusillus and the pale form of C. viminalis.

Chaitophorus macrostachyae (Essig) Apterae are broadly oval, greyish to brownish or black, often with a pale central dorsal area; BL 2.0-2.8 mm. Alatae have dark dorsal abdominal cross-bands and marginal sclerites. Usually found on roots and parts of trunk of Salix spp. at or below ground level, but colonies may also form at tips of twigs (Essig 1912). In western North America. Four other species have been described from this microhabitat and resembling C. macrostachyae in most characters with the exception of the degree of sclerotisation and pigmentation of the dorsum, some differences in shape and length of hairs, and in chaetotaxy of abdominal tergite 8; they are C. abditus (Hottes), C. canens Richards, C. knowltoni Hille Ris Lambers and C. salicicorticis (Essig). None of these differences give reliable separation of paratype and other material in the BMNH collection, so they are all treated here as synonyms of macrostachyae. Ch. utahensis, however, synonymised with macrostachyae by Richards (1972a), is a distinct species (q.v.).

Chaitophorus matsumurai Hille Ris Lambers (= salicicola Matsumura 1917 nec. Essig 1911) Apterae are rather elongate oval, fuscous (according to original description, Matsumura 1917); BL 1.0-1.4 mm. Alatae have greenish-yellow abdomen with dark dorsal cross-bands. Other morphs and biology unknown. Originally described from Salix bakko in Japan, and specimens from Salix sp. in China and Korea are also this species, but many of the other host records (as salicicola Matsumura) in the literature are probably based on misidentifications (Pintera 1987).

Chaitophorus melanosiphon Pintera Apterae are short-bodied, oval, white; BL 0.8-1.7 mm. Alatae have not been recorded. On Populus alba in Romania, Iran, Turkey (Kök & Özdemir 2021), Kazakhstan (Kadyrbekov 2017a) and Pakistan (Baluchistan; Naumann-Etienne & Remaudière 1995). Oviparae and both apterous and alate males occur in October (Pintera 1987).

Chaitophorus minutus (Tissot) Apterae are uniformly yellow according to Tissot (1932b); BL 0.7-1.1 mm. Alatae are without distinct dorsal cross-bands. In dense colonies on undersides of leaves of Salix caroliniana in eastern USA (Florida, Louisiana, North and South Carolina).

Chaitophorus miyazakii Pintera Apterae are elongate oval, colour in life unknown, probably darkish; BL 1.5-1.8 mm. Other morphs and biology not known. On an unidentified Populus sp. in Hokkaido, Japan (Pintera 1987).

Chaitophorus monelli (Essig) Apterae are “transparently light green throughout” (Essig 1912a); BL 1.0-1.2 mm. Alatae have median dorsal black patches on pale green abdomen, only extending laterally as cross-bands on on posterior tergites. Found on tender shoots and suckers of Salix lasiolepis and S. laevigata in California, USA, and also now recorded from Oregon and New Mexico (aphidtrek.org). Hille Ris Lambers (1960a) redescribed this species. Sexuales and biology are unknown.

Chaitophorus mordvilkoi Mamontova Apterae are broadly oval, dark with a yellowish-green spinal stripe from back of head to middle of abdomen; BL 1.2-2.1 mm. Alatae have dark dorsal abdominal cross-bands. In small colonies on Salix purpurea, on shoot terminals and undersides of leaves, without ants (Pintera 1987), and also recorded from S. viminalis (Wieczorek & Osiadacz 2007). In central, southern and eastern Europe, and eastward to Mongolia (BMNH collection; leg. H. Szelegiewicz).

Chaitophorus murreensis Nasir & Yousuf Apterae are dark green: BL 1.9-2.1 mm. Described from 25 specimens collected on “Platinum” (error for Platanus) orientalis (Chanar) in Murreee, Pakistan (Nasir & Yousuf 1994). The host is anomalous for the genus and requires confirmation.

Chaitophorus nassonowi Mordvilko Apterae are brownish-red to brown, oval; BL 1.5-2.8 mm. Alatae have dark dorsal abdominal cross-bands and marginal sclerites. In ant-attended colonies on bark of young and older twigs or leaf petioles of Populus nigra and its varieties and hybrids, in eastern and central Europe. Apterous males and oviparae occur in October-November (Pintera 1987). [Chaitophorus chrysanthemi Hille Ris Lambers, described from apterae, oviparae and apterous males collected in September in Italy, with the host given (erroneously?) as Chrysanthemum sinense, has been synonymised by some authors with this species, but Pintera (1987) itemised some differences.]

Chaitophorus neglectus Hottes & Frison Apterae are usually yellowish-green to green with brownish head, prothorax and posterior abdomen, and typically with a pair of dark pleural stripes of varying width running from thorax to abdominal tergite 6; BL 1.7-2.5 mm. Alatae have dark dorsal abdominal cross-bands and dark siphunculi. On upper and undersides of leaves, or in leaves folded and stuck together by other insects, of aspens (Populus tremuloides, P. grandidentata) and of P. deltoides. Colonies are often quite large, not attended by ants (but sometimes mixed with C. stevensis which is ant-attended). Oviparae and both apterous and alate males occur in September-November. Widely distributed in North America. Records from Turkey (Görür et al. 2012) need confirmation. This aphid is very similar to, and often confused with, C. populifolii, but is probably a good species; see Richards (1972a), and Hille Ris Lambers (1960a; as populifolii sspp. neglectus and simpsoni). 2n=12.

Chaitophorus niger Mordvilko Apterae are blackish-brown with mainly pale antennae and legs, a thin pale ring around the base of each siphunculus, and a pale cauda (see influentialpoints.com/Gallery); BL 1.0-2.4 mm. Nieto Nafrìa & Mier Durante (1998), however, found considerable variation in dorsal sclerotisation in Spanish populations, with some individuals entirely pale. Immatures are wine-red. Alatae have brown dorsal abdominal cross-bands. Living separately or in small colonies on leaves of various Salix spp., only rarely visited by ants. Oviparae and males occur in September-November; whether males are apterous or alate seems to depend on geographical location (Lampel 1983). In Europe and across Asia to east Siberia (Pashchenko 1988a), but apparently not in China or Japan. Closely related to, and often treated as a subspecies of, C. salijaponicus. 2n=30.

Chaitophorus nigrae Oestlund (= salicicola Essig) Apterae are usually black, often with a paler spinal area which rarely extends to abdominal tergite 7, with femora of mid and hind legs usually dark brown, but paler specimens may also occur with variably developed dark pigmentation of metathorax and abdomen confined to pleural areas only (see influentialpoints.com/Gallery); BL 1.7-2.5 mm. Alatae have dark dorsal abdominal cross-bands, often coalescent beween abdominal tergites 2-6. First instar are usually black, brown or dark green (cf. C. pallipes). In sometimes large colonies on leaves and petioles of new growth of various Salix spp. Ant attendance is not recorded. Large, pale oviparae and alate males occur in September-November (BMNH collection). Widely distributed in North America. This is an extremely variable species, or possibly a complex of species, varying not only in pigmentation but in the reticulation of the dorsal cuticle, the form of the dorsal hairs and the shape of the cauda. We follow Richards (1972a) in treating Ch. salicicola, and its subspecies described by Hille Ris Lambers (1960a), as synonyms. 2n=24*.

Chaitophorus nigricantis Pintera Apterae are elongate oval, colour in life unknown but probably pale; BL 1.4-2.3 mm. Alatae have brownish dorsal abdominal cross-bands, coalescent on abdominal tergites 3-6. Recorded from Salix spp. (nigricans, phylicifolia, triandra) in Sweden, Finland (Pintera 1987) and north-west Russia (Stekolshchikov & Buga 2018; Stekolshchikov & Khureva 2020). Apterous and alate males collected in July-August on Salix phylicifolia were described by Stekolshchikov & Buga (2018), indicating an abbreviated life cycle. A form on Salix sp. in Mongolia is distinguished as a subspecies, mongolicus Pintera. This species has sometimes been confused in the literature with C. truncatus (q.v).

Chaitophorus nigricentrus Richards Apterae are broadly oval, dark yellow-brown to blackish-brown, with central part of dorsum darker than margins (see influentialpoints.com/Gallery); BL 1.5-1.8 mm. Alatae have dark dorsal abdominal cross-bands. Found only on second-year growth of (unidentified) Salix sp(p); recorded from Ontario, Canada (Richards 1972a) and Pennsylvania, USA (BMNH collection). Biology and sexual morphs are unknown.

Chaitophorus nigritus Hille Ris Lambers Apterae are rather elongate oval, black with pale antennae and legs; BL 1.0-1.6 mm. Alatae have not been recorded. In compact colonies around galls on the leaves of unidentified Salix sp(p). in Pakistan and Iran (Hille Ris Lambers 1966c, Pintera 1987), and Turkey (Güçlü et al. 2015), and since identified from Salix alba in Iraq and Himachal Pradesh, India. Oviparae and one alate male were collected in Iran in December (BMNH collection; leg. S. Hodjat). 2n=18 (Dutta & Gautam 1993).

Chaitophorus nodulosus Richards Colour of apterae in life is not recorded probably dark brown to black; BL about 1.5-1.7 mm. Alatae and sexual morphs are unknown. Described from an unidentified Populus sp. in Utah, USA (Richards 1972a), and subsequently found in Colorado (aphidtrek.org).

Chaitophorus nudus Richards Apterae are broadly oval, dark brown to shiny black, with antennae and legs also dark (see influentialpoints.com/Gallery); BL 2.0-2.6 mm. Alatae have dorsal abdomen blackish or dark green with black spots or incomplete cross-bands, and forewing veins black and strongly bordered, with a very conspicuous black pterostigmal spot. In rather small colonies girdling trunk of saplings of Populus tremuloides (Richards 1972a). Large pale oviparae were collected in September in Colorado (BMNH collection). Recorded from Canada (Manitoba, Ontario) and USA (Colorado, Pennsylvania; BMNH collection).

Chaitophorus pakistanicus Hille Ris Lambers Apterae are oval, pale green; BL 1.2-1.7 mm. Alatae are undescribed; two specimens in the BMNH collection have no dorsal abdominal markings. In dense colonies along the veins on both sides of leaves of Salix spp. in central and east Asia (Afghanistan, Iran, India, Pakistan, China, Thailand). Oviparae were collected on S. acmophylla in December (Hille Ris Lambers 1966c, Pintera 1987). Bhagat (1982b) described a new species of parasitoid from Ch. pakistanicus in Kashmir. Ch. dulciculus Zhang, Qiao & Chen, described from Salix sp. in north-west China (G. Zhang 1999), is very similar and likely to be a synonym.

Chaitophorus pallipes Richards Apterae are yellow with broad black pleural stripes, and femora usually pale, at least on fore and middle legs (see influentialpoints.com/Gallery); BL 1.8-2.4 mm. Alatae have black dorsal abdominal cross-bands. First instars are yellow (cf. nigrae). On young growth of Salix discolor (BMNH collection; R.L. Blackman), and possibly other Salix spp., in Ontario, Canada, and also recorded from British Columbia, Prince Edward Island, and in USA: North Carolina (Richards 1972a), Idaho, Montana, California and Oregon (aphidtrek.org). Ant-attended. The synonymy of pallipes with saliciniger by Eastop & Hille Ris Lambers (1976) was in error.

Chaitophorus parvus Hille Ris Lambers Apterae are black with pale antennae and legs; BL 1.2-1.8 mm. Alatae have narrow dorsal abdominal cross-bands. On undersides of leaves of Salix repens (= rosmarinifolia) and S. viminalis, attended by ants. Recorded from northern Europe (Poland, Latvia, Denmark, Norway and Sweden), and also from Spain (on S. purpurea ssp. lambertiana; Nieto Nafría & Mier Durante 1998). Pintera (1987) synonymised this species with niger, but it appears distinct.

Chaitophorus pentandrinus Ossiannilsson Apterae are oval, pale green or yellow; BL 1.4-2.3 mm. Alatae have dark dorsal abdominal cross-bands and marginal sclerites. Only known from Salix pentandra and S. myrsinites in Sweden, and also found in Poland (Wieczorek & Osiadacz 2007), Russia (Altai Krai) and Kazakhstan (Kadyrbekov 2017a). Ant-attended. Males are apterous (see Heie 1982, Pintera 1987).

Chaitophorus pheleodendri Juchnevitch Apterae are elongate oval, colourless; BL 1.2-1.4 mm. Alatae have brown dorsal abdominal cross-bands. In colonies on undersides of leaves of Phellodendron amurense (Rutaceae), introduced as an ornamental into east Kazakhstan (Juchnevitch 1970). (Possibly these were aberrant colonies of C. horii.)

Chaitophorus populeti (Panzer) Apterae are oval, shiny dark green to black, with antennae and legs mid-green to black (with ANT III paler); BL 1.5-2.9 mm. Immatures usually have a distinctive dorsal pattern of dark markings (see influentialpoints.com/Gallery). A golden yellow colour variant has also been reported (Dransfield & Brightwell 2015). Alatae have broad brown dorsal abdominal cross-bands. On young shoots and terminal leaf petioles of various Populus spp. (especially of the alba, tremula groups), usually attended by ants, throughout the palaearctic region. Oviparae and both apterous and alate males occur in October-November. A subspecies, Ch. populeti sensoriatus Mimeur, is believed to occur on P. alba in the Mediterranean region and eastward to Afghanistan (see Pintera 1987) and this form is also recorded from western Siberia (Stekolshchikov & Novgorodova 2013), although populations with intermediate characters occur in Spain (Nieto Nafría & Mier Durante 1998). Ch. frigidus Zhang, Qiao & Chen, described from P. ×canadensis in north-west China (G. Zhang 1999), may be related. The male genitalia were described and illustrated by Wieczorek et al. (2011). 2n=12 (samples from Iran and China; but Pal & Khuda-Bukhsh record 2n=10 from India).

Chaitophorus populialbae (Boyer de Fonscolombe) Apterae are short-bodied, oval, greenish to yellowish white, often with small green spots (see influentialpoints.com/Gallery); BL 1.0-2.3 mm. Alatae have dark brown dorsal abdominal cross-bands, often coalescent on abdominal tergites 2-6. In small colonies usually on undersides of leaves of various Populus spp. sometimes ant-attended. Throughout the palaearctic region, in northern, western and southern Africa, and introduced and widespread in North America. Oviparae and both apterous and alate males occur in September-November. A subspecies, yomefuri Shinji, occurs on P. alba and P. sieboldii in Mongolia, Korea and Japan (see Pintera 1987). Ch. diversifolii Juchnevitch, described from P. diversifolia (= P. euphratica) in Kazakhstan, is probably a synonym of Ch. populialbae, and Ch. floris Zhang, Qiao & Chen, described from north-west China (G. Zhang 1999), is also not clearly distinct. 2n=28 (Chen & Zhang 1982) or 30.

Chaitophorus populicola Thomas Apterae yellow-brown to shiny black, with dark antennae and legs (see influentialpoints.com/Gallery); BL 2.0-2.5 mm. Alatae have dorsal abdominal cross-bars or bands and forewing veins dark and conspicuously brown-bordered. In often dense, usually ant-attended colonies on young shoots, developing leaves and leaf petioles of numerous Populus spp. throughout North America, sometimes attaining pest status in plantations of P. deltoides (Coleman & Jones 1988). Large, pale oviparae and alate males occur in September-November. This species is very similar to Ch. nudus, but differs in chaetotaxy and feeding habit (Richards 1972a). A subspecies, patchae Hille Ris Lambers, was erected for populations with long, pointed dorsal hairs. 2n=18, 28 and 32 in different populations, indicating that a complex of species is involved under the name populicola.

Chaitophorus populifolii Essig (= balsamiferinus Hille Ris Lambers; = populellus Gillette & Palmer; = essigi Gillette & Palmer) Apterae are yellowish, sometimes with variably developed brown to black pigmentation of head, pro- and mesothorax and abdominal tergites 7 and 8, and occasionally with a pair of dark, usually incomplete, pleural stripes; BL 1.0-2.3 mm (dwarf apterae occur in summer and have dark legs and 5-segmented antennae). Alatae have variably developed dark dorsal cross-bands and pale siphunculi. On leaves of Populus spp., especially angustifolia and balsamifera, sometimes on deltoides or tremuloides (Richards 1972a). Widespread in USA and Canada, Mexico, and introduced into Europe (Germany), probably on balsam poplar cuttings (Pintera 1987). Recently it has been reported from Serbia (Poljaković-Pajnik & Petrović-Obradović 2009) and Turkey (Kök & Özdemir 2021). Oviparae and both apterous and alate males occur in September-November. There has been considerable confusion of nomenclature of this taxon – see Sorenson (1989). 2n=12.

Chaitophorus populihabitans Zhang Apterae (BL c. 2 mm) and alatae were described from Populus alba in Yunnan, China (G. Zhang & Zhong 1985g), and there is a subsequent record from P. tomentosa (G. Zhang et al. 1992c). This species could not be included in the key to aphids on Populus on the basis of the published description.

Chaitophorus pruinosae Narzikulov Apterae are dark brownish-green; BL 1.6-2.1 mm. On undersides of leaves of Populus pruinosa in Tadzikhistan, and also collected from P. euphratica in Kazakhstan (Kadyrbekov 2017a) and P. nigra in Afghanistan (BMNH collection). Males are alate. Pintera (1987) described the fundatrix. Ch. xinjiangalbus Zhang, Qiao & Chen, described from north-west China on P. euphratica, and Ch. dabanchengus, described by the same authors from both Populus sp. and Salix sp. (in G. Zhang 1999), will probably key to Ch. pruinosae, and may be this species.

Chaitophorus purpureae Lee & Seo Apterae are green; BL c. 1.6 mm. Alatae have dark head and thorax but are apparently without dorsal abdominal sclerotisation. On undersides of leaves of a Salix species identified as “S. purpurea var. smithiana” (= S. sinopurpurea?) in Korea (Lee & Seo 1992). In the key to Chaitophorus on Salix this species will probably go to C. viminalis (a North American species with apterae dark in later generations, and alatae with dark dorsal abdominal sclerotisation).

Chaitophorus pusillus Hottes & Frison Apterae are whitish, BL 0.9-1.3 mm. Alatae have dark dorsal cross-bands coalescing into a central patch. On upper and undersides of leaves of Salix amygdaloides in Illinois, USA (Hottes & Frison 1931), and also recorded from S. discolor and unidentified Salix spp. in other parts of USA and Canada. C. macgillivrayae Richards, described from a Salix sp. in eastern Canada is likely to be a synonym.

Chaitophorus pustulatus Hille Ris Lambers Appearance in life has not been recorded but apterae are probably variably pigmented, brown to black with paler central dorsal area; BL 1.0-1.9 mm. Alatae have not been described; those in the BMNH colection have dark dorsal abdominal cross-bands. On leaves of Salix sp. in Utah, Coloradoa and Wyoming (Hille Ris Lambers 1960a), and also collected in British Columbia (Richards 1972a), Idaho and Oregon (BMNH collection) – in all cases from unidentified Salix. Small pale oviparae and apterous males have been collected in Colorado in October (BMNH collection, leg. F.C. Hottes).

Chaitophorus quinquemaculatus Bozhko Apterae are elongate oval, whitish with three longitudinal stripes and two (?pleural) spots (Bozhko 1976); BL 1.2-2.2 mm. Alatae have brownish dorsal abdominal cross-bands. In dense colonies on twigs, or rarely on undersides of leaves, of Salix acutifolia, and also recorded from S. appenina, S. caprea, S. caspica, S. fragilis and S. pentandra. In Italy (Sicily), Ukraine and Central Asia. Pintera (1987) regarded it as a subspecies of C. saliapterus.

Chaitophorus ramicola (Börner) Apterae are broadly oval, mainly very dull greyish olive to black with a more-or-less distinct paler spinal stripe, and dark antennae and legs; BL 1.4-2.6 mm. Alatae have narrow dorsal abdominal cross-bands. In ant-attended colonies on bark of twigs of mainly broad-leafed Salix spp., in north, central and eastern Europe, Turkey (Şenol et al. 2014b), Iran (Hodjat 1993) and Kazakhstan (Kadyrbekov 2017a). Apterous males and oviparae occur in October (Heie 1982). C. bruneolineatus Juchnevitch 1968, described from S. rossica (= viminalis) in Kazakhstan, is probably a synonym.

Chaitophorus remaudierei Pintera Apterae are elongate oval, BL 1.4-2.0 mm, probably variably dark-pigmented but colour in life has not been recorded. Alatae are undescribed. On an unidentified Salix sp. in Iran (Pintera 1987). Apterous males and oviparae were collected in October.

Chaitophorus saliapterus Shinji Apterae are elongate oval, pale green; BL 1.0-2.0 mm. Alatae have dark dorsal abdominal cross-bands. On undersides of leaves of Salix spp. (especially integra), attended by ants, in eastern and western Siberia (Pashchenko 1988a, Stekolshchikov & Novgorodova 2013), Japan and Korea (Pintera 1987). 2n=30 (Shinji’s record of 2n=14 (n=7) should probably be applied to another species of Chaitophorus).

Chaitophorus saliciniger (Knowlton) Apterae are variably pigmented but often black, with a paler, spindle-shaped, spinal stripe; BL 1.7-2.2 mm. Alatae have dark dorsal abdominal cross-bands. On bark of twigs of unidentified Salix sp(p). in Colorado and Utah, USA. A record from Turkey (Eser et al. 2009) is doubtful and requires further confirmation.

Chaitophorus salicti (Schrank) Apterae are black with a pale spinal stripe in spring, but small apterae in late summer are yellowish white or pale yellowish green with reddish-brown or greyish-black dorsal markings (see influentialpoints.com/Gallery); BL 1.3-1.8 mm. Alatae are dark, with broad dorsal abdominal cross-bands. On undersides of leaves of broad-leafed Salix spp., usually along main veins, attended by ants. Throughout Europe, south-west and central Asia, and also recorded from China (Zhang et al. 1986). Apterous males and oviparae occur in September-October (Heie 1982, Pintera 1987). 2n=28.

Chaitophorus salijaponicus Essig & Kuwana Apterae are presumably dark brown or black, sometimes with paler narrow spinal stripe; BL 1.3-1.9 mm. Alatae have dark dorsal abdominal cross-bands. On leaves of Salix spp. in Japan, China and Siberia. Very similar to C. niger, which is often treated as a subspecies. Two other forms in this group are distinguished as subspecies by Pintera (1987); Ch. salijaponicus ssp. stroyani from S. phylicifolia in Italy, and Ch. salijaponicus ssp. szelegiewiczi from Salix sp. in Mongolia, the latter now also recorded from S. caprea in western Siberia (Altai Republic; Stekolshchikov & Novgorodova 2013) and Kazakhstan (Kadyrbekov 2017a).

Chaitophorus saliniger Shinji Apterae are shining blackish-brown to black with legs and distal parts of antennae dark and cauda conspicuously paler; BL 1.0-1.8 mm. Alatae have broad black dorsal abdominal cross-bands. On young leaves, petioles or shoot tips of narrow-leafed Salix spp., often ant-attended, in Japan, China and Korea. Apterous males and oviparae occur in November-December (Higuchi 1972, Pintera 1987). Genetic variation of this species in China has ben investigated by F. Fang et al. (2017) and Zhu et al. (2017), providing evidence of divergence between northern and southern populations. 2n=8.

Chaitophorus shaposhnikovi Mamontova Apterae are whitish with five greenish spots on dorsum (Mamontova 1955); BL c.1.2-1.3 mm. Alatae are undescribed. On undersides of leaves, along main veins, of an unidentified Salix sp. in Ukraine and Kazakhstan (Shaposhnikov 1964a). Hosts subsequently named are Salix caspica and S. turanica (Kadyrbekov 2017a).

Chaitophorus similis Pintera Colour of aptera in life is unknown, presumably pale; BL 0.8-1.3 mm. Alatae are unknown. On Salix incana in southern France (Pintera 1987). Apterous males occur in September.

Chaitophorus stevensis Sanborn Apterae are pale green with darker dorsal markings (see influentialpoints.com/Gallery); BL 1.6-2.0 mm. Alatae have black head and thorax, greenish abdomen with dark cross-bands often coalesced into a central patch, and darkish siphunculi (Richards 1972). Ant-attended, on leaves of Populus spp., apparently restricting its feeding to aspens and cottonwoods; records from balsam poplars are probably misidentified populifolii. It is frequently associated with leaf-fold galls of Thecabius populiconduplifolius. Widely distributed in North America, aand also recorded from Costa Rica (Villalobos Muller et al, 2010). 2n=14.

Chaitophorus tremulae Koch Apterae are elongate oval, dark brown to black, sometimes with a paler spinal stripe, legs brown with hind pair darker (see influentialpoints.com/Gallery); BL 1.2-2.5. Alatae have very broad and nearly coalescent dorsal abdominal cross-bands. Immatures are bright green. In small colonies on undersides of leaves of P. tremula and a few related species, often in leaves spun together by other insects, or in leaf-nest galls of Eriosomatinae. Not usually ant-attended. Oviparae and alate males occur in October. The male genitalia were described and illustrated by Wieczorek et al. (2011). Gärdenfors (1986) described a specific parasitoid in Sweden. Ch. tremulae occurs throughout Europe (Heie 1982, Pintera 1987, Nieto Nafría & Mier Durante 1998), and in Turkey, Transcaucasia and Centra Asia (Kadyrbekov 2017a). Pintera classified two related East Asian forms as subspecies of tremulae; shantungensis Tseng & Tao 1936 on Populus spp. in China and Japan, and a pale form living on leaves or in Pemphigus galls on Populus laurifolia in Japan and Mongolia (ssp. sorini Pintera). The latter form is also recorded from P. tremula in west Siberia and Kazakhstan (Stekolshchikov et al. 2008a, Kadyrbekov 2014d). A third subspecies, Ch. tremulae spp. lanzhouensis Zhang, Qiao & Chen, was described from north-west China (G. Zhang 1999). 2n=18*.

Chaitophorus truncatus (Hausmann) Apterae are elongate oval, pale green in spring and often with three dark green longitudinal stripes (see influentialpoints.com/Gallery), but in summer sometimes much darker or even blackish, with mainly pale legs and antennae; BL 1.2-2.4 mm. Alatae have variably-developed dorsal abdominal markings, usually not forming solid cross-bands. In small colonies on leaves of various narrow-leafed Salix spp., throughout Europe and east to Iran and Kazakhstan (Heie 1982, Pintera 1987, Kadyrbekov 2005c). Not ant-attended. Apterous males and oviparae in October-November. 2n=30.

Chaitophorus utahensis (Knowlton) Apterae are broadly oval, colour in life not noted but presumably rather dark; BL 2.0-2.5 mm. Alatae have dark dorsal abdominal cross-bands, and dark wing veins with narrow fuscous borders. On bark of twigs and extending onto leaf petioles, or on trunk 15-20 cm above ground level (BMNH collection; leg. G.F. Knowlton), of unidentified Salix sp(p)., ant-attended, in western USA (Utah, Colorado, California, Idaho, Oregon, Washington; Hille Ris Lambers 1960a and BMNH collection).

Chaitophorus variegatus Szelegiewicz Apterae are greenish-yellow with sometimes darker head and thorax; BL 1.4-2.3 mm. Other morphs are unknown. On undersides of leaves of Populus koreana in Korea, attended by ants (Szelegiewicz 1981b).

Chaitophorus viminalis Monell Apterae are of varying pigmentation in life, usually pale green to yellow in spring with darker green pleural stripes but often dark brown to black in summer (see influentialpoints.com/Gallery); BL 1.3-1.9 mm. Alatae have dark dorsal abdominal cross-bands often coalesced into a central patch. On leaves and young growth of various narrow-leafed Salix spp. throughout North America. Oviparae and alate males occur in October-November. Probably often misidentified in the literature; differences from similar-looking species (e.g. nigrae, pusillus) are discussed by Hille Ris Lambers (1960a). See also Richards (1972a). 2n=18.

Chaitophorus viminicola Hille Ris Lambers Apterae are rather elongate oval, brown or blackish, usually with a paler spinal stripe from pronotum to about abdominal tergite 3 (see influentialpoints.com/Gallery) ; BL 1.9-2.5 mm. Alatae have broad dark dorsal abdominal cross-bands. On Salix nigra (BMNH collection), S.alba var. sericea (Favret et al. 2010) and Salix sp(p). in eastern USA, and Canada (Ontario). Biology and sexual morphs are unknown.

Chaitophorus vitellinae (Schrank) Apterae are broadly oval, yellowish-green with two broad darker green to greenish-brown pleural longitudinal stripes, and pale antennae and legs (see influential points/gallery); BL 1.3-2.3 mm. Alatae have dark dorsal abdominal cross-bands. On bark of young twigs and leaf petioles of narrow-leafed Salix spp., especially S. alba, attended by ants. Throughout Europe, and east to west Siberia, Iran and Kazakhstan (Kadyrbekov 2017a). Pintera (1987) described a subspecies (danubicus), with a distinctly knobbed cauda and thickened dorsal hairs, from S. alba in Rumania, Czechoslovakia and Turkey. Apterous males and oviparae occur in September-October (Heie 1982). [The molecular phylogeny of Shingleton & Stern (2003) indicates that C. vitellinae is closely related to Populus-feeding species, and may have crossed to Salix relatively recently.]

Chaitoregma Hille Ris Lambers & Basu

Hormaphidinae: Cerataphidini

Bamboo-feeding aphids characterised by the rounded frontal horns and the heavily sclerotised dorsum.

Chaitoregma tattakana (Takahashi) Apterae are yellowish-brown to deep violet-black, covered with fine wax dust; BL 1.5-1.9 mm. In large colonies on undersides of leaves of bamboos, (Bambusa, Arundinaria, Phyllostachys and Yushania). Recorded from India, Taiwan and China (Hille Ris Lambers & Basu 1966, Liao 1976, Huang et al. 2012, Qiao et al. 2018). C. aderuensis (Takahashi 1935f), from Bambusa sp. in Taiwan, is not clearly distinct. The life cycle is unknown. [The galls on Ligustrum in Hong Kong ascribed to a Chaitoregma sp. by Blackman & Eastop (1994) are now thought to be those of Ceratopemphigus zehntneri.]

Chakrabartiaphis Remaudière

Aphidinae: Macrosiphini

One species on Hydrangea in India, very close to Eumyzus and forming similar leaf-roll galls, but with longer, finely pointed dorsal hairs and a cauda similar to that found in moss-feeding aphids. Chakrabartiaphis is a replacement name for Tubicauda Chakrabarti & Bhattacharya 1982, which is preoccupied (Remaudière 2001).

Chakrabartiaphis hydrangeae (Chakrabarti & Bhattacharya) In reddish tubular marginal leaf galls on upper surfaces of leaves of Hydrangea sp. in Uttar Pradesh, India. The host was subsequently identified as H. altissima (= anomala) (Chakrabarti & Banerjee 1993a, as Tubicauda hydrangeae). The colour of apterae in life is unrecorded; BL c.1.6 mm. Alatae produced in early June have 25-28 secondary rhinaria on ANT III and 1-4 on IV. The life cycle is unknown.

Chitinosiphon Yuan & Xue

Aphidinae: Macrosiphini

(= Radiaphis Pashchenko 2000; = Radicisiphum Zhang, Chen, Zhong & Li, in Zhang 1999)

Five or six species in east Asia somewhat resembling Aulacorthum but adapted for root feeding and ant attendance on Asteraceae, with rather low, divergent antennal tubercles, ANT III of aptera with 3-16 rhinaria spread out on basal half, extensive dorsal sclerotisation, siphunculi with distal part heavily imbricated, and a short dark cauda. Pashchenko (2000c) keyed the 5 species known from east Siberia (as Radiaphis).

Chitinosiphon abdomenigrum Yuan & Xue (= Radiaphis saussureae Pashchenko 2000) Apterae are white with a large greyish green dorsal abdominal patch; BL c.2.0 mm. On Saussurea nigrescens in Shaanxi Province, China, and also found on basal part of stem of Saussurea sp., attended by ants, in Amur Province, east Siberia (Pashchenko 2000c, as Radiaphis saussureae).

Chitinosiphon cardui (Pashchenko) Apterae are yellow-green with a sclerotised dorsal abdominal patch; BL c. 2.3 mm. On roots of Carduus crispus, attended by ants (Pashchenko 2000c, as Radiaphis). In east Siberia (Primorskii).

Chitinosiphon cirsorhizum (Zhang, Chen, Zhong & Li) (= Radiaphis cirsii Pashchenko 2000; = Unisitobion cirsiariston Zhang, Chen, Zhong & Li, in Zhang 1999) Apterae are yellow-green with a large greenish-brown dorsal abdominal patch, shining or pruinose; BL 1.9-2.5 mm. On underground part of stem and roots of Cirsium spp. in north-west China (Zhang 1999, as Aulacorthum) and east Siberia (Pashchenko 2000c, as Radiaphis cirsii). Monoecious holocyclic with oviparae and alate males in October (An ovipara with barely swollen hind tibiae bearing few scent glands, indicating lack of host alternation, was on a slide labelled as the holotype of Unisitobion cirsiariston.)

Chitinosiphon doellingeriae (Pashchenko) Apterae are whitish green with a brownish green sclerotised dorsal abdominal patch; BL c.2.0 mm. On basal part of stem and roots of Aster scaber (= Doellingeria scabra), attended by ants (Pashchenko 2000c, as Radiaphis). In east Siberia (Primorskii Territory).

Chitinosiphon kalimeris (Pashchenko) Apterae are green with a dark greenish gray dorsal abdominal patch; BL c.2.4 mm. On roots of Kalimeris incisa attended by ants (Pashchenko 2000c, as Radiaphis). In east Siberia (Primorskii Territory.).

Chomaphis Nevsky

Aphidinae: Aphidini

One species in Europe and west Asia on roots of Asteraceae, related to Aphis but with first tarsal segments bearing 4-4-3 hairs, a very short cauda, long siphunculi bearing small hairs, and very large marginal tubercles.

Chomaphis mira Mordvilko (Fig.54b,c) Colour of apterae in life is unrecorded; BL 2.0-2.5 mm. On roots of Petasites, Tussilago and Arctium. In Russia, Ukraine and Kazakhstan, and an alata has been recorded from northern Italy by van Harten & Cocceano (1981), who provided a redescription. 2n=8.

Chondrillobium Bozhko

Aphidinae: Macrosiphini

One species in Europe on Chondrilla (Compositae/Asteraceae) superficially resembling Hyalopteroides but perhaps closer to Pleotrichophorus.

Chondrillobium blattnyi (Pintera) Apterae are pale green, BL 1.8-2.9 mm. On undersides of leaves of Chondrilla juncea, causing yellowing. In southern Europe and eastward to Bulgaria, Greece, Turkey, Iran and Ukraine, Russian Volga region, and recorded from C. canescens in south-east Kazakhstan (Kadyrbekov & Aoitzhanova 2005). Nieto Nafría (1974) provided a detailed redescription of apterae and alatae. Oviparae and apterous males were described from Turkey (Tuatay & Remaudière 1965). Populations studied by Caresche et al. (1974) in the Mediterranean area were anholocyclic.

Chromaphis Walker

Calaphidinae: Panaphidini

Two palaearctic species of small pale yellow aphids, related to Panaphis and likewise associated with Juglans. Both are also recorded from Aleurites moluccana (Euphorbiaceae), which presumably acts as a “reserve host”. Trioxys pallidus and Praon flavinode specialise on Chromaphis and the related genera Eucallipterus, Myzocallis, Tinocallis and Tuberculatus (Starý 1976). Aphelinus perpallidus attacks Melanocallis and Monellia as well as Chromaphis. Quednau (2003) reviewed and illustrated the genus.

Chromaphis hirsutustibus Kumar & Lavigne Adult viviparae all alate, pale yellow, often with black patches on abdominal tergites 4 and 5, and with dark spots at ends of wing veins; BL 1.6-2.5 mm. On Juglans regia in India, Pakistan, Bhutan (Chakrabarti & Das 2014), Nepal and China (G. Zhang & Zhong 1985); both immatures and adults are recorded also from Aleurites moluccana in India and China. Chakrabarti (1988) described the oviparae, collected in September. The sexual morphs described under this name by Chakrabarti (1978) were C. juglandicola.

Chromaphis juglandicola (Kaltenbach) Adult viviparae all alate, pale lemon yellow or yellowish-brown with pale brown thoracic lobes and (in autumn) paired brown spots on abdominal tergites 4 and 5 (see influentialpoints.com/Gallery); BL 1.2-2.3 mm. Immatures can have paired brown spots on more of the abdominal tergites, and oviparae (the only apterous morph) have broad black bands across the thorax and abdomen. Living scattered under leaves on Juglans regia, with occasional records from other Juglans spp., and there is also a record from Aleurites moluccana (of oviparae; Chakrabarti 1978, as hirsutustibus). In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), Middle East, Central Asia, India, Pakistan, China and North and South America. Oviparae and males occur in May-June in India, and August-November in Europe, in April in Chile (Salas et al. 2016) and in late September-October in USA. Biology and ecology have been studied intensively, especially in California (e.g. Nowierski & Gutierrez 1986), where biological control has been successful (Hoy & Cave 1988, Hougardy & Mills 2009). 2n=8.

Chromocallis Takahashi

Calaphidinae: Therioaphidini

Two species on Ulmus in east Asia. They are plump-bodied aphids with rather short antennae, thick dark hairy tibiae and a deeply cleft anal plate. Quednau (2003) reviewed the genus.

Chromocallis nirecola (Shinji) Adult viviparae are all alate, green, with black tibiae and forewing veins bordered with fuscous; BL 3.0-3.2 mm. On Ulmus spp (japonica, pumila) in Japan, Korea, China and east Siberia (Higuchi 1972, Pashchenko 1988a, Qiao et al. 2005b). Biology is little known; alate males were collected in September in Japan (BMNH collection, leg. M. Sorin), and Paik (1972) recorded oviparae and males in Korea. C. pumili Zhang in Zhang & Zhong 1982c is a synonym (Quednau 2003). 2n=18*.

Chromocallis similinirecola Zhang Appearance in life is unknown; BL c.2.9 mm. On Ulmus pumila in China (Zhang & Zhong 1982c).

Chuansicallis Tao

Calaphidinae: Panaphidini

A genus for one east Asian species with a well-defined median suture on the head, a very long third antennal segment with regularly-spaced, slit-like secondary rhinaria and a rather short processus terminalis. Quednau (2003) reviewed the genus.

Chuansicallis chengtuensis Tao All adult viviparae are alate, green, secreting profuse cottony wax threads, with dark hind legs and wings held horizontally over the abdomen when at rest; BL 1.6-2.0 mm. The life cycle and host relationships are unclear; numerous alate males and apterous oviparae were described from Rhus chinensis (= javanica) in December in Taiwan (Tao 1963), but alate viviparae have been collected from Dalbergia hupeana in November in Taiwan, and also in May-June in China (Kwangtung, Szechuan; Zhang & Zhong 1980c, as Sinotherioaphis pterothorax). Alatae were also recorded from Phyllanthus sp. in Bhutan (A.K. Ghosh & Quednau 1990).

Chucallis Tao

Calaphidinae: Panaphidini

A genus for two east Asian bamboo-feeding species with very large dorsal abdominal tubercles. Quednau (2003) reviewed the genus.

Chucallis bambusicola (Takahashi) (fig. 30E) Adult viviparae are all alate, dark purple to black with pale antennae and legs and clear wings; BL 1.6-2.0 mm. On shoots and undersides of leaves of bamboos (Bambusa stenostachya, Dendrocalamus latiflorus, Pseudosasa cantorii) in Taiwan and China (Kwangtung, Szechuan, Tao 1964; Hong Kong, Martin & Lau 2011). Very active, jumping when disturbed. Anholocyclic in Taiwan (Takahashi 1923; as Myzocallis bambusicola), and sexual morphs are unrecorded.

Chucallis latusigladius Jiang, Chen & Qiao Adult viviparae are all alate, with pale brown head and thorax, dark green abdomen bearing very large sword-shaped dark green marginal processes on abdominal tergite 4, and a conspicuous dark brown crescent-shaped mark on the pterostigma of each forewing; BL 1.6-2.7 mm. On undersides of leaves of Indocalamus tessellatus in Zhejiang and Fujian provinces, China (Jiang et al. 2011a). Sexual morphs and life cycle are unknown.

Chusiphuncula Zhang

Aphidinae: Macrosiphini

A genus for one species in China on Sorbaria, compared by the author with Rhopalosiphoninus, but probably more closely related to Wahlgreniella.

Chusiphuncula sorbarisucta Zhang (= Himalayaphis sorbarisorbifoliae Zhang, Chen, Zhong & Li) Apterae are green; BL c.2.0 mm. An alata had 7 secondary rhinaria on ANT III. On Sorbaria sorbifolia in north-west China (Zhang & Chen 1998).

Cinara Curtis

Lachninae: Eulachnini

A very large genus resulting from a highly successful adaptive radiation on Coniferae of the families Pinaceae and Cupressaceae. As understood here, Cinara includes Cedrobium, Cupressobium and Schizolachnus as subgenera, and contains more than 200 described species, about 150 of them native to North America, 30 in Europe and the Mediterranean, and 25 in the Far East. Many of the North American species were described in a long series of short papers by Hottes in 1951-1964, and a number of these are only known from the original description. The European species fall into fairly well-defined groups that have been dignified by some authors as separate genera or subgenera, but there are difficulties with applying most of these groupings to the world fauna, possibly in part due to the inadequate descriptions available for many species, and they are not used here.

Schizolachnus was for a long time treated as a full genus, but recent molecular phylogenetic studies combining both nuclear and mitochondrial data indicate that it should be regarded as a subgenus nested within Cinara (R. Chen et al. 2016, Havelka et al. 2020), with distinctive features that may be associated with its pine needle-feeding microhabitst.

All Cinara live without host alternation on the roots, trunk, branches, twigs, shoots or foliage of their hosts. They are often regularly attended by ants, and have specialised parasitoids, Pauesia spp. (Smith 1944, as Protaphidius; Starý 1976). Different Cinara species are specialised for different feeding sites, and Favret & Voegtlin (2004a) showed by studying DNA phylogenies that feeding site specificity has played a greater part than host specificity in the speciation of Cinara on North American pinyon pines. A similar conclusion was reached by Durak et al. (2014) studying the DNA phylogeny of Cupressobium species. More extensive phylogenetic studies of European and North American species (Jousselin et al. 2013, Meseguer et al. 2015) provided evidence that geographic isolation has been a major factor driving speciation in this genus. Males may be apterous or alate depending on species, and oviparae often differ from viviparae in the presence of a perianal ring of wax. The male genitalia of six species were compared by Wieczorek et al. (2012). R. Chen et al. (2012) compared the use of three mitochondrial DNA regions as barcodes for identification of Cinara species.

For an introduction to the genus see Eastop (1972). Regional accounts are available for Britain (Eastop 1972, Carter & Maslen 1982, Blackman et al. 2019b); Fennoscandia and Denmark (Heie 1995); central Europe (Pintera 1966, Szelegiewicz 1962a); Germany (F.P. Müller 1969, Heinze 1962); Lithuania (Havelka et al. 2020); Austria (Fossel 1970); Switzerland (Lampel & Meier 2003); Italy (Binazzi 1978); the Iberian peninsula (Nieto Nafría et al. 2002a); Pakistan (Remaudière & Binazzi 2003 a,b); India (A.K. Ghosh 1982b); India, Bhutan and Nepal (Chakrabarti et al. 2020); east Siberia (Pashchenko 1988a); China (G. Zhang et al. 1993b); Korea (Paik 1972, Seo 1994, Lee et al. 1994)); Japan (Inouye 1970, Eastop et al. 1998); New Zealand (Baker 1986); eastern Canada (Bradley 1951); eastern USA Pinus-feeders (Pepper & Tissot 1973); USA pinyon pine feeders (Favret & Voegtlin 2004b) and USA Rocky Mountain Region (Palmer 1952). Danielsson & Carter (1993) keyed the subgenus Cupressobium, and Zhuravlev (2003) reviewed and keyed the Cupressobium of eastern Europe, Caucasus and Central Asia, treating it as a full genus. Voegtlin & Bridges (1988) provide a catalogue and bibliography of the North American species. Mamontova (2008) discussed the morphology, evolution and phylogenetics of the genus. For illustrations of and more detailed biological information on European species see Binazzi & Scheurer (2009), and http://www.influentialpoints.com/Gallery/Cinara_aphids.htm.

Cinara abietihabitans Zhang & Zhong BL of aptera c. 6.0 mm; appearance in life is not recorded. Described from Abies sp. in Yunnan, China (Zhang & Zhong 1985e). Related to C. confinis but with longer antennae, shorter hind tibial hairs and a longer R IV.

Cinara abietisibiricae Binazzi & Battisti Apterae are very large, uniformly dull bronze with diverging rows of small intersegmental round spots, prominent dark siphuncular cones and blackish legs; BL 4.8-6.4 mm. On bark of needle-bearing twigs of Abies sibirica in central Siberia (Binazzi & Battisti 1998).

Cinara abietispindrow Remaudière & Binazzi Apterae (only fundatrices and oviparae are described) are very large, brown; BL of fundatrices 6.7-7.7 mm, of oviparae 5.3-6.0 mm. On Abies pindrow in Pakistan (Remaudière & Binazzi 2003a). The original description also includes the alate vivipara. This species is close to C. confinis but could not be included in the key to Abies-feeders as the apterous vivipara is undescribed. The fundatrix differs from that of C. confinis by lacking spinal sclerites on abdominal tergites 1-7 and having cylindrical dorsal hairs with blunt, rounded apices. Oviparae were collected in early November.

Cinara acadiana Hottes Appearance in life is unknown: BL of aptera 2.9-3.2 mm. Described from Picea glauca in New Brunswick, Canada (Hottes 1956), and not found since.

Cinara acutirostris Hille Ris Lambers Apterae and alatae are dark brown to pale bronze, with a pattern of dark markings and wax dust similar to C. pini (see influentialpoints.com/Gallery); BL of aptera 2.6-4.1 mm. On twigs of Pinus nigra and P. pinea in western, southern and central Europe (England, Wales, France, Netherlands, Bulgaria, Czechoslovakia, Italy, Spain, Portugal), Turkey, Kazakhstan (Kadyrbekov 2017a); also reported from P. yunnanensis in China (G. Zhang et al. 1993b), and it has more recently been introduced into Argentina, where it is reported from P. patula and P. taeda (Delfino & Binazzi 2002). Alatae are produced in May-June (Italy) or June-July (England). Apterous males and oviparae occur in October-November, and eggs are laid on the needles (Binazzi 1973, Eastop 1972).

Cinara alacra Hottes & Essig Colour in life is not known; BL of aptera 4.6 mm. Described from a single specimen on Abies concolor in Nevada, USA (Hottes & Essig 1953). There is also a record from California (Smith & Parron 1978).

Cinara anelia Favret & Voegtlin Apterae are dark brown to dark grey or black, mottled with a more-or-less extensive pattern of grey wax; BL 2.8-4.6 mm. In small to large colonies on needle-free or sparsely needled branches, twigs or trunks of young Pinus monophylla, sometimes in mixed colonies with C. edulis or C. terminalis, frequently attended by ants. It may also sometimes occur on P. lambertiana (Jousselin et al. 2013). This species was previously confused with C wahtolca Hottes, but DNA analysis has demonstrated that this aphid on P. monophylla with almost identical morphology should be treated as a distinct species (Favret & Voegtlin 2004b). Alatae were collected in June. In western USA (California, Utah, Nevada).

Cinara antennalis Remaudière & Binazzi Colour of apterae in life is unknown; BL 3.3-4.9 mm. On Abies pindrow in Pakistan (Remaudière & Binazzi 2003a). The original description includes an alata collected in late June, and oviparae collected in September.

Cinara anzai Hottes & Essig (fig. 74B) Apterae are yellowish-brown, varying in darkness in different individuals, with interrupted white powdery dorsal abdominal cross-bands giving a whitish or brownish mottled appearance. Siphuncular cones dark each surrounded by a pale area, appendages light brownish-yellow with distal parts of femora, tibiae and tarsi dark; BL 3.5-3.9 mm. On branches of Pinus albicaulis in California and Oregon, USA. Oviparae and alate males were collected in September (Hottes 1956e).

Cinara apini (Gillette & Palmer) Apterae are dark brown mottled with wax powder on spinal and marginal areas and intersegmentally, with appendages mainly yellowish to dusky, siphunculi and distal two-thirds of tibia black (see influentialpoints.com/Gallery); BL 3.2-4.0 mm. In large colonies on bark of twigs and small branches of Pinus flexilis (Palmer 1952), and also sometimes on P. albicaulis, in western North America. (The record of this species from P. banksiana in Quebec (Quednau 1966) is a misidentification of C. banksiana.) Oviparae and alate males occur in September-October, and eggs with a light covering of wax-wool are laid on the needles.

Cinara arizonica (Wilson) Apterae dark brown to greyish-black with patches of wax powder on thorax and sides of abdomen, the largest ones being just anterior to siphunculi and tending to extend across abdominal tergite 4 ((Palmer 1952, Voegtlin 1976); BL 4.5-6.0 mm. On bark of stems of P. ponderosa, and there are also arboretum records from several other pines in subsections Ponderosae, Sabinianae and Australes. In western North America.

Cinara (Cupressobium) asishghoshi Chakrabarti, Medda & Kanturski Apterae are brownish, with immatures greenish; BL 3.2-3.9 mm. On apical twigs of Juniperus squamata in India (Uttarakhand). Differences from other Cupressobium species are tabulated by Chakrabarti et al. (2020).

Cinara atlantica (Wilson) (fig. 80E) Apterae are greyish brown with dark brown head, pro- and mesothorax, and paired dark brown irregular dorsal patches on metathorax and abdominal tergite 1, extensive more-or-less circular dark brown siphuncular sclerites, and legs dark except for a pale section on basal part of each tibia; BL 2.2-3.3 mm (Pepper & Tissot 1973). Forming dense colonies in spring on new growth, or on bark of small branches among the needle bases, of many Pinus spp., mostly in subsection Australes. In USA (especially common in the south-east) and Canada (Ontario), Cuba (Hernandez & Rodriguez 1985), Costa Rica (Villalobos Muller et al. 2010) and Jamaica (BMNH collection). C. atlantica has more recently been found in southern Brazil, causing significant damage to P. elliotti, P. taeda and P. caribaea (Lazzari & de Carvalho 2000), and Argentina (Delfino & Binazzi 2005), and has been introduced to Korea (Seo 1994). Oviparae and alate males occur in October, and eggs are laid on the needles. Anholocycly occurs in warmer regions. Oliver & Chapin (1988) summarise biology and control of C. atlantica in P. virginiana plantations in Louisiana. Patti & Fox (1981) studied seasonal trends, and Carner et al. (1977) record fungal attack, in populations on P. taeda in South Carolina. 2n=10.

Cinara atra (Gillette & Palmer) (fig. 74C) Apterae are rather long-bodied with narrow head and prothorax, shining black or dark brown without any wax, the appearance being superficially ant-like; BL 2.0-3.0 mm. Immatures are orange-yellow to olive-brown or reddish. In small colonies on smooth bark of small branches of young pinyon pines, Pinus edulis and P. monophylla, in Colorado and Utah, USA. Favret & Voegtlin (2004b) redescribed this species and made C. poketa Hottes a synonym. Apterous males and oviparae occur in October-November (Palmer 1952).

Cinara atripes Hottes Colour of apterae in life is not recorded, probably dark brown or black., with wholly dark legs; BL 1.6-2.3 mm (Hottes 1958a). On Picea glauca in Maine, USA. Biology and other morphs are unknown, and apparently not collected since the original description.

Cinara atroalbipes David, Narayanan & Rajasingh Apterae are green to dark green with a longitudinal spinal pale streak, appendages mainly pale brown except for the wholly black fore-tibiae; BL 2.7-3.2 mm. Originally described from “Pine” (David et al. 1971c) in Himachal Pradesh, India, but the host plant of apterous specimens from the type locality in the BMNH collection (leg. A.N. Chowdhuri) was identified as Picea smithiana, and this is probably the true host. Biology and sexual morphs unknown.

Cinara atrotibialis David & Rajasingh Apterae are brown with darker antennae and legs, the tibiae and tarsi almost black; BL 3.3-4.3 mm. On young shoots at needle bases of Pinus kesiya (= insularis), and also recorded from P. roxburghii (A.K. Ghosh 1982) and P. leboqi. In India (Himachal Pradesh, Meghalaya), and probably as geographic variants in Thailand (as C. khasyae Robinson), China (G. Zhang et al. 1993b) and the Philippines (Eastop 1976). Agarwala (1990) studied population trends in the field, Samonta & Raychaudhuri (1984) reported the effects of a parasitoid (Pauesia laricis) and Agarwala (1988) studied development in the laboratory. Sexuales and life cycle are unknown; active stages are present in December, so it is presumably at least partially anholocyclic. 2n=10 (Kurl 1988).

Cinara azteca Hottes Appearance in life is unknown; BL of aptera 2.0-2.3 mm (Hottes & Essig 1953). On Pinus ?oocarpa in Mexico. Not recorded since the original description; possibly these were small specimens of C. atlantica.

Cinara balachowskyi Remaudiere Apterae are rather elongate oval with narrow head and thorax, chocolate brown with uniformly dark legs and a light dusting of white wax powder on the ventral abdomen (Remaudiere 1974, Melia Masia 1978); BL 3.3-4.9 mm. In compact colonies in bark crevices on the trunk of Pinus halepensis in the western Mediterranean (France, Spain). Life cycle and sexuales are unknown; active stages have been collected in December, suggesting anholocycly.

Cinara banksiana Pepper & Tissot (fig. 75E) Apterae have a dark head and thorax and paler brown abdomen with paired dark markings and large, prominent shiny black siphuncular cones; BL 2.4-3.6 mm. On previous year’s growth or in dense colonies on new shoots, producing numerous alatae in spring and early summer; on Pinus banksiana, and apparently specific to this host, in Canada and northern USA. Oviparae and alate males occur in October, and eggs are laid on the needles. Bradley & Hinks (1968) studied the relationship with ants in Manitoba.

Cinara bhutanica Chakrabarti, Medda & Kanturski Apterae have a dark brown head and thorax and brownish abdomen with paired black spinal and marginal patches and variably developed white wax longitudinal stripes; BL 3.9-4.7 mm. Feeding at bases of twigs of Pinus wallichiana and Pinus sp(p). in Bhutan. Apterae and alatae are described by Chakrabarti et al. 2020), who tabulate characters distinguishing it from two similar species (eastopi and tenuipes).

Cinara bonica Hottes Colour in life is not recorded, probably brownish with some wax dust; BL 3.4-3.6 mm. Collected in Alaska on an unknown host, probably either Picea glauca or P. mariana, and not found since (Hottes 1956g).

Cinara bonita Hottes Colour in life is not recorded, probably brownish with some wax dust; BL about 5.7 mm. Collected in Alaska on an unknown host, probably either Picea glauca or P. mariana, and not found since (Hottes 1956g, 1964).

Cinara braggii Gillette Apterae in life are covered in white wax powder, with legs pale; BL 3.2-3.8 mm. Feeding solitarily on bark of twigs, and on new growth, of Picea spp., with a wide distribution in North America. Numerous alatae are produced in the second and third generations. Oviparae and alate males were collected in October (Gillette 1917). 2n=10.

Cinara brauni Börner (fig. 78) Apterae are golden brown with a dusting of wax powder, except for an extensive shiny dark brown to black patch on abdominal tergites 5-7 encompassing the siphunculi (see influentialpoints.com/Gallery), although this patch is reduced in some populations (Binazzi 1978); BL 2.7-3.8 mm. On current year’s growth and one-year-old twigs of Pinus spp. of the nigra group, attended by ants (usually Formica rufa), in Europe and east to Turkey and Crimea. Oviparae and alate males in October-November. Scheurer (1971b) described the fundatrix, alata and sexuales. Binazzi & Roversi (1987) studied seasonal differences in abdominal sclerotisation, including holocyclic and anholocyclic populations.

Cinara brevipilosa Voegtlin, Remaudiere & Pena Martinez Colour of apterae in life is unknown, probably dark, with an extensive black dorsal abdominal sclerite that does not however encompass the small siphuncular cones; BL 2.0-3.5 mm. A species with very short dorsal abdominal hairs, collected from an unidentified Pinus sp. in Mexico (Voegtlin et al. 1986). Biology is unknown.

Cinara brevisaeta Zhang & Zhang BL of aptera c. 3 mm. On Pinus armandii in Yunnan, China (G. Zhang et al. 1993b).

Cinara brevispinosa (Gillette & Palmer) (fig. 75C) Apterae are brown lightly dusted with wax powder medially and marginally; BL 3.0-3.5 mm. In colonies on bark of twigs and small branches of Pinus contorta (incl. var. murrayana) in western North America. Oviparae and alate males occur in late August-October (Gillette & Palmer 1931). C. sclerosa Richards is a synonym.

Cinara bungeanae Zhang & Zhang BL of aptera c. 3 mm. On Pinus bungeana in Beijing, China. Sexual morphs were collected in September (G. Zhang et al. 1993b).

Cinara burrilli (Wilson) Apterae are, according to the original description, black “with pruinose patches that produce a calico effect” (but see Hottes 1951d); BL 2.5-3.0 mm. On bark of twigs of Juniperus spp. in the Rocky Mountain region, USA (Palmer 1952). Biology and sexuales are unknown. C. pulverulens may be a synonym.

Cinara caliente Hottes Apterae are dusky grey, variably dusted with wax powder, with black sclerotisation of thorax and abdominal tergite 1 often in form of an inverted “V” (Hottes 1955f); BL 1.6-2.2 mm. On bark of twigs, especially on needle-free sections, of Pinus edulis and P. monophylla in western USA (Hottes 1960b). Favret & Voegtlin (2004b) provided a redescription. Oviparae and alate males were collected in September.

Cinara californica Hottes & Essig Appearance in life is unknown, in cleared and mounted specimens all tibiae are uniformly dark (Hottes & Essig 1953); BL 3.1-3.5 mm. This species is something of a mystery, being described from the Chinese pine P. tabuliformis in California. Long-haired Cinara on other Pinus spp. from California in the BMNH collection, including H.G. Walker’s material previously identified as californica (Walker et al. 1978), have two-toned tibiae and short tarsi and seem to be the East Coast species atlantica (q.v.). C. californica is included in the key to species from oriental Pinus, in case it is a previously unrecognised oriental species introduced into California. This seems to be now confirmed by a record of C. californica from north-west China (G. Zhang 1999), although with the host identified as P. sylvestris.

Cinara canatra Hottes & Bradley (fig. 75A) Apterae are shining black, without wax; BL 2.3-3.5 mm. In spring in dense colonies on bark of smaller branches some distance from the tips, later moving to form clusters on the bark of the trunk (Hottes & Bradley 1953). Probably specific to Pinus banksiana, where it may occur in mixed colonies with, and be confused with, C. nigra (Pepper & Tissot 1973); records from other hosts are referable to C. gracilis or C, nigra. In Michigan, USA, and Ontario, Manitoba and Newfoundland, Canada. Oviparae were collected in October, males are unrecorded.

Cinara carnica Binazzi Apterae are light bronze to dark brown with a slight dusting of wax; BL 3.5-4.2 mm. On shoots and 1- to 3-year-old twigs of Pinus mugo in mountains of north-east Italy, in small colonies and sometimes mixed with C. neubergi (Binazzi 1995). Oviparae were collected in August and October, an an alate male in late August.

Cinara caudelli (Wilson) Appearance in life is unknown, and not found since original colection, on an unidentified Picea sp. in British Columbia (Wilson 1919). BL of aptera about 2.4 mm. Life cycle and sexuales are unrecorded.

Cinara cedri Mimeur Apterae are dark bronze or reddish-brown with darker segmental markings and a distinctive but variable pattern of white wax, usually comprising a spinal stripe and paired dorsolateral segmental patches, as well as a ventral dusting of wax powder (see Plant Parasites of Europe website and influentialpoints.com/Gallery); BL 2.2-3.8 mm. In compact colonies on branches of Cedrus spp.. It is widely distributed in Europe, and in North Africa (Morocco, Algeria), the Mediterranean area including Cyprus (F. Binazzi et al. 2015), south-west and Central Asia, North America (California, New York), and Argentina (Delfino & Binazzi 2002). Oviparae and alate males occur in October-November, sometimes concealed under growths of lichen on the branches (Carter & Maslen 1982). See Covassi & Binazzi (1974), Stroyan (1979), Notario et al. (1984) and F. Binazzi et al. (2015) for fuller accounts. Populations on Cedrus libani ssp. brevifolia in Cyprus are now regarded as an endemic subspecies, C. cedri ssp. brevifoliae A. Binazzi (F. Binazzi et al. 2017), alatae of which were described by F. Binazzi & Roversi (2018). The endosymbionts of C. cedri (Buchnera and Serratia) have been the subject of recent research as they are at an interesting stage of co-evolution (Lamelas et al. 2011). A draft assembly of the genome of C. cedri has been published and compared phylogenetically with those of four species of Aphidinae and one of Chaitophorinae (Julca et al. 2019). 2n=10.

Cinara cembrae (Seitner) Apterae are shiny brown, BL 3.3-4.8 mm. On bark of two-year-old or older parts of branches of Pinus cembra in mountainous areas of central Europe (Austria, Czech Republic, Switzerland), and in Russia (Leningrad district), Ukraine and Kazakhstan (Kadyrbekov 2017a). Oviparae and both apterous and alate males are produced in late August (Pintera 1966). Records of cembrae from east Asia should probably all be referred to C. mongolica. 2n=10.

Cinara chibi Inouye Apterae have a brown head and thorax and shiny dark brown to black abdomen, with a pale brown spinal stripe (Inouye 1962); BL 2.4-3.1 mm. On stems of young trees, or on 3- to 4-year-old parts of branches, of Larix spp. in Japan. Oviparae and alate males occur in October (Inouye 1970). It was placed as a subspecies of C. lariciola by Inouye (1970) and could be small specimens of that species (which might itself be a synonym of C. cuneomaculata).

Cinara chinookiana Hottes Apterae are shiny apple-green to brownish green with brown-black head and dark markings on thorax and posterior abdomen; BL 3.3-3.5 mm. On Abies lasiocarpa and A. amabilis (BMNH collection), living singly on upper sides of small twigs near their tips (Bradley 1961). In western North America. Apterous males and oviparae occur in October (Hottes 1957).

Cinara cognita Hottes & Essig Appearance in life is unknown: BL of aptera 3.6 mm (one specimen), BL of alatae 2.4-3.3 mm (Hottes 1955d). On Juniperus communis in western USA (California, Washington). Biology and sexuales are unknown.

Cinara coloradensis (Gillette) (= caliginosa Hottes) Apterae have the abdomen dullish black dorsally, somewhat shiny black ventrally, and a reddish-brown head and thorax (Gillette 1917); BL 2.2-3.4 mm. In large, dense colonies on undersides of small 2 or 3-year-old branches, and on upper trunk (Bradley 1961), of Picea spp. Widely distributed in western USA and across Canada. 2n=10.

Cinara comata Doncaster Apterae are pale brown; BL 4.1-4.5 mm. Alatae have the media only once-branched. On Picea smithiana in north-west India (A.K. Ghosh 1982) and Pakistan (Remaudière & Binazzi 2003a). Biology and sexual morphs are unknown.

Cinara commatula Hottes & Essig Appearance in life is not recorded; BL of aptera 4.3-4.7 mm. One aptera and one alata were described, from Pseudotsuga taxifolia in California, USA (Hottes & Essig 1954b).

Cinara confinis (Koch) (= abieticola Cholodkovsky; = C. indica Verma) (fig. 11B) Apterae and alatae are large, dark brown or greenish-black with a double row of blackish slightly shining spots around a paler central area, and small flecks of wax powder in transverse rows (see influentialpoints.com/Gallery); BL 3.8-7.8 mm. Usually in spring colonies on trunk, branches and twigs of Abies spp., and in summer on the roots; sometimes also on Cedrus spp., where it may build up large spring colonies on trees growing in urban areas. Recorded from Europe, south-west and Central Asia, India, Pakistan, Siberia (Pashchenko 1988a, as piceae), China (G. Zhang et al. 1993b), Canada, western USA and Argentina (Blanchard 1939; as grossa). Struble et al. (1976) studied its biology in Maine, USA. DNA barcoding data indicate that North American and European populations of C. confinis have been separated for a long time (Skvarla et al. 2017). Oviparae and alate males are recorded in September-October in northern Europe, Pakistan, India and North America, but are not recorded from other countries where the species is common, suggesting that anholocycly occurs in mild winter climates (see also Eastop 1972). Heikinheimo (1963) records natural enemies in Finland. C. tenuipes, described as a subspecies of confinis, is here regarded as a distinct species. 2n=12*.

Cinara contortae Hottes (fig. 75D) Colour in life is not recorded; BL 2.5-3.2 mm. On Pinus contorta in California, USA (Hottes 1958e). Biology and sexuales unknown. This species is keyed on the basis of specimens in the BMNH collection (leg. D.J. Voegtlin) which, unlike the original description, have well-developed anterior dorsal abdominal sclerites, and in this respect resemble C. medispinosa, but have much shorter dorsal abdominal hairs. This species is also very close to C. brevispinosa and C. banksiana; taxonomic relationships in this group of species require investigation.

Cinara costata (Zetterstedt) Apterae are wax-covered, light brown or yellow-brown, sometimes with a dull metallic golden sheen, and with a pair of dark bottle-green dorsal longitudinal stripes which sometimes coalesce at about the level of the siphunculi (see influentialpoints.com/Gallery); BL 2.0-4.0 mm. Alatae have characteristically pigmented forewings with the media only once-branched. In small colonies on smaller woody twigs (which receive a deposit of mealy wax) on lower branches of Picea spp., usually not attended by ants (Pintera 1966; but see Eastop 1972). It occurs in Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), across Asia, Australia, Greenland, North and South America. Oviparae and alate males occur in the northern hemisphere from late August to mid-November. 2n=10.

Cinara covassii Binazzi Apterae are shimmering light bronze; BL 4.1-5.0 mm. On older branches of Pinus mugo in the eastern Alps, northern Italy. Binazzi & Covassi (1994) compared it with other species on P. mugo in Italy. Ovipara occur in August (Binazzi 1991a). The alate vivipara (Binazzi 1992) and the alate male (Binazzi 1994b) were subsequently described.

Cinara cronartii Tissot & Pepper Apterae are dark brown with mainly dark legs; BL 2.6-4.2 mm. This aphid seems normally to restrict its feeding site to lesions and cankers of the rust Cronartium fusiforme on trunks and branches of Pinus spp. (taeda, elliotti, clausa, serotina)in eastern USA (Pepper & Tissot 1973). Introduced into South Africa, where severe infestations cause economic damage to P. taeda (Raubenheimer & Shaw 1987). Biology was studied in the laboratory in South Africa by van Rensburg (1981), and biocontrol with a parasitoid (Pauesia sp.) has been attempted (Kfir et al. 1985, Kfir & Kirsten 1991). Sexuales are unrecorded; in Florida the aphids remain in rust lesions throughout the year. 2n=10*.

Cinara cuneomaculata (Del Guercio) (= boerneri) Apterae and alate are of variable colour, darkish brown to orange-reddish, sometimes with dark green segmental markings, and with a ventral dusting of greyish wax powder that sometimes extends dorsally (see influentialpoints.com/Gallery); BL 2.4-4.6 mm. In small colonies on young twigs and shoots of Larix spp., usually but not always ant-attended, in Europe, Central Asia (Kazakhstan; Kadyrbekov 2017a), Mongolia (see Eastop 1972; as boerneri), west and east Siberia (Pashchenko 1988a; Stekolshchikov & Khureva 2020) and China (G. Zhang et al. 1993b). Oviparae in October-November, with alate males having been recorded as early as June. It is virtually indistinguishable from C. laricicola Matsumura in Japan (Inouye 1962) and Korea (Paik 1972, Seo 1994), which may be a synonym. 2n=10.

Cinara (Cupressobium) cupressi (Buckton) Plate 16b Apterae and alatae are orange-brown to yellowish-brown, the dorsum dusted with pale grey wax making a pattern of cross-bands, with two dark wavy longitudinal stripes, close together on the thorax and diverging on the abdomen, where they merge with a broad brown-black cross-band at the level of the siphunculi (see influentialpoints.com/Gallery); BL 1.8-3.9 mm. Most commonly on Cupressus spp. but also on other Cupressaceae, feeding on smaller twigs in the foliated parts of the crown, and frequently causing branch die-back (Inserra et al. 1979). It is a vector of Seiridium cardinale (Sutton & Gibson 1972). In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), south-west and central Asia, India (Agarwala & Raychaudhuri 1982), Pakistan (Remaudière & Binazzi 2003b), China (G. Zhang et al. 1993b), North America, and also introduced into Africa south of the Sahara (Mills 1990) and South America (Brazil, Sousa-Silva & Ilharco 2001a; Columbia, BMNH collection, leg. A. Lopez; Argentina, Delfino & Binazzi 2005). A population on Juniperus chinensis in north-west China was described as a subspecies, C. cupressi ssp. langzhouensis Zhang, Chen, Zhong & Li (G. Zhang 1999). Watson et al. (1999) used multivariate analysis to distinguish populations introduced from the Middle East to Africa as a separate species, C. cupressivora Watson & Voegtlin, but Remaudière & Binazzi (2003b) found morphologically intermediate populations in Pakistan and considered the distinction from C. cupressi to be unjustified. DNA analysis is badly needed to clarify the situation. Populations identified as C. cupressivora are a major pest in plantations of Cupressus lusitanica in Malawi, and biological control was attempted in the 1990s using the parasitoid Pauesia juniperorum (Kairo & Murphy 2005; Missanjo & Kamanga-Thole 2015). Ruiz et al. (2014) analysed morphometric variation of Chilean populations of C. cupressi. Gunkel (1963 a,b) studied biology, ecology and natural enemies (including an aphidiine parasitoid) in Germany, Durak et al. (2007) studied its ecology on Thuja occidentalis in Poland, and Middle Eastern populations were studied in Jordan (Mustafa 1987) and Israel (Mendel & Golan 1983). Oviparae and alate males occur in September-October in Europe, but anholocycly is prevalent in regions with mild winters. Durak & Durak (2015) provided a redescription of the male. Ciesla (1992) provided an account with special reference to Africa. Some of the literature records were based on misidentified C. fresai or C. tujafilina. 2n=12.

Cinara curtihirsuta Hottes & Essig Apterae have an orange-red head and thorax and shiny black abdomen (Voegtlin 1976); BL 3.0-3.9 mm. On Abies concolor in California and Oregon, USA, on branches, or on trunks of young trees. Oviparae occur in September-December (Voegtlin 1976), males unrecorded.

Cinara curvipes (Patch) (fig. 11A) Apterae are shiny or dull grey-black (see influentialpoints.com/Gallery), in early summer sometimes (perhaps in the absence of ants) developing a covering of pale grey wax; BL 3.4-5.7 mm. Early instars may have an extensive covering of wax. On trunk or branches of Abies spp., and also sometimes found feeding on Cedrus and Picea. Voegtlin (1976) found a small colony on Pinus contorta. Widely distributed in USA and Canada, also in Mexico (Ortiz 1982), and now also widely distributed in Europe (UK, Martin 2000, Baker 2009b; Germany, Scheurer & Binazzi 2004; Switzerland, Angst et al. 2007; Austria, Perny 2014; Hungary, Bodor 2013; Serbia, Poljaković-Pajnik & Petrović-Obradović 2002; Czech Republic, Olbrechtová 2007; Slovenia, Jurc et al. 2009; Slovakia, Nakládal 2007; Turkey, Görür et al. 2015; Poland, Halaj & Osiadacz 2015). Oviparae and alate males occur in September-October, but it can also be anholocyclic in mild winters, building up large colonies in spring (Scheurer & Binazzi 2004, Scheurer et al. 2004).

Cinara (Schizolachnus) curvispinosa (Hottes, Essig & Knowlton) Colour of aptera in life is not recorded, ‘probably a dusky green’, with black hind tibiae; BL 2.1-2.8 mm. On needles of Pinus ponderosa, and occasionally found on non-native pines, in western North America. Oviparae and alate males occur in October (Hottes et al. 1954). The alata was described by Hottes (1956d).

Cinara (Cupressobium) dahurica Szelegiewicz & Holman Apterae are shining brownish-yellow, without wax: BL 3.9-4.3 mm. Collected from twigs of Juniperus pseudosabina in Mongolia (Szelegiewicz & Holman 1980). Biology and other morphs are unknown.

Cinara deodarae Seo Apterae have BL c. 2.2-2.3 mm, alatae have BL 2.5-3.3 mm. Described from apterae and alatae on Cedrus deodara in Korea (Seo 1994). Biology and other morphs are unknown.

Cinara diabola Hottes Appearance in life is unknown, probably dark-coloured: BL about 3.4 mm. Described from apterae and alatae collected on Pinus coulteri in California (Hottes 1961b), and not since recorded.

Cinara dubia Hottes & Essig Appearance in life is unknown, probably dark-coloured; BL 3.0-3.5 mm (Voegtlin 1976). Only the aptera is described (Hottes & Essig 1954), from Pseudotsuga taxifolia in Oregon, USA. Also recorded from California (Voegtlin & Bridges 1988). Probably a synonym of C. pseudotaxifoliae (q.v.).

Cinara eastopi Pintera (fig. 80A) Apterae are brownish, with a series of dark spinal sclerites; BL 4.0-5.0 mm. On Pinus wallichiana, “under bark” (BMNH collection, leg. S.N. Chatterjee), in north-west India, Bhutan, and recorded as sometimes very abundant on this host in the Muree Hills of Pakistan, so that honeydew which had crystallised as large lumps on the branches could be collected (Naumann-Etienne & Remaudière 1995). Biology and sexuales are unknown. Vagrant alatae have been found on several different plants, and on snow, suggesting that this species may be common but often overlooked on pine. Differences from two very similar species (tenuipes and bhutanica) are tabulated by Chakrabarti et al. (2020).

Cinara edulis (Wilson) (fig. 74D, as C. pinona) Apterae are cinnamon brown to dark brown to gun-metal or black, often with a pale narrow spinal stripe and paired patches of whitish wax powder on thorax and abdomen, and legs bicoloured black and orange; BL 2.5-4.0 mm. On Pinus edulis and P. monophylla in western USA, in large colonies on bark of branches and older twigs, or on young tree-trunks, or feeding among needles on the younger branches of young healthy trees with greenish bark. Oviparae and apterous and/or alate males occur in September-October, eggs being laid along the needles (Palmer 1926). Using multivariate morphometric analysus, Favret & Voegtlin (2004b) demonstrated that five Hottes species (apacheca, metalica, pinata, pinona and rustica), treated separately in Blackman & Eastop (1994), are probably synonyms of C. edulis.

Cinara engelmanniensis Gillette & Palmer Apterae are “light cinnamon-brown to tan or pinkish-buff, with a pair of longitudinal rows of green or black spots” (Gillette & Palmer 1925); BL 2.2-2.7 mm. Scattered or in small groups on bark of branches or twigs of Picea engelmannii, usually close to the ground (Hottes 1961f). In western USA, and now also recorded from eastern USA on P. rubens (Favret et al. 2010). Oviparae and alate males occur in September-October.

Cinara essigi Hottes Appearance in life is unknown; BL of aptera c.4.5 mm. Described from Pinus attenuata in California, USA (Hottes 1961a). Biology and sexuales are unknown. Apart from its larger size this aphid is, to judge from its description, closely related to and possibly synonymous with either C. cronartii or C. newelli.

Cinara etsuhoe Inouye Apterae elongate oval, dark brown; BL 4.5-5.8 mm. Collected on bark of trunk of Pinus densiflora in Japan, near the ground and often in ant shelters (Inouye 1970). Life cycle and sexuales are unknown. It is very similar to and perhaps conspecific with C. sorini (see Eastop et al. 1998).

Cinara ferrisi (Swain) Apterae are rather elongate oval, brown mottled with black, slightly pruinose; BL 4.3-6.5 mm. On Pinus albicaulis and P. monticola in western USA and British Columbia. The only recorded feeding site is the roots (Bradley 1961). Life cycle and sexuales are unknown. Records from Abies concolor should probably be referred to other species, eg. C. curvipes. [A record from Tunisia (Ben Halima-Karmel 2012) needs further confirmation.]

Cinara flexilis Gillette & Palmer Apterae brownish-black, dull or shining, without wax; BL 4.0-5.0 mm. Scattered on bark of twigs and small branches of Pinus flexilis and P. strobiformis in Arizona (BMNH collection; leg. F.C. Hottes), Colorado and New Mexico, USA. Oviparae were collected in Colorado in September (Palmer 1952).

Cinara (Schizolachnus) flocculosa (Williams) Apterae have a black head and thorax with grey wax markings, abdomen light brown or green with narrow cross-bands of wax (Hottes 1952a; as Schizolachnus wahlea) and black hind legs; BL 2.4-2.9mm. On needles of Pinus arizonica and P. ponderosa in US Rocky Mountain region (Colorado, Utah) and Arizona; there are now also records from the Great Smoky Mountains in eastern USA, on P. rigida and P. virginiana (Favret et al. 2010). Oviparae and alate males in September-November (Hottes 1954b; as S. wahlea).

Cinara formosana (Takahashi) Apterae are broadly oval, yellowish-brown with dusky yellow legs mottled with dark brown; BL 4.0-5.5 mm. Immatures are somewhat greenish (Takahashi 1924a). On bark of 1- or 2-year-old shaded branches (Inouye 1970) of Pinus spp. (especially massoniana, thunbergii) in east Asia (Japan, Korea, Thailand, China, Taiwan). Oviparae and alate males in November (BMNH collection). C. pinitabulaeformis Zhang & Zhong 1989 is a synonym. Wu et al. (2017) have sequenced the transcriptome of this species (as C. pinitabulaeformis). 2n=10.

Cinara fornacula Hottes Apterae are pale, translucent green, or sometimes pale brown or orange, with slight pruinosity (see influentialpoints.com/Gallery); BL 3.1-4.5 mm. On small branches and current year’s growth of Picea spp., living at the needle bases, not attended by ants (see Johnson & Lyon 1988, p.83). Widely distributed in North America. Oviparae and alate males were collected in late August, with eggs laid on the needles (Hottes 1933a). 2n=10.

Cinara (Cupressobium) fresai Blanchard Apterae are pinkish grey to dark brownishgrey or greenish grey, dusted with white wax, especially on sides of thorax, and with paired black patches on thoracic and anterior abdominal tergites diverging in an inverted “V” (see influentialpoints.com/Gallery); BL 2.2-4.2 mm. Alatae are unusual in having the radial sector not reaching the apex of the forewing. On foliage and adjacent woody shoots and branches of various Cupressaceae (Juniperus spp., Cupressus spp.; see Eastop 1972), and also recorded from Cryptomeria japonica (Taxodiaceae; Bradley 1965b, as C. maui). Known distribution is remarkably disjunct; England, Wales, Spain, Israel (Mendel & Zehavi 1981), USA, Central and South America, Australia, New Zealand, Japan (Eastop et al. 1998), and Korea (Seo 1994). C. fresai is evidently anholocyclic; no sexual morphs have been recorded, and the karyotype is structurally heterozygous (2n=13).

Cinara gentneri Hottes Apterae are light yellowish-brown to brown (Hottes 1957): BL 5.5-5.8 mm. On Abies grandis and A. concolor, only recorded from Oregon, USA. Oviparae in October. Alate viviparae and males, and biology, are not known.

Cinara glabra (Gillette & Palmer) (fig. 77) Apterae are dusky olive-green, olive-brown or black, without dorsal wax; BL 2.5-3.0 mm. Singly or in small groups on twigs at bases of needles of Pinus ponderosa in western USA; Colorado, Arizona, New Mexico and Wisconsin (BMNH collection; leg. F.C. Hottes). Oviparae and alate males occur in October (Gillette & Palmer 1924, 1931).

Cinara glacialis Hottes Appearance in life is not recorded; BL of aptera c.2.7 mm. Other morphs and life cycle unknown. Collected on Picea sitchensis in Alaska (Hottes 1964). Closely related to, if not synonymous with, C. glehna.

Cinara glehna (Essig) Apterae are reddish-brown to dusky brown, but in life covered with white wax (Essig 1915); BL 2.9-4.0 mm. On twigs of Picea spp., feeding on bark at bases of needles. First described heavily infesting the east Asian species P. glehna in California, but since reported mainly from P. pungens, which is presumably the native host. In western North America. Sexuales and life cycle unknown. Closely related to C. braggii, and possibly the two are variants of a single species. C. mariana, described from Picea mariana in Saskatchewan, Canada, and not found since, keys to C. glehna on the basis of the original description (Bradley 1956b). and the only clear difference seems to be its smaller size (BL c.2.0 mm). C. yukona Hottes, described from one alate vivipara on P. glauca in Alaska (Hottes 1964) is also very similar and may be a synonym.

Cinara gracilis (Wilson) Apterae are shiny black, sometimes with a whitish wax spinal stripe on thorax and abdominal tergites 1-2; BL 2.7-3.5 mm. It lives in dense colonies on bark among needles, often near the bases of twigs and small branches, and is easily disturbed when approached (Pepper & Tissot 1973). The normal host is Pinus virginiana, and it is also found on P. taeda (Patti & Fox 1981), but the more northerly records from P. banksiana (eg. Bradley & Hinks 1968) seem to be misidentifications of canatra and/or nigra. In eastern USA.

Cinara grande Hottes Appearance in life is unknown; BL of aptera 4.8-5.1 mm. On trunk of young Abies concolor in Arizona, USA, not collected since the original description (Hottes 1956), although the earlier records of Lachnus abietis on A. concolor in California by Davidson, referred to C. ferrisi by Swain (1918), possibly apply to this species. Other morphs and life cycle are unknown.

Cinara guadarramae Mimeur (fig. 80H) Apterae are dark brown, often with two dorsal longitudinal rows of small black sclerites; BL 2.3-3.7 mm. On young branches of Pinus sylvestris and P. pinaster in Spain. Very similar to C. pini; a detailed morphometric comparison is made by Garcia Sanchez & Nieto Nafria (1978). Sexuales and life cycle are not known.

Cinara gudaris Binazzi Apterae are greyish brown; BL 3.9-4.2 mm. Described from the Sierra de Gúdar, Spain, on the Spanish black pine Pinus nigra var. salzmannii, living among the needles of 1- or 2-year-old twigs (Binazzi & Sanchis 1994). Binazzi et al. (1995a) subsequently described the alata from material collected in northern Spain. Sexuales and life cycle are unknown.

Cinara harmonia Hottes Apterae are dark greenish-brown or brownish (Hottes 1958m), with broad but fairly low black siphuncular cones; BL 2.9-4.1 mm. Usually in colonies, feeding at needle bases on 2- to 3-year-old wood of Pinus resinosa, in eastern USA and Canada. Oviparae occur in October, males are unrecorded (Pepper & Tissot 1973).

Cinara hattorii Kono & Inouye Apterae are dark brown to black, rather elongate oval; BL 3.2-4.4 mm. Alatae have forewing membrane pigmented between Cu_1aandCu_1b. On Abies sachalinensis and A. mariesii in Japan, forming colonies on bark of trunk or old twigs 1 m or less above ground. Oviparae and alate males occur in October-November. C. konoi is a synonym (Inouye 1956), and its separation from longipennis needs further confirmation.

Cinara (Cupressobium) himalayaensis Chakrabarti, Medda & Kanturski Apterae (and also immatures) are brownish: BL 4.8-5.2 mm. On shoots of Taxus baccata in India (Uttarakhand). Morphological differences from other Cupressobium species are tabulated by Chakrabarti et al. (2020).

Cinara hirsuta Hottes & Essig Appearance in life is unknown; BL of aptera 2.9-3.3 mm. Described from apterae on Pinus monticola in Oregon, USA (Hottes & Essig 1954a), and there has been a subsequent collection from the same host in Idaho (aphidtrek.org). C. kuchea is possibly a synonym.

Cinara horii Inouye Apterae are rather elongate oval, with yellow head, light green thorax and green abdomen (Inouye 1956); BL 4.4-5.3 mm. Resembling C. pilicornis, but without a dark cross-band on abdominal tergite 8. On twigs and young shoots of Picea jezoensis in Japan and Sakhalin (Inouye 1970). Eggs and fundatrices have been observed in April-May, but sexuales are unrecorded.

Cinara hottesi (Gillette & Palmer) Apterae are dull bluish-black with mainly yellowish orange appendages (see influentialpoints.com/Gallery); BL 3.0-3.5 mm. In large dense colonies on bark of twigs of Picea spp., moving to larger branches or trunk as season progresses (Bradley 1961). Widespread in North America. Oviparae and apterous males in September-October in Colorado (Hottes 1955a). C. alaskana, described from Picea sp. in Alaska (Hottes 1964), is probably a synonym; its BL is recorded as 2.2 mm, but this is possibly a misprint.

Cinara hylikos Zhang & Zhang Appearance in life is unknown, apterae have paired dark sclerites on abdominal tergites 3-4 and 6-8 and a broad quadrate patch beween the siphunculi; BL c. 2.5 mm. On an unidentified Pinus sp. in Guangxi, China (G. Zhang et al. 1993b)

Cinara hyperophila (Koch) Aptera are dark brown to black, shining, with legs mainly or wholly dark, sometimes secreting abundant wax (Binazzi & Scheurer 2009); BL 2.7-3.6 mm. On 1- to 3-year-old shoots of Pinus sylvestris, and possibly sometimes on P. mugo, ant-attended, never in large colonies. In Scandinavia, central Europe, and also now recorded from Spain (Binazzi et al. 1995b). Heikinheimo (1984) redescribed the viviparous morphs (as diversiseta), and Danielsson (1987) discussed the synonymy. Biology is little known; Koch (1855) collected oviparae in September-October and found numerous males in April-May, perhaps due to exceptional conditions (see Eastop 1976). 2n=10.

Cinara inscripta Hottes & Essig Fundatrices have a dark brown head and thorax and shiny amber-brown abdomen with large prominent dark brown siphuncular cones (Bradley 1961), appearance of later generations in life unrecorded; BL 3.7-4.2 mm. On Pinus albicaulis in Oregon and Idaho, USA and British Columbia, Canada, colonising young branches in spring; otherwise biology unknown, and sexuales are undescribed.

Cinara intermedia (Pasek) Apterae are mainly pale brown, with two brownish or greenish-black dorsal longitudinal stripes, converging on abdominal tergites 5-8 (Heinze 1962); BL 2.5-2.7 mm. On bark of trunk of young Pinus sylvestris in Slovakia, as yet only collected (twice) as single apterae within colonies of C. nuda (Pintera 1966). A record from Turkey (Görür et al. 2017) requires further confirmation. Life cycle and other morphs are unknown.

Cinara jianglensis Zhang & Qiao Apterae are brown in life, BL c.2.6 mm. On Pinus massoniana in China (Fujian province). Other morphs and biology are unknown (Qiao & Zhang 1999b).

Cinara jucunda Hottes Appearance in life is unknown; BL of aptera 2.7-3.0 mm. Apterae described from either Picea pungens or P. glauca (or both; Hottes 1958k) in North Dakota USA, and also recorded from Colorado and S. Dakota (Voetglin & Bridges 1988). Other morphs and biology are unknown.

Cinara juniperensis (Gillette & Palmer) Apterous viviparae are undescribed (keyed according to oviparae); BL 3.5-4.0 mm. On bark of tender twigs of Juniperus communis in Colorado and Oregon, USA. Records from Turkey (Şenol et al. 2014b, Akyildirim Beğen & Görür 2019) require additional conformation (the latter publication reports on the mitochondrial COI sequence of an aphid collected on Juniperus that was identified as C. juniperensis, but was more likely to be C. juniperi). Oviparae were collected in late August (Gillette & Palmer 1931).

Cinara (Cupressobium) juniperi De Geer (= sibiricae Gillette & Palmer) Apterae are quite plump-bodied, pinkish-brown with variable wax dusting, not forming a pattern, and dark legs (see influentialpoints.com/Gallery); BL 2.1-3.4 mm. On undersides of young shoots of Juniperus communis and its varieties, feeding on the needles; occasionally on other Juniperus spp., but most records from other hosts are likely to be misidentifications of tujafilina, fresai or cupressi. In Europe (incl. Iceland), North Africa (Algeria; Laamari et al. 2013), the Middle East (Turkey, Lebanon), Central Asia (Kazakhstan; Kadyrbekov 2017a), Siberia, Mongolia, ?China (G. Zhang et al. 1993b), Japan (Eastop et al. 1998) Australia, New Zealand, western USA, and Canada (Ontario). Oviparae and apterous males occur in September-October in Colorado, USA (Palmer 1952, as sibiricae), elsewhere it is apparently anholocyclic (Eastop 1972). Alatae occur mainly in May-June. Durak (2011) explored differences in mitochondrial DNA and morphology between C. juniperi and C. mordvilkoi. 2n=12.

Cinara juniperivora (Wilson) Apterae are dull dusky to cinnamon or greyish-brown with a waxy bloom, with legs mainly dark; BL 2.2-3.0 mm. On Juniperus virginiana and J. silicicola in eastern USA, living under bark on trunk or branches, often in ant shelters of Crematogaster – see account by Hottes & Frison (1931; as C. difficilis). Sexuales and life cycle are unknown. Durak (2011) compared mitochondrial DNA sequences with those of C. juniperi and C. mordvilkoi.

Cinara keteleeriae Zhang Described on the basis of a single aptera (BL about 3.5 mm) and two immatures collected on Keteleeria fortunei in Yunnan Prov., China (Zhang & Zhong 1985e). Appearance in life is unrecorded.

Cinara kiusa Hottes Appearance in life is not recorded. Not collected since the original description; BL of aptera 3.2-3.7 mm, other morphs are unknown. On Abies concolor in Oregon, USA.

Cinara kochiana Börner Apterae are greyish-brown to dark grey, slightly wax-powdered, with a darker pattern of small sclerites and often a spinal stripe (see influentialpoints.com/Gallery); BL 4.7-6.1 mm. On Larix spp., forming ant-attended colonies in bark crevices on lower part of trunk or bases of older branches, or (especially in midsummer) on large partly exposed roots. Recorded from most parts of Europe except the Iberian peninsula, but rather rare (Eastop 1972). Also in Russia (Siberia) and Kazakhstan (Kadyrbekov 2017a). Oviparae and males (which can be either apterous or alate) occur in October-November, and eggs are laid in bark crevices. Kloft et al. (1960) summarize biological information and Pontin (1960) found a heavily parasitized population on larch roots. 2n=10. In Japan and Korea, and possibly in China (G. Zhang et al. 1993b), similar aphids have longer antennal hairs and are regarded as a subspecies, C. kochiana kochi Inouye (see Inouye 1962, 1970).

Cinara kuchea Hottes Appearance in life is not recorded; BL of paratype apterae 2.3-2.6 mm. Described from Pinus monticola in Idaho, USA (Hottes 1958g), also on this host in Oregon (aphidtrek.org), and similar but larger aphids (BL up to 3.5 mm) have been collected in British Columbia Canada; “Cinara sp. 3″ of Bradley (1961) is possibly this species. Except for differences in hair length, C. kuchea resembles and may prove to be synonymous with another little-known species on P. monticola, C. hirsuta (q.v.). Biology is unknown. (Bradley’s species formed large, dense colonies on small branches and roots of young trees.)

Cinara lachnirostris Hille Ris Lambers Apterae are darkish-brown; BL 2.4-2.8 mm. On Pinus wallichiana, on branches of young trees near ground level attended mostly by ground-dwelling ants (Myrmica sp.; Naumann-Etienne & Remaudière 1995). Bhagat (1985) records it also on P. roxburghii. In Pakistan, Nepal, India (Jammu and Kashmir; BMNH collection). The alata and first instar larva were briefly described by Remaudière & Binazzi (2003a). The ovipara (collection date unknown) was described and illustrated by Chakrabarti et al. (2020). Bhagat (1981a) reared a parasitoid (Pauesia pini) from this aphid. 2n=8 (Dutta & Gautam 1993).

Cinara (Cupressobium) lalazarica Remaudière & Binazzi Colour of apterae in life is unknown; BL 2.9-3.6 mm. On Juniperus communis (var. saxatilis) in Pakistan, and also recorded from J. sabina in Kazakhstan (Kadyrbekov 2017a). Remaudière & Binazzi (2003b) discuss how this species differs from eight other species of the subgenus Cupressobium with dark tibiae. Other morphs are as yet unknown.

Cinara (Cedrobium) laportei (Remaudière) Apterae are broadly oval and dorso-ventrally flattened, pale brown and grey with black segmental markings and often with a whitish spinal stripe (see influentialpoints.com/Gallery); BL 1.5-2.0 mm. In small dense colonies on twigs and on small shoots of lower branches of Cedrus spp. in Europe, Turkey, North Africa (Algeria, Morocco) and South Africa. Sexuales are produced in October at high altitudes in Morocco, but in many places this aphid is probably anholocylic. Notario et al. (1984) provided a detailed account. Biological control with a specific parasitoid from Morocco was undertaken in southern France (Fabre & Rabasse 1987). Remaudière (1954a) erected the genus Cedrobium for this species which differs from other Cinara in its 5-segmented antennae, strongly sclerotic tergum and curious, mace-like dorsal hairs. However on phylogenetic grounds and taking into account biochemical evidence (Lampel & Burgener 1987), Cedrobium should clearly be treated as a subgenus within Cinara.

Cinara largirostris Zhang & Zhang BL of aptera c. 4.6 mm. On Pinus tabuliformis in Liaoning, China (G. Zhang et al. 1993b). These are possibly small specimens of C. sorini or C. etsuhoe.

Cinara laricifex (Fitch) Apterae are dark brown to bronze, with little or no wax, although immatures often have wax markings (see influentialpoints.com/Gallery) ; BL 2.5-3.5 mm. On Larix laricina, in small numbers on new growth near tips of twigs, or later also forming colonies on branches or trunk (Bradley 1959). Males are alate, and eggs are laid on the bark (Bradley 1961). In Canada and north-eastern USA. This species has sometimes been confused in the North American literature with the European C. laricis (Hartig) – see Hottes (1953a). 2n=10.

Cinara laricifoliae (Wilson) Apterae are dark greenish-brown to black with a dusting of white wax and a pale spinal stripe (Wilson 1915, Bradley 1961); BL 4.2-4.6 mm. On Larix occidentalis, in small colonies on small and medium-sized branches, feeding at bases of needle clusters. Recorded from Oregon, Washington and Idaho, USA and British Columbia, Canada. Life cycle is unknown.

Cinara laricionis Binazzi Apterae are dark bronze; BL 2.8-3.7 mm. In colonies on young growth and one-year-old branches of Pinus nigra var. maritima (= P. laricio) in Italy and Corsica (Binazzi 1980a, b), and apparently specific to this host. Very similar to C. pinimaritimae. Oviparae and alate males occur in October-December (Binazzi 1983c).

Cinara laricis (Hartig) Apterae are dark greyish-brown speckled with black, usually with a wax bloom (see influentialpoints.com/Gallery); BL 3.0-5.1 mm. In small dense colonies on twigs of lower (3- to 5-year old) branches, or on trunks of young larch trees (decidua, kaempferi, × eurolepis, gmelinii). Throughout Europe, and also recorded from Siberia, Kazakhstan (Kadyrbekov 2017a), Mongolia (Szelegiewicz 1963), China (Fang et al. 1983), Korea and Japan (Inouye 1962; but Japanese specimens have fewer siphuncular hairs and are often more heavily pigmented suggesting that they are at least subspecifically distinct). Oviparae and alate males occur in October-November. The copious honeydew crystallizes to form the “Lärchenmanne” which is highly regarded by beekeepers in central Europe (eg. Manino et al. 1985).

Cinara longipennis Matsumura Apterae are dark brown to black with dark legs, body dusted with wax laterally and ventrally (Moritsu 1983); BL 6.0-7.0 mm. On Abies spp. (firma, holophylla, sachalinensis), in large spring colonies with abundant alatae on trunk and older branches of young trees, apterae moving down onto the roots in summer (Inouye 1970). In Japan and Korea. Oviparae occur in Korea in late October (Paik 1972); but the aphid illustrated by Paik as longipennis is much paler, yellow-brown with black markings).

Cinara longirostris Börner Appearance in life is unknown; BL of aptera c. 4.7 mm. Only known from type specimens from Pinus sylvestris in French Alps, which have recently been redescribed by Binazzi (1990), and judged to be a good species. The available material (one aptera and one ovipara) is inadequate for inclusion in a key. Resembling C. pini, except for large size and absence of a mesosternal tubercle.

Cinara (Cupressobium) louisianensis Boudreaux Apterae are green, with a dorsal pattern of white wax (Boudreaux 1948), normally well camouflaged, but more conspicuous on cultivated varieties with yellow foliage; BL 1.7-2.2 mm. On branches of Thuja sp. in southern USA, on a Cupressus sp. in Mexico (Voegtlin et al. 1986, who provided a redescription) and also now recorded from Thuja sp. in Costa Rica (Villalobos Muller et al. 2010). Recorded also from Taiwan (Tao 1958, on Thuja orientalis); Korea (Paik 1972, Seo 1994 as idahoensis); China (G. Zhang 1999, as idahoensis); Tasmania (Eastop 1966) and New Zealand (BMNH collection, leg. V.F. Eastop; on Th, occidentalis and Chamaecyparis lawsoniana). Biology and sexuales are unknown. 2n=12.

Cinara lyallii Bradley Apterae are dark brown to dull black, dusted with grey wax powder; BL c.4.0 mm. In small loose colonies on upper sides of twigs of previous year’s of Larix lyalli growing at high elevations (above 2,000 feet) in Alberta, Canada. Oviparae and both apterous and alate males are recorded, and there is possibly an abbreviated life cycle (Bradley 1961). Closely related to C. laricifoliae.

Cinara maculipes Hille Ris Lambers Apterae are pale brown with legs, especially femora, mottled with dark brown in pantherine fashion; BL 3.7-4.9 mm. On Pinus wallichiana in northern India (Himachal Pradesh, Uttar Pradesh, Jammu, Kashmir), Bhutan (Chakrabarti et al. 2020), Pakistan (Murree, Dungagali, Changla Pass) and Afghanistan. Feeding site is not recorded. Oviparae and apterous males were found in December (Chakrabarti & Raha 1988); the ovipara was described by Remaudière & Binazzi (2003a) and the male by A.K. Ghosh (1982). Das & Chakrabarti (1989) reared a new Pauesia sp. from this aphid. 2n=12 (Kurl & Chauhan 1986a).

Cinara maghrebica Mimeur Apterae are chocolate brown with a dorsal pattern of white wax dust (Mimeur 1934); BL 2.0-2.9 mm. Living, often in dense colonies, on young twigs of Pinus spp. (canariensis, halepensis, pinaster, pinea). In the Mediterranean area (Italy, France, Spain, Morocco, Malta); reported also to occur in Turkey (Akyürek et al. 2010) and Iran (Mortazavi et al. 2015), and also recorded from Argentina (Delfino & Eastop 1982). Garcia Sanchez & Nieto Nafría (1978) provide a detailed morphological comparison with the closely-related C. pinimaritimae. Binazzi (1983a) differentiated populations on P. halepensis in Italy as a subspecies, C. maghrebica ssp. garganica. 2n=10*.

Cinara maritimae see Cinara pinimaritimae

Cinara manitobensis Bradley Apterae are reddish-brown covered with dense grey wax powder; BL 2.4-3.2 mm. Described from Juniperus horizontalis in Manitoba, Canada. Oviparae and apterous males occur in October (Bradley 1963).

Cinara matsumurana Hille Ris Lambers Apterae have a greenish-brown head and thorax and green abdomen; BL 2.3-3.0 mm. On young twigs and branches of Abies spp. (homolepis, sachalinensis) in Japan, feeding among needles, usually on upper parts of young trees. Oviparae and alate males occur in October (Inouye 1970). Records from Turkey (Görür et al. 2012) require further confirmation. C. togyuensis, described from A. holophylla in Korea (Seo 1994) is possibly this species. 2n=10.

Cinara medispinosa (Gillette & Palmer) (fig. 75F) Apterae are light cinnamon-brown to dark metallic brown, with a pattern of black dorsal sclerotic markings (may not be evident in midsummer populations), and some wax powder spinally and laterally (Palmer 1952); BL 3.0-4.0 mm. On bark of tender twigs and young branches of Pinus contorta and its varieties (plus one record from P. albicaulis), in western USA and Canada. Oviparae and males were collected in October. Closely related to brevispinosa and murrayanae; the taxonomy of this group of species requires clarification.

Cinara melaina Boudreaux Apterae are shiny dark brown, almost black; BL 2.8-3.3 mm. Normally apterae have an uninterrupted cross-band across abdominal tergite 8, whereas in C. atlantica, with which it may form mixed colonies, this band is usually interrupted medially. On current and previous year’s growth of Pinus spp. of subsect. Australes, feeding at needle bases; there are also single records from P. resinosa and P. virginiana (Pepper & Tissot 1973). In south-eastern USA, north to Delaware. Males are unrecorded, and oviparae are rare; probably anholocyclic in southern states. Patti & Fox (1981) studied populations on P. taeda in South Carolina. Very similar when slide-mounted to the western US species, C. ponderosae.

Cinara micropunctata Remaudière & Binazzi Colour in life is unrecorded; BL 2.6-3.4 mm. On Abies pindrow at high altitudes in Pakistan. The original description (Remaudière & Binazzi 2003a) includes the alata and first instar. The name draws attention to the finely punctuated dorsal cuticle, a character shared with another Abies feeder, C. tistaensis.

Cinara minoripinihabitans Zhang Body of aptera has a thick covering of wax powder, head and thorax pale, legs black (G. Zhang & Zhong 1989); BL c. 2.4 mm. On Pinus koraiensis in Heilongjiang Prov., China. There is also a record from Juniperus rigida ( Other morphs and life cycle unknown.

Cinara minuta Hottes & Knowlton Appearance in life is unknown, perhaps rather pale; BL (ovipara) 2.3-2.4 mm. Only known from the oviparous morph, collected on Abies lasiocarpa in Utah, USA (Hottes & Knowlton 1954).

Cinara moketa Hottes Apterae are shining dark brown with cross-bands of grey pulverulence; BL 3.3-4.1 mm (Voegtlin 1976). In colonies on branches, or on trunks of young trees, of Pinus lambertiana in California and Washington, USA. Sexuales unknown, apparently anholocyclic in California (Voegtlin 1976). Tilles (1984) studied ant attendance, and predation by an Elaterid beetle. C. hirticula Hottes is probably a synonym.

Cinara mongolica Szelegiewicz & Holman Apterae are brownish, without wax; BL 3.7-3.9 mm. Type specimens were from a small colony, not attended by ants, on young twigs of Pinus sibirica in Mongolia (Szelegiewicz & Holman 1980). Oviparae and alate males were collected in August. Kadyrbekov (2013d) also recorded it from the same host in Kazakhstan. Very similar to C. cembrae, and possibly all records of C. cembrae in the Far East (e.g. Inouye 1970, Pashchenko 1988a, Seo 1994) should be referred to mongolica, in which case P. cembra, P. pumila and P. koraiensis are additional hosts.

Cinara montanensis (Wilson) Appearance in life is unknown, legs are almost entirely black; BL c. 4.0 mm. Known only from the original collection of alatae and apterae on Pinus “laricis” (= P. nigra var.?) in Massachusetts, USA (Wilson 1919).

Cinara montanesa Hottes Appearance in life is unknown; BL of aptera c. 5.7 mm (but that of alata is given as only c. 3.5 mm). Known only from original collection of apterae and alatae on Pinus coulteri in California, USA (Hottes 1961c).

Cinara montanicola (Börner) Apterae are grey-brown with variably-developed black dorsal markings, sometimes lightly dusted with wax; BL 1.9-3.9 mm. On Pinus mugo, preferring undersides of 2- to 3-year old branches including sections without needles, often in sheltered positions near the ground. Colonies may be very dense and are attended by ants. Alatae are frequent in summer, and oviparae and males are produced in September to October. The males are apterous (cf. C. pini), small and slender, with the posterior part of the abdomen woolly and white (Binazzi & de Silva 1993). In mountainous regions of south and central Europe. Binazzi & Covassi (1994) compared it with other species on P. mugo in Italy.

Cinara (Cupressobium) mordvilkoi Pasek Apterae have a light coffee-brown wax-dusted head and thorax, and darker shining bronze abdomen; BL 2.7-3.3 mm. On Juniperus communis in Europe (Sweden, Poland, Slovakia, Italy, Bulgaria, Ukraine, Belarus, Latvia, Lithuania, Estonia, Poland), and also recorded from J. turkestanica (= pseudosabina) in Kazakhstan. In spring and autumn they feed on young shoots and needles, but spend summer months on roots and lower parts covered in soil (Durak 2014a). The life cycle was studied in Poland by Durak (2014a); males are usually apterous (cf. C. cupressi), but alate males are produced in some years. Klimaszewski et al. (1977) described biochemical differences from cupressi, and Durak (2011) explored differences in mitochondrial DNA and morphometrics between C. mordvilkoi and C. juniperi.

Cinara murrayanae (Gillette & Palmer) Apterae are yellow-brown to dark brown; BL 3.0-4.0 mm. On bark of twigs and small branches of Pinus contorta and its varieties, in western North America; records from other hosts are probably misidentifications of other species. Oviparae and alate males in October (Gillette & Palmer 1924). Difficult to distinguish from less sclerotic midsummer individuals of C. medispinosa, and resembling C. atlantica except in length and shape of hairs.

Cinara nepticula Hottes Only known from alate viviparae, BL 3.4-3.6 mm, collected on Picea rubens in Nova Scotia, Canada (Hottes 1958, 1961). Probably closely related to, if not synonymous with, C. braggii and/or C. glehna.

Cinara neubergi (Arnhart) Apterae are of variable colour, light to dark brown, with a slight dusting of wax (original description); BL 4.0-5.0 mm. On bark of twigs of Pinus mugo, feeding among needle bases, with a boreomontane distribution in Europe (Austria, Bulgaria, Germany, Italy, Switzerland, Ukraine, Lithuania). A record from P. sylvestris in Turkey (Akyürek 2019) is doubtful and requires further confirmation. Oviparae occur in September (BMNH collection); males are alate (Lampel & Meier 2003). A member of the C. pinea group. Binazzi & Covassi (1994) compared it with other species on P. mugo in Italy.

Cinara newelli Tissot Apterae are yellowish-brown to dark brown with a covering of grey wax (Tissot 1939); BL 2.7-3.1 mm. Like C. cronartii, this species feeds at rust lesions (Cronartium fusiforme) on Pinus spp. (elliotti, palustris, taeda). In south-eastern USA (Florida, North Carolina, Tennessee); sexuales and the life cycle are not recorded (Pepper & Tissot 1973).

Cinara nigra (Wilson) (fig. 75B) Apterae are shining chocolate-brown, without wax; BL 2.5-3.0 mm. In large dense colonies on undersides of branches, or on trunk of young trees, of Pinus banksiana and P. contorta. In north-western USA east to Michigan, and in Canada from British Columbia to Ontario. Apterous males and oviparae in late September (Pepper & Tissot 1973). Eggs are laid in the cleft between two needles just above the sheath (Bradley 1961, as canatra). Foottit & Mackauer (1990) studied morphological variation within and between populations on P. contorta.

Cinara nigripes Bradley Apterae are dark brown to bluish-black, with scattered white wax (Bradley 1962); BL 2.8-3.4 mm. In dense colonies on trunk of young trees of Picea spp. (glauca, mariana, sitchensis) in Canada (British Columbia, Ontario) and Alaska. Oviparae and small apterous males were collected in mid-August in Ontario.

Cinara nigrita Hottes & Essig Only known from a single alata, distinguished by its uniformly dark wings, collected in Arizona, USA. The host is unknown but surmised to be Pinus ponderosa (Hottes & Essig 1953b).

Cinara nigritergi Mamontova Apterae elongate oval, black, BL c.3 mm. On Pinus sylvestris in Russia (Karelia), Ukraine, Czech Republic, and now also recorded from Sweden (Albrecht 2017) and Kazakhstan (Kadyrbekov 2017a, as C. nigra Mamontova-Soluhka). It was originally described as C. nigra (a homonym of nigra Wilson), and provided with a replacement name by Mamontova (2002). An ovipara (BL 3.6 mm) collected in late August in southern Karelia was described by Stekolshchikov (2011b).

Cinara nimbata Hottes Apterae have a black head and thorax dusted with grey wax, and abdomen grey-green dorsally with lateral areas cinnamon-brown (Hottes 1954c); BL 3.7-4.2 mm. The legs are characteristically patterned, with a contrasting pale band across the dark distal part of each femur and tibiae spotted with black. Alatae have diffuse patches of fuscous on forewings as in C. costata, to which it is closely related. Living singly particularly on stubby branches close to trunk of tree, and attended by ants (Hottes 1954c). On Picea engelmanni in western North America from Colorado to Alaska; an ovipara from an unknown host in Labrador, and an alata collected on P. mariana in Quebec (BMNH collection, leg. F.C. Hottes) also seem to be this species. Oviparae and alate males occur in September in Colorado (Hottes 1954c).

Cinara nuda (Mordvilko) (= escherichi Börner) Apterae are brownish to bronze, very shiny, with little or no wax; BL 3.4-4.3 mm. In large colonies on Pinus sylvestris, on trunk and basal parts of older branches of young trees, or on 2- to 8-year-old parts of leading shoots of older trees. Widely distributed in Europe (England, Finland, Poland, Lithuania, Germany, Czechoslovakia, Austria, Spain), and also reported from Kazakhstan (Kadyrbekov 2017a) and China (G. Zhang et al. 1993b). Apterous males and oviparae occur in central Europe in August-October. Scheurer (1971) studied populations over a 3-year period in Germany. Danielsson (1987) established the correct name for this species. 2n=10.

Cinara obovatae Binazzi & Battisti Apterae are very large, uniformly blackish or dull bronze with two longitudinal rows of small intersegmental black spots, with small dark brown siphuncular cones and mostly pale brown legs; BL 4.2-6.3 mm. On Picea obovata in Central Siberia (Binazzi & Battisti 1998).

Cinara obscura Bradley Apterae are dark brown to black, with pale sections on tibiae; BL c. 2.5 mm. In large dense colonies on bark of small branches of Picea glauca in Canada, from Alberta to Newfoundland (Bradley 1953, 1961), and also recorded from P. engelmannii in British Columbia. Fundatrices were found on P. glauca in Ontario (BMNH collection, leg. Bradley), but sexuales are not recorded. C. pallidipes Hottes, described from P. glauca in Maine USA, may be a synonym.

Cinara (Schizolachnus) obscura (Börner) Apterae are brownish with greyish white covering of wax powder (see influentialpoints.com/Gallery); BL 1.9-2.7 mm. On needles of Pinus nigra (also sometimes on mugo, pinaster and pinea) in Europe, east to Turkey (Klimaszewski & Wojcieckowski 1976) and Georgia (Barjadze et al. 2010a). Specimens collected from P. sylvestris in Ontario, Canada, originally identified as C.(S.) pineti, have now been assigned to this species (see Skvarla et al. 2017). In fact C.(S.) obscura was originally described as a subspecies of C.(S.) pineti, and the distinction between these two species is not at all clear, several authors having regarded them as synonymous. In Spain, fundatrices were found in April-May, alatae in May, and oviparae and males from the end of October to December (Nieto Nafría et al. 2002a).

Cinara occidentalis (Davidson) (fig. 12B) Apterae are golden brown with bluish-white wax either in cross-bands or forming a flocculent covering of body and appendages; BL 2.1-3.2 mm. On bark of one-year-old twigs of Abies spp. in western USA and Canada (British Columbia, ?Quebec). There is one record from a Pseudotsuga sp. in New Mexico (Jousselin et al. 2013). A record from Turkey (Akyürek et al. 2019) is doubtful and needs confirmation. Oviparae and alate males in September-November. Ant attendance was studied by Tilles & Wood (1986). Heavy parasitisation was noted by Knowlton (1930). (N.B. An Indian Abies-feeder, C. tistaensis (q.v.), would probably key out to occidentalis, although clearly not this species.)

Cinara oregonensis (Wilson) Apterae are light brick-red to rust-red, shining; BL 2.5-3.0 mm. On Pinus contorta and P. ponderosa and their varieties in western USA and Canada, living among scales of young cones. The second generation are nearly all alate and fly to new cones. Oviparae and apterous males occur in September-October; eggs are laid on undersides of scales of new cones.

Cinara oregoni Hottes & Essig Appearance in life is not recorded; BL of aptera 4.0-4.4 mm (Hottes & Essig 1953a). On Pinus albicaulis in Oregon USA, and also recorded from this host in British Columbia, Canada. Other morphs and biology are unknown.

Cinara (Schizolachnus) orientalis (Takahashi) Apterae are yellowish-brown in original description from Taiwan (Takahashi 1924a), but recorded as blackish-brown in India (A.K. Ghosh 1982b), covered in fine, dirty white wax meal; BL 1.9-2.4 mm. Feeding in rows along needles of Pinus spp. in India, Japan, Korea, China and Taiwan. A record from Turkey (Şenol et al. 2014b) requires further confirmation. Oviparae and alate males occur in October-November (Inouye, 1970).

Cinara osborni Knowlton (fig. 13A) Apterae are buff-coloured to grey, dusted with wax, with four black spots on thorax (Hottes 1960); BL 3.3-4.2 mm. Living solitarily on the twigs of Abies spp. (concolor, lasiocarpa) in western USA. (Originally described from Pseudotsuga (Knowlton 1942), but this is unlikely to be a normal host.) Sexuales and life cycle are unknown.

Cinara (Cupressobium) oxycedri Binazzi Apterae are greyish-white due to covering of wax powder, with two diverging rows of blackish patches from thorax to abdominal tergites 2 or 3, prominent black siphuncular cones and dark legs; BL 2.2-3.1 mm. On Juniperus oxycedrus, including ssp. macrocarpa in Italy (Tuscany, Sardinia), Corsica and Spain (Binazzi 1996). Alatae were found in the second half of May. A record from Turkey (Şenol et al. 2014b) requires additional confirmation. Sexuales and life cycle are not known.

Cinara pacifica (Wilson) Only known from original collection of 3 alatae and 11 immatures from Abies grandis in northern California. Distinguished by the unusual siphunculi which are flattened and inconspicuous, without hairs or pigmented conical base (Palmer 1945). Hind tibiae are entirely dark.

Cinara palaestinensis Hille Ris Lambers Apterae are chestnut-brown to yellowish-green (Binazzi 1978); BL 2.0-3.3. mm. In large dense ant-attended colonies on new growth of Pinus halepensis (incl. var brutia) in the Mediterranean area and south-west Asia (Israel, Greece, Italy, Malta, Spain, Turkey, Iran). Sexuales have been described from Italy (Binazzi et al. 2016). Binazzi (1983a) differentiated some Italian populations as a subspecies, C. palaestinensis ssp. apulica. 2n=10*.

Cinara (Schizolachnus) parva (Wilson) Apterae are brownish tinged with green, covered with bluish-white wax threads which also extend onto needles; BL 1.6-1.8 mm. On needles of Pinus spp. in eastern USA, and recently recorded from Costa Rica (Villalobos Muller et al. 2010). Oviparae and alate males occur in USA in late October (Wilson 1915).

Cinara parvicornis Hottes Apterae are dark brown, shining, without wax; BL 2.4-2.8 mm. On Pinus banksiana in Canada (Ontario, Manitoba), feeding on needle bases of 1- and 2-year-old branches, and causing yellowing of needles in late spring and summer (Bradley 1952, as ontarioensis). Numerous alatae are produced in the second generation. It was originally described from oviparae and alate males collected on P. contorta in Montana, USA, in September (Hottes 1958d). Bradley (1962, as ontarioensis) also recorded oviparae and alate males on P. banksiana in Ontario in autumn.

Cinara paxilla Zhang Appearance in life unrecorded; BL of aptera c. 4.4 mm. On Pinus sp. in Tibet (Zhang & Zhong 1981b). A short-haired species, possibly close to C. formosana.

Cinara pectinatae (Nördlinger) (fig. 12A) Apterae are usually bright olive-green with three diffuse paler green longitudinal bands, red eyes and brown head and appendages (see influentialpoints.com./Gallery); BL 2.8-5.0 mm. A brown form, similar in colour to the buds and stem, can occur in late summer/autumn. On Abies spp., especially A. alba (= pectinata). They feed singly on small branches where they sit on the needles, feeding at the junction of petiole and stem. It occurs throughout Europe, and also in Turkey, Kazakhstan and Siberia. Oviparae and alate males occur in mid-September to October. It is a copious honeydew-producer, important to forest bee-keepers in Central Europe, and its population dynamics has been much studied because of this (e.g. Bloc 1987 in France; Liebig 1988 in Germany). 2n=6* (Germany, 2 samples; but Rukavishnikov (1979) recorded 2n=12 from Russia).

Cinara pergandei (Wilson) (fig. 80D) Apterae are rather globose, shiny, brown to bronze, with or without wax powder (see influentialpoints.com/Gallery); BL 3.0-4.4 mm. Active insects living singly on twigs and new shoots of many Pinus spp. in subsections Australes and Contortae. In eastern USA, across Canada, and in Cuba (Hernandez & Rodriguez 1985) and Costa Rica (Villalobos Muller et al. 2010). Oviparae and alate males in September-November (Pepper & Tissot 1973). Relations with parasitoids and ants were recorded in Manitoba (Bradley & Hinks 1968). 2n=14*.

Cinara (Cupressobium) petersoni Bradley Apterae have dark greyish-brown head and thorax and greenish-brown abdomen, and are dusted with greyish-white wax (Bradley 1963); BL 2.9-3.4 mm. Described from Juniperus horizontalis in Manitoba, Canada, and also on Juniperus sp. in California (aphidtrek.org) ; specimens conforming to this description have also been collected in Majorca and Israel (on J. oxycedrus; BMNH collection). Alatae are undescribed, and sexual morphs and life cycle are unknown.

Cinara piceae (Panzer) (= grossa Kaltenbach) Apterae are uniformly jet black “resembling the texture and shape of old droplets of tar” (Carter & Maslen 1982, and see influentialpoints.com/Gallery); BL 3.2-6.7 mm. Forming large colonies in spring on undersides of older branches and on trunks of Picea spp., often moving to ground level or roots in summer (Pintera 1966; as grossa). Throughout Europe, and what is nominally the same species also occurs in Kazakhstan (Kadybekov 2017a), east and west Siberia (Pashchenko 1988a; Stekolshchikov & Khureva 2020), China and Japan, and has more recently been reported from Argentina (Delfino & Binazzi 2002). Numerous alatae are produced in May-June. Oviparae appear in September-November, move to current year’s growth and lay wax-dusted eggs on needles. Males are apterous, small and usually overlooked according to Carter & Maslen (1982). Inouye (1970; as grossa) reported alate males in Japan, but this may be an error as the four males in his collection are all apterous (V.F. Eastop, personal observation). Starý (1976) studied the natural enemy complex in central Europe. 2n=10 (Blackman & Eastop 1994, England) or 12 (Rukavishnikov 1979, European Russia).

Cinara piceicola Cholodkovsky (= viridescens, = stroyani) Apterae have dark brown head and thorax and pale olive-buff abdomen with two longitudinal faint greyish-green stripes (see influentialpoints.com/Gallery); BL 2.1-4.2 mm. On Picea spp., especially abies, in colonies on bark of woody shoots between needle-bases in spring, moving to older branches and roots in summer (Börner & Franz 1956). In north, west and central Europe; also in Turkey, Iran (Rezwani et al. 1994) and Kazakhstan (Kadybekov 2017a), and G. Zhang et al. 1993b) recorded it from China. Numerous alatae are produced in May-June. Oviparae and apterous males occur from July onwards (Pintera 1966). Kunkel (1990) studied its relationship with ants. Danielsson (1987) discussed the confused nomenclature of this species. C. alba Zhang 1982, described from P. asperata in China, is very similar and possibly a synonym. 2n=8.

Cinara pilicornis (Hartig) Apterae are either orange-brown or greyish-green, usually with an overall dusting of wax (see influentialpoints.com/Gallery); BL 2.1-4.7 mm. On Picea spp., initially in spring in small colonies on undersides of the previous year’s twigs, moving onto new growth after bud-burst. It may also colonise Tsuga heterophylla (Carter & Maslen 1982, Forbes & Chan 1976). Throughout Europe (incl. Iceland), Turkey, Iran (Rezwani et al. 1994), Kazakhstan, India (one alata; A.K. Ghosh 1982b), China (G. Zhang et al. 1993b), Japan (as C. nopporensis Inouye, which may be at least subspecifically distinct; see Eastop et al. 1998), and introduced to Australia, New Zealand, North and South America (Eastop 1972, Sunde 1984, Carrillo 1977). Numerous alatae are produced in May-July, oviparae and alate males in early August-November. Kettner (1985) studied populations in relation to honeydew production in Germany, Parry (1979) studied egg survival, Michelena Saval & Gonzalez Funes (1988a) recorded parasitoids, and Edwards (1981) described a fungal infection. 2n=10 (in UK and New Zealand) or 14 (Rukavishnikov 1972, European Russia).

Cinara pilosa (Zetterstedt) Apterae are rather pale yellowish- to reddish brown or olive green, with spotted legs; BL 3.1-5.2 mm. On Pinus sylvestris, feeding singly or in small groups on current-year growth of older branches of mature trees (eg. among male cones), or on stunted twigs of younger trees (Holopainen & Heikinheimo 1983). In north, east and central Europe, and also recorded from Italy (Binazzi 1988), Andorra and Spain (Binazzi et al. 1995b), and western Siberia (Stekolshchikov & Novgorodova 2013). Oviparae are recorded fom Italy and Spain in September, and from Switzerland in November. Danielsson (1987) discussed the nomenclature and differences from the very similar C. pinea. The discriminants between these two species given in the key are based on his work, but do not work for all populations, and additional studies are needed to clarify the number and relationships of species in the “pinea group”. 2n=8*.

Cinara pinea (Mordvilko) (fig. 75G, fig. 80C) Apterae are shiny orange-brown in spring, to grey or dark brown in summer, finely spotted with dark brown or black and dusted with wax (see influentialpoints.com/Gallery); BL 3.1-5.2 mm. On new shoots of Pinus sylvestris, and sometimes other Pinus spp. including nigra in dry areas. Throughout Europe, Turkey, Georgia, Kazakhstan (Kadyrbekov 2017a), eastern Siberia, (Pashchenko 1988a), China (G. Zhang et al. 1993b), and introduced to North America (see Voegtlin & Bridges 1988). Oviparae and alate males occur in October. Biological studies include Kidd (1985, 1990; population dynamics), Kidd & Tozer (1984, 1985a,b; induction of alatae, overwintering survival, distribution within canopy), Sudd (1983; ant attendance) and Michelena Saval & Gonzalez Funes (1988a; parasitoids). 2n=10, 11 or 14 (Blackman 1990), suggesting that there may be sibling species still to be recognised (see also C. pilosa).

Cinara (Schizolachnus) pineti (Fabricius) Plate 16e Apterae are dark greyish-green covered in wax meal giving a light bluish-grey appearance (see influentialpoints.com/Gallery); BL 1.2-2.5 mm. On numerous Pinus spp., especially on young trees, forming dense colonies in rows along the previous year’s needles. Oviparae and alate males occur in October-December; in some years colonies persist into early winter and possibly anholocyclic overwintering occurs in mild winters (Carter & Maslen 1982). Common and widespread in Europe; also in Turkey, Kazakhstan, east Siberia (Pashchenko 1988a) and China (Zhang & Zhong 1985e) although some of these records may apply to C. (S.) obscura (q.v.). Holman (1990) studied morphological changes in spring generations, Kidd et al. (1985) studied the association with Eulachnus agilis and Thompson (1977) studied its effect on tree growth. 2n = 18 (Russia; Rukavishnikov 1972) or 2n = 10 (England; Blackman 1980); the first chromosome number may possibly apply to C. (S.) obscura.

Cinara pini (Linnaeus) (fig. 80G) Apterae are grey or greyish-green with black markings, and with either a slight bronze iridescence or a dusting of grey wax (see influentialpoints.com/Gallery); BL 1.9-3.7 (fundatrices up to 4.4 mm). On young shoots of Pinus sylvestris in spring (Pintera 1966), later on undersides of older foliated twigs or branches. There are also records from many other Pinus spp., but many of these are likely to be misidentifications or misapplications of the name pini (Eastop 1972). Throughout Europe, and also in Turkey, Iran (Nazemi et al. 2013), Kazakhstan (Kadyrbekov 2017a), Siberia (Pashchenko 1988a) and Japan (Moritsu 1983). North American records cited as pini are probably of pinea or a native species. Oviparae and apterous and/or alate males occur in September-October. Larsson (1985) studied within-crown distribution of C. pini, Sudd (1983) studied ant attendance, and Michelena Saval & Gonzalez Funes (1988a) recorded parasitoids. 2n=10.

Cinara piniarmandicola Zhang & Zhang BL of aptera c. 3.1 mm. On Pinis armandii in Hebei, China (G. Zhang et al. 1993b). Alatae are undescribed.

Cinara pinidensiflorae (Essig & Kuwana) Apterae are dark reddish-brown with silvery wax dorsal markings (Essig & Kuwana 1918); BL 2.3-3.5 mm. In dense colonies on twigs and branches of oriental Pinus spp. in Japan, Taiwan, Korea and China. Oviparae and alate males occur in Japan in October-November (Inouye 1970). C. atratipinivora Zhang (Zhang & Zhong 1982) seems to be a synonym. 2n=10.

Cinara piniformosana (Takahashi) Apterae are blackish-brown dusted with grey wax dorsally, orange ventrally, with black legs; BL 2.8-4.2 mm. On young trees of Pinus densiflora and P. thunbergii; in spring at bases of new shoots, later moving to 1- or 2-year-old parts of branches (Inouye 1970, Furvie et al. 1990). In Japan, Korea, Taiwan, Siberia (Pashchenko 1988a), China (G. Zhang et al. 1993b), Malaya (BMNH collection), and on introduced oriental pines in Brazil (Eastop 1976). Holocyclic in Japan, with alate males (Inouye). 2n=10.

Cinara pinihabitans (Mordvilko) Apterae and alatae are reddish-brown with dark segmental markings, often made more distinctive by a contrasting pattern of white wax; BL 3.1-4.6 mm. On bark of thin 2- to 8-year-old branches of Pinus sylvestris, and also recorded from P. mugo and P. uncinata. Colonies are rarely seen, possibly because they tend to occur high in crown of tree (Stroyan 1973). Widely distributed in Europe (Eastop 1972), and in Turkey, Iran (Heidari Latibari et al. 2016), in Kazakhstan (Kadyrbekov 2017a), and G. Zhang et al. (1993b) record it from China. Oviparae (see influential points/gallery) and alate males in October.

Cinara pinikoraiensis Zhang Appearance of apterae in life is unknown; BL c. 3.5 mm. On Pinus koraiensis in China (Zhang & Zhong 1989). Oviparae and males (undescribed) were collected in October. Perhaps this is a long-haired form of C. watanabei.

Cinara pinimaritimae (Dufour) (= maritimae of authors including Blackman & Eastop 1994) Plate 16a, fig. 80B Apterae are brownish, covered in greyish wax dust; BL 2.6-4.1 mm. Living separately or in small groups among needle bases on young twigs of Pinus spp., with a relatively wide range of hosts in subsect. Pinus. In the Mediterranean area and Middle East, and introduced into South America (Argentina, Brazil, Chile). Oviparae and alate males in October in Italy (Binazzi 1978). Anholocyclic overwintering probably occurs frequently. Mustafa (1987) studied population biology in Jordan, and Michelena Saval & Gonzalez Funes (1988a) recorded parasitoids (Pauesia spp.) in Spain. Probably this aphid is confused in earlier literature with C. pinea (Eastop 1976). Nieto Nafría et al. (2002a) corrected the name of this species, previously known as C. maritimae. 2n=16.

Cinara piniphila (Ratzeburg) Apterae are greyish-brown with a rusty tinge, dusted with wax (Heinze 1962); BL 3.9-4.5 mm. On bark of 1- or 2-year-old twigs of Pinus sylvestris, feeding among needles. In northern, central and southern Europe, Turkey (Şenol et al. 2014), and G. Zhang et al. (1993b) record it from P. tabuliformis in China. Sexual morphs occur in northern Europe from mid-August to October, and were fully described by Danilov et al. (2019a). Tsinovsky & Yegina (1979) used Entomophthora virulenta in biocontrol of this species in Latvia. [Molecular studies of both mitochondrial (COI) and nuclear (EF-1a) indicate a close relationship and possibly synonymy of C. piniphila with C. pinea, and there is a strong possibility that the morphological differences between them may be due to differences in habitat and feeding site (see Havelka et al. 2020).]

Cinara (Schizolachnus) piniradiatae (Davidson) Plate 16f Apterae are dark olive green to brown, with grey flocculent wax coating; BL 2.0-2.5 mm. On needles of numerous Pinus spp., widely distributed in North America. Oviparae and alate males occur in September-November. The male genitalia were described and illustrated by Wieczorek (2012). There have been studies in Canada of its life history and ecology (Gröbler 1962), population dynamics on Pinus resinosa (Sharma & Laviolette 1968a), oviposition behaviour (Sharma & Laviolette 1968b), predators (Gagne & Martin 1968) and fungal diseases (Gröbler et al., 1962; Tyrrell & Macleod, 1975).

Cinara piniradicis Bradley Apterae and alatae are dark brown, without wax; BL 4.0-4.8 mm. In colonies on roots of young Pinus banksiana in central Canada and USA (Michigan). Apterous males and oviparae occur in September-October; eggs are laid singly on inner side of needles just above the follicle sheath (Bradley & Wighton 1959, Pepper & Tissot 1973).

Cinara pinivora (Wilson) Apterae have a shiny dark brown head, lighter brown thorax and abdomen with dark dorsal sclerites and spots of grey wax, black steep-sided siphuncular cones, and all legs with extensive pale yellow sections (Tissot 1944; as osborni); BL 3.3-4.2 mm. In dense colonies at tips of branches, or scattered along older sections of twigs, of Pinus spp. of subsections Australes and Contortae in eastern USA (plus one record from Montana – aphidtrek.org) and Canada; introduced to Argentina (Delfino & Binazzi 2002), Brazil (Penteado et al. 2004), and to plantations of P. sylvestris and P. oocarpa in Africa (Kenya, Malawi; BMNH collection). A record from Turkey (Görür et al. 2011a) requires further confirmation. Oviparae and alate males occur in October-November (Pepper & Tissot 1973).

Cinara polymorpha Remaudière & Binazzi Apterae are large, long-bodied, pale green, with dorsum covered with a fine layer of transparent wax, and with small pale siphuncular cones; BL 5.1-6.3 mm. The species is remarkable in that alatae have much longer hairs than apterae on the antennae and legs. On Abies pindrow in Pakistan (Dungagali), presumably feeding on the needles. Fundatrices were collected in mid-May, and a single ovipariform individual in late May (Remaudière & Binazzi 2003a).

Cinara ponderosae (Williams) (fig. 80F) Apterae have a dull brown head and thorax and shiny golden brown abdomen, with a dorsal pattern of wax patches; BL 2.6-4.2 mm (Voegtlin 1976). On bark usually near tips of twigs and small branches of Pinus ponderosa and its varieties; sometimes on other Pinus spp. in subsection Ponderosae, and also recorded from planted P. canariensis and P. roxburghii. Records from other pines may be misidentifications. In western North America. Oviparae and alate males occur in October-November in Colorado (Palmer 1926), but populations studied in the Californian Sierra Nevada were anholocyclic (Voegtlin & Dahsten 1982). 2n=10.

Cinara pruinosa (Hartig) (= bogdanovi Mordvilko) Apterae are dark green or brown, sometimes with a bronze metallic tinge, with blackish markings, lateral patches of wax which can be quite extensive, and prominent black siphuncular cones (see influentialpoints.com/Gallery); BL 2.4-5.0 mm. In small colonies on woody twigs of Picea spp. in spring, but later found at base of trunk and on roots in ant shelters (Pintera 1966). Oviparae and alate males occur in September-October, but anholocyclic overwintering on roots also occurs. Throughout most of Europe, Turkey, Iran (Rezwani 2004), Kazakhstan (Kadyrbekov 2009b), and in North America (most records as palmerae; see Voegtlin & Bridges 1988) and now also South America (Chile: Nieto Nafría et al. 2018). C. rara Bradley, described from sexuales on Picea mariana in Saskatchewan, is also a synonym. It has also recently been recorded from South America (Argentina: Mier Durante et al. 2011). Similar populations in Japan (Inouye 1970) and Siberia (Pashchenko 1988a) tend to be shorter-haired and are regarded as a subspecies, C. pruinosa ezoana Inouye, the population dynamics of which was studied by Furuta & Takai (1983). However there is also considerable variation in hair length in European populations. 2n=10.

Cinara (Cupressobium) pseudosabinae (Nevsky) Apterae are pale brown to bluish green with mainly pale antennae and legs; BL 2.0-2.9 mm. Described from Juniperus pseudosabina in Central Asia (Uzbekistan). Nevsky’s specimens were redescribed by Zhuravlev (2003), who recorded it also from Russia (north Caucasus), and this species has also been redescribed from J. excelsa in Pakistan (Remaudière & Binazzi 2003b). Oviparae were collected in Pakistan in October.

Cinara pseudoschwartzii Palmer Apterae are brownish-black to dull black with white wax markings; BL 3.3-4.0 mm. On bark of twigs of Pinus ponderosa in Colorado USA (Palmer 1936). Holocyclic (fundatrices in May), but sexuales are undescribed.

Cinara pseudotaxifoliae Palmer Apterae are dark brown to black with burnt orange mid-dorsal area, and wax powder on head, prothorax and in large patches anterior and posterior to siphunculi (Palmer 1952, Voegtlin 1976); BL 2.2-3.4 mm. On two-year-old or older twigs and branches of Pseudotsuga menziesii in western North America. Oviparae and alate males occur in October-November. Schowalter et al. (1988) studied ant attendance. C. dubia Hottes & Essig, described from Oregon, is probably a synonym, and C. vagabunda (q.v.) is possibly only a long-haired variant population.

Cinara pseudotsugae Wilson Apterae are rather shiny amber to golden brown, paler medially, sometimes tinged with green with paired dusky to black dorsal markings and/or slight wax dusting laterally and intersegmentally; BL 2.4-3.7 mm. On terminal shoots and branches of Pseudotsuga menziesii in western North America, attended by ants (Johnson & Lyon 1988, p.85). Oviparae and alate males occur in October (Wilson 1912). Smith & Schowalter (2001) studied the effects of this species on shoot and root growth of Douglas-fir seedlings. C. taxifoliae Swain is a synonym (Voegtlin 1976). Palmer (1926, 1952) and Hottes (1961) apparently had a different species with darker legs, possibly a form of pseudotaxifoliae. See also C. splendens.

Cinara puerca Hottes Apterae are shiny dark grey to brownish-black dorsally, sometimes with a paler abdominal spinal stripe, and pale reddish-brown ventrally, without wax (Hottes 1954c); BL 3.0-4.4 mm. Favret & Voegtlin (2004b) provided a redescription. On trunk and branches of Pinus edulis, in crevices in bark and/or in ant shelters, or on roots; also collected on shoots of P. monophylla. Recorded from Arizona and Colorado, USA. No sexuales known, and the populations studied in Colorado were anholocyclic (Hottes 1955a).

Cinara pulverulens Gillette & Palmer Apterae are yellowish-brown with black pleural markings, covered with dense white wax powder; BL 2.2-2.9 mm. On bark of small twigs of Juniperus spp. in western USA (Colorado, Idaho, Utah, New Mexico). Oviparae and alate males in September-October (Palmer 1952). It seems likely to be a synonym of C. burrilli, as it differs only in hair length, colour and amount of wax.

Cinara radicivora Voegtlin Apterae are shining dark greenish-black with scattered brown patches; BL 4.4-6.1 mm. On root crown of Abies concolor, attended by Camponotus spp., in California and Oregon, USA (Voegtlin 1983). Life cycle and sexuales are unknown.

Cinara rigidae Hottes Apterae are “probably black” (Hottes 1958a); BL c. 2.4 mm. Known only from one aptera and two alatae collected on Pinus rigida in New Hampshire, USA; type specimens were redescribed by Pepper & Tissot (1973). Very similar to C. westi. Biology and sexuales are unknown.

Cinara rubicunda (Wilson) Apterae are light brown or pinkish, covered with white powder except for thin spinal stripe (Wilson 1915); BL c. 3.5-3.6 mm. On twigs of Juniperus occidentalis in Oregon, USA. Sexuales and life cycle unknown. Bradley’s (1951) account was based on a misidentification of C. juniperi (Voegtlin & Bridges 1988).

Cinara russellae Pepper & Tissot Apterae are rather shiny black; BL 2.0-2.5 mm. Found in July in small groups at bases of needles of Pinus banksiana, feeding on the sheath area rather than on the branch, but in autumn oviparae and apterae were found in dense colonies with C. nigra on the trunk and branches (Pepper & Tissot 1973). Only known from two collections in Michigan, USA. Oviparae and alate males occur in late September.

Cinara saccharinipini Hottes Appearance in life is not recorded: BL 3.4-4.0 mm. On Pinus lambertiana in California, USA. Apterous and alate viviparae collected in December, indicating anholocycly (Hottes 1958j). Biometric data given by Voegtlin (1976) indicate a close relationship to C. moketa, except for a greater number of accessory hairs on R IV (14-16, as opposed to 8-9 in moketa).

Cinara saraswatae Das & Raychaudhuri Apterae are long-bodied, dark brown; BL 2.8-3.3 mm. Feeding at bases of needles on tender stems of an unidentified Pinus sp. in Nepal (Das & Raychaudhuri 1983). No other morphs are known.

Cinara saskensis Bradley Apterae are dark grey, without wax; BL 5.1-5.6 mm. On bark of main roots of Picea glauca, near the trunk and just below ground level (Bradley 1962). In Manitoba and Saskatchewan, Canada. Life cycle and sexuales are unknown.

Cinara schimitscheki Börner Apterae and alatae are broadly oval and somewhat flattened dorso-ventrally, dark brown covered with wax powder; BL 3.3-5.2 mm. In spring at twig apices among previous year’s needles, later on older branches under bark (Pintera 1966). The host is usually Pinus nigra (incl. var. maritima), sometimes P. mugo, P. pinea or P. leucodermis. It occurs throughout Europe (except Scandinavia and the Baltic), Turkey, Crimea, and is also recorded from P. tabuliformis and P. massoniana in China (G. Zhang et al. 1993b). Scheurer (1976) found and described oviparae (and “sexuparae”) in Germany. The oviparae lack a perianal wax ring and occur from August to October in central Europe, and in October-November in southern Europe (Binazzi & Scheurer 2009). Males are undescribed. Binazzi (1978) gave an account of schimitscheki in Italy, and Michelena Saval & Gonzalez Funes (1988a) recorded its parasitoids in Spain. 2n=10.

Cinara schuhi Hottes Apterae probably have a light brown abdomen but darker head, thorax and legs (Hottes 1957a); BL 2.5-2.9 mm. On Abies concolor in Oregon, USA, and also recorded from A. lasiocarpa in Washington (aphidtrek.org). Biology and sexuales are unknown.

Cinara schwarzii (Wilson) Apterae are cinnamon-brown, with a black area between the siphuncular cones, sometimes extending further over abdominal dorsum, and white spots on thorax and in a spinal row on abdomen (Voegtlin 1976); BL 2.2-4.2 mm. In medium to large colonies, mostly on 1- to 3-year-old branches of Pinus ponderosa and its varieties, also on P. jeffreyi and P. cooperi, in western USA and Mexico (Ortiz 1982). Records from other hosts and localities are suspect. Oviparae and males occur in September-October; eggs are laid in rows on the needles (Palmer 1926, and fig. 7).

Cinara setosa (Börner) Apterae are probably dull brown speckled with wax powder; BL 3.1 -3.8 mm. On older twigs of Pinus mugo in alpine Europe (Austria, Italy). A record from Turkey (Şenol et al. 2014b) requires further conformation. Binazzi (1991) redescribed and illustrated the apterae, and subsequently (Binazzi 1994a) described an alata, as well as an oviparae and alate males collected in late October. [C. meridionalis Mamontova, described from P. nigra var. pallasiana in Ukraine, is similar but with more accessory hairs on R IV and a relatively shorter ANT V (Mamontova 2012).]

Cinara setulosa Hottes & Essig Appearance in life is unknown, probably rather pale (males have a bright green abdomen, Hottes 1957b); BL of aptera 4.5-5.0 mm. On lower branches of Abies magnifica var. shastensis in Oregon USA. Oviparae and alate males occur in late September.

Cinara shinjii Inouye Apterae have a pale brown head and thorax and dark greyish-brown abdomen, with an extensive pattern of white wax powder (Moritsu 1983); BL 2.5-3.0 mm. On twigs, branches and stem of Pinus parviflora in Japan, forming large colonies on lower parts of tree in summer (Inouye 1970). Also recorded from P. parviflora in Korea (Seo 1994), although Paik’s (1965) account of C. shinjii on P. koraiensis in Korea is of a different, shorter-haired species. Alatae are produced in July, oviparae and alate males in September-October. Recently this species has been found in Germany on planted P. densiflora (Scheurer et al. 2020); these authors provide redescriptions of apterae, alatae and oviparae. Similar to C. pinidensiflorae, and their separate identity requires additional confirmation (Eastop et al. 1998).

Cinara sitchensis Hottes Appearance in life is unknown; BL of aptera 2.4-2.6 mm. Other morphs are unknown. Described from Picea sitchensis in California USA (Hottes 1958b); also collected in Oregon (BMNH collection) and Washington (aphidtrek.org).

Cinara (Cupressobium) smolandiae Danielsson & Carter Apterae are pale brown to dusky brownish grey with dull bronze metallic highlights in sunlight, and brown-black siphuncular cones (see influentialpoints.com/Gallery); BL 2.9-4.4 mm. Feeding in bark crevices or rust cankers 1-2 m above ground on stems of old (50-year plus) trees of Juniperus communis (including var. sibirica). Described from Sweden (Danielsson & Carter 1993), recorded also from Finland (Albrecht 2010) and north-west Russia (Zhuravlev 2003), and recently found to occur in Scotland, where small specimens were found on thin, lignified stems (Baker & Blackman 2014). Sexuales and life cycle are unknown.

Cinara solitaria (Gillette & Palmer) Apterae are tick-like in behaviour and appearance, dull brown to yellow-brown, slightly wax-dusted, with six longitudinal rows of black dots (Palmer 1952); BL 2.5-3.0 mm. On Pinus ponderosa, living singly on bark of tender shoots, positioned head downwards at bases of needles. In Arizona (BMNH collection), Colorado and Utah, USA. Oviparae occur in early October, males not recorded.

Cinara sonata Hottes (fig. 13B) Apterae are broadly oval, dull reddish-brown to dary grey with numerous small irregularly distributed black spots, and at least in the ovipara there are patches of white wax powder on pronotum and narrow cross-bands of wax on abdominal tergites (Hottes 1955c, 1957b); BL of aptera 5.3-7.0 mm. On undersides of small branches of Abies magnifica var. shastensis, and also recorded from A. concolor, in western North America. Oviparae and alate males occur in late September.

Cinara soplada Hottes Appearance in life is unknown; BL of aptera c. 3.1 mm. Apterae collected on an unidentified scrub spruce (Hottes 1956c), but the alate male was subsequently described from Picea glauca (Hottes 1958a). Only known from Maine, USA. Alate viviparae and oviparae are not recorded.

Cinara sorini Inouye Apterae are long-bodied, brown to dark brown: BL 6.1-6.8 mm. On Pinus thunbergii in Japan, living on bark of trunk of old trees, usually close to the ground in ant shelters (Inouye 1970). Life cycle and sexuales are unknown. Closely related to, and perhaps synonymous with, C. etsuhoe.

Cinara spiculosa Bradley Apterae are dark brown; BL c. 3.0 mm. On small twigs of Larix laricina, often in loose colonies on lower branches, closely resembling old buds (Bradley 1961). Oviparae and alate males occur in autumn (Bradley). In Canada, from Alberta across to Nova Scotia.

Cinara splendens (Gillette & Palmer) Apterae are rusty-brown, slightly dusted with wax on head, lateral areas and intersegmentally, with conspicuous black patches on metanotum and abdominal tergite 1, and a pale median dorsal area on anterior abdomen; BL 2.2-2.6 mm. On bark of twigs of Pseudotsuga menziesii in Colorado USA. Oviparae and alate males occur in September-November. Very similar to C. pseudotsugae, but apparently with consistent differences in size and pigmentation (Gillette & Palmer 1924).

Cinara strobi (Fitch) Apterae are long-bodied, shiny metallic to dull grey-black, with a spinal stripe and spots of white wax (see influentialpoints.com/Gallery); BL 3.0-3.4 mm. Often in large colonies on branches and upper trunk of Pinus strobus throughout its range in eastern North America. Records from P. banksiana and P. resinosa need confirmation (MacGillivray 1955). Oviparae and alate males occur in September-November. Eggs are laid in rows on needles (Bradley 1951). Weigel & Baumhofer (1948) described damage to white pines. 2n=10*.

Cinara subapicula Zhang Appearance in life is unknown; BL of aptera c. 3.3 mm. A long-haired species described (alatae only) from unidentified Pinus spp. in Tibet (Zhang & Zhong 1981b).

Cinara subterranea Bradley Only the sexuales are recorded (Bradley 1956). Oviparae are greyish-brown with dark brown legs and antennae, BL c. 3.3 mm, collected mid-September. On roots of Larix laricina in Alberta and Manitoba, Canada, forming dense colonies (Bradley 1961). Males are alate.

Cinara taedae Tissot Apterae are dark brown to brownish-black, often shining; BL 2.1-3.1 mm. Feeds on bark between needle bases of 1- or 2-year-old growth, or sometimes on current terminal shoots (Pepper & Tissot 1973). On Pinus spp. of subsections Australes and Contortae, especially P. taeda and P. rigida, in western USA south from Pennsylvania. Oviparae and alate males were collected on P. rigida in Pennsylvania in October-November.

Cinara taiwana (Takahashi) Apterae are blackish-brown, shining; BL c. 3.8 mm. On Tsuga formosana at a high altitude in Taiwan (Takahashi 1925), possibly also on Tsuga and Torreya nucifera in Japan (as Cinara “sp. E” of Eastop et al. 1998), and there is a record from Pinus massoniana (an unlikely host) in China (G. Zhang et al. 1993b). Biology and sexuales are unknown. The description of C. ozawai, on Tsuga diversifolia and Chamaecyparis obtusa in Japan (Inouye 1970) does not distinguish it from C. taiwana.

Cinara takahashii Chakrabarti, Medda & Kanturski [Replacement name for Cinara orientalis (Takahashi 1925), by Chakrabarti et al. (2020).] Apterae are blackish-brown, shining, not waxy; BL 4.5-5.0 mm. Described from Pinus sp. in Taiwan (Takahashi 1925); in Japan it feeds on branches of Pinus densiflora, in ant shelters constructed by Lasius niger (Inouye 1970). Also recorded from Korea (Paik 1972), India (Manipur), Bhutan, and there are specimens in the BMNH collection from China (on P. armandii) and Nepal (1 alata). Sexuales and life cycle are unknown. G. Zhang (1985) described a subspecies (C. takahashii ssp. lijiangensis) from P. yunnanensis in China, and this was given full species status by G. Zhang et al. (1993b), but the differentiation seems to be based on differences in antennal sensoriation that are probably within the normal range of the species.

Cinara tanneri (Knowlton) Appearance in life is not recorded; BL of aptera 3.3-4.0 mm. On pinyon pines (P. edulis, P. monophylla) in Utah USA. Sexuales and biology are unknown, and the alatae in the original description were edulis according to Hottes (1960b). Favret & Voegtlin (2004b) reviewed this species.

Cinara (Cupressobium) taurica (Zhuravlev) Apterae are dark reddish brown with a waxy coat; BL 2.0-3.2 mm. Found in May-June on young branches of Juniperus excelsa in Ukraine, Russia and Armenia (Zhuravlev 2003). Sexuales and life cycle are unknown.

Cinara tellenica Binazzi & Strangi Apterae are blackish, without wax markings; BL 2.7-3.4 mm. In ant-attended colonies on branches of Cedrus atlantica in the Tell Atlas mountains, Algeria (Ayache et al. 2020). Morphological differences from the closely related C. cedri (including its subspecies C. cedri brevifoliae) are tabulated by Ayache et al. (2020), who also studied the molecular phylogeny. Sexuales and life cycle are unknown.

Cinara tenuipes Chakrabarti & Ghosh Appearance in life is not recorded; BL of aptera 3.6-4.3 mm. Originally described as a subspecies of abieticola (= confinis), from a host plant uncertainly identified as Abies (Chakrabarti et al. 1974) in Himachal Pradesh, India. However, Stary & Raychaudhuri (1982) described two new species of parasitoid (Pauesia) from this aphid, and the host is given as Pinus sp. There is also a record from P. smithiana (Chakrabarti & Sarkar 2001), and an unlikely record from Juniperus communis (Kar et al. 1990), so the true host association is very much in doubt. Differences from two very similar species (eastopi and bhutanica) are tabulated by Chakrabarti et al. (2020). 2n=12.

Cinara terminalis (Gillette & Palmer) Apterae are light cinnamon brown, sometimes tinged with green, darkest in centre of dorsum, with some wax dusting on head, thorax and sometimes intersegmentally on abdomen (Gillette & Palmer 1924); BL 2.0-3.4 mm. Described from specimens living solitarily and inconspicuously on bark of young tender twig terminals of Pinus edulis in Colorado. However, Favret & Voegtlin (2004b) noted that C. terminalis frequently aggregated into colonies. They considered C. nitidula Hottes to be a synonym, although this is described as a darker aphid colonising older terminal branches of mature trees (Hottes 1960b). C. terminalis also occurs on P. monophylla, and is recorded from California, Utah, Arizona, Nevada, New Mexico and Mexico. [The Mexican records (Ortiz 1982) are from Pinus cembroides and several other Pinus spp., including michoacana and pseudostrobus in subsect. Ponderosae. Paratypes of C. terminalis in BMNH collection resemble small C. ponderosae, so there may be confusion between these species.]

Cinara thatcheri Knowlton & Smith Appearance in life is unknown; BL of aptera 3.2-4.5 mm (Knowlton & Smith 1938b). On Pinus ponderosa in western USA and British Columbia. Biology and sexuales are unknown. Very similar to, and possibly synonymous with, C. schwartzii.

Cinara tibetapini Zhang Appearance in life unknown; BL of aptera c. 6.5 mm. On Pinus sp. in Tibet (Zhang & Zhong 1981b). Other morphs and biology are unknown.

Cinara tistaensis Agarwala & Raychaudhuri Appearance in life is unknown; BL of aptera 3.7-4.0 mm. On on unidentified Abies sp. in Sikkim, India (Agarwala & Raychaudhuri 1982). Other morphs and biology are unknown.

Cinara todocola (Inouye) Apterae have a brown head and dark green to greenish-black abdomen, lightly dusted with wax (Inouye 1936); BL 2.6-3.0 mm. On trunk and branches of young todo-fir trees (Abies spp.) in Japan and Sakhalin. Often in ant shelters. Injurious. Oviparae and alate males occur in September-November (Inouye 1970). Yamaguchi (1976) studied ecology and morph determination, and Furuta (1984) studied the impact of natural enemies. C. smaragdina is a similar species on Abies spp. in Siberia, but apparently with a different biology as it feeds on shoots between needle bases (Pashchenko 1988a).

Cinara tonaluca Hottes & Wehrle Apterae are dusted with white wax, with a black stripe and two longitudinal rows of black spots showing through the wax (Hottes 1952b); BL 2.1-2.6 mm. On woody stems of young trees of Juniperus monosperma in Arizona USA. Not recorded since the original description (Hottes & Wehrle 1951a), sexuales and life cycle are unknown.

Cinara tsugae Bradley Apterae have the abdomen chestnut-brown, head and thorax somewhat lighter brown; BL 2.9-3.6 mm. In dense spring colonies on branches of Tsuga heterophylla, and later on roots or lower part of trunk in ant shelters (Bradley 1960). In British Columbia, Canada and Oregon USA (BMNH collection, leg. Bradley). Oviparae deposit eggs on needles in autumn, males are unrecorded. C. wahsugae, described from Tsuga mertensiana in Oregon (Hottes 1960a), may be a synonym, although oviparae and alate males of wahsugae were collected in August (and see Hottes 1961c).

Cinara (Cupressobium) tujafilina (Del Guercio) Apterae are reddish-brown with a dorsal pattern of bluish-white wax, and two dark brown divergent curved bands running from head to about level of siphunculi (see aphids of Karnataka website and/or influentialpoints.com/Gallery); BL 1.7-3.5 mm. On Cupressaceae: Callitris, Chamaecyparis, Cupressus, Juniperus, Libocedrus, Thuja (usually orientalis), Thujopsis and Widdringtonia. On foliated branches, undersides of branches near the trunk, on wound tissue, or in midsummer on roots (Bray 1953, Colombo & Parisini 1984). Virtually cosmopolitan (but not in northern Europe), including many warmer regions. Apparently almost entirely anholocyclic, but an ovipara has now been recorded from Iran (Remaudiėre & Binazzi 2003b), Zhuravlev (2003) described the (alate) male from Kirghizia, and males were recently reared in Poland under artificial conditions and redescribed by Durak & Durak (2015). Overwintering generations occur on the roots (Durak 2014b). Durak & Borowiak-Sobkowiak (2007) studied development and found that there were only three larval instars, a feature shared with C. cupressivora and possibly other Cinara species not yet studied in this respect. There are population studies by Furuta (1988; Japan) and Mohammed et al. (1988; Iraq). Similar but shorter-haired specimens in south-west Asia may be referable to C. mediterraneum Narzikulov. 2n=12.

Cinara vagabunda Hottes & Essig Appearance in life is unknown; BL of aptera c. 3.6 mm. On Pseudotsuga menziesii in Arizona and New Mexico, USA (Hottes 1961d). See C. pseudotaxifoliae.

Cinara vandykei (Wilson) Apterae are dark brown to black with little or no wax: BL 1.8-3.0 mm. In colonies on bark of twigs of Picea spp. in western USA (incl. California and Oregon: BMNH collection), and across Canada. Oviparae and apterous males are produced early, in July-August (Palmer 1926). C. wanepae Hottes, described from Picea pungens in Colorado, is a probable synonym (Palmer 1936b).

Cinara villosa Gillette & Palmer Apterae are described as indistinguishable in life from those of C. apini (Gillette & Palmer 1924), and therefore presumably dark brown, mottled with wax powder; BL c. 3.2-4.0 mm. On bark of twigs of Pinus flexilis in western North America. Originally described as a long-haired form of apini in Colorado, and possibly the two are not specifically distinct. Hottes (1955b) described two variant populations of C. villosa from P. flexilis var. reflexa in Arizona as subspecies.

Cinara wahhaka Hottes Apterae are shining dark brown without wax; BL 1.5-1.8 mm. On foliage of small twigs of Juniperus osteosperma (= utahensis), only known from Colorado USA. All the second generation were alate (Hottes 1952c). Oviparae and apterous males (Hottes 1953b) occur in September-October.

Cinara wahluca Hottes Apterae are reddish-brown without wax; BL 2.7-3.0 mm. On bark of trunk and larger branches of Juniperus scopulorum, under scales or in crevices or wounds. In western USA (Colorado, Wyoming); the record from Turkey (Akyürek et al. 2010) is almost certainly a misidentification. Oviparae and apterous males occur in September (Hottes 1952a).

Cinara wahtolca Hottes Apterae are brownish to dark grey with black marking and black siphunculi, usually extensively patterned with greyish white powder; BL 2.6-4.6 mm. In small to large colonies on trunk and needle-free or sparsely-needled limbs of pinyon pines (Pinus edulis, P. cembroides, P. discolor) in south-western USA. Populations on P. monophylla previously regarded as C. wahtolca are referable to C. anelia, although hybrids between P. edulis and P. monophylla may be colonised by C. wahtolca (Favret & Voegtlin 2004b). Oviparae and alate males occur in October (Hottes 1954b, 1956e).

Cinara watanabei Inouye Apterae have head and thorax dark brown, and abdomen shiny brown; BL 4.8-5.4 mm. In large colonies on 3- to 5-year-old wood of branches of older trees, or on trunk of young trees, of Pinus spp. (e.g. koraiensis, parviflora) in Japan, Korea and Sakhalin (Pashchenko 1988a). A record from Turkey (Akyürek et al. 2011) is likely to be a misidentification. Oviparae and alate males occur in Japan in October-November (Inouye 1970). [C. stekolshchikovi Mamontova, collected on a Pinus sp. in east Siberia (Primorskiy Kray), is described as similar but with a different pattern of dorsal sclerotisation, shorter antennal hairs and smaller siphuncular cones (Mamontova 2012)].

Cinara watsoni Tissot (fig. 75H) Apterae have head and thorax dark chestnut brown and abdomen cinnamon brown with dark brown markings, the whole body dusted with greyish-white wax (Tissot 1939); BL 2.7-4.3 mm. In small loose groups on new shoots, or near ends of previous year’s growth, of Pinus spp. of subsections Australes and Contortae in eastern USA. Records from P. radiata in California (Jousselin et al. 2013) require confirmation, as do records from unidentified Pinus spp. in Central America (Villalobos Muller et al. 2010), and from P. sylvestris in China (G. Zhang et al. 1993b, G. Zhang 1999). Oviparae and alate males occur in October-November (Pepper & Tissot 1973). Population trends of C. watsoni in South Carolina were studied by Brooks & Warren (1964), and by Patti & Fox (1981).

Cinara westi Tissot & Pepper Apterae are dark brown; BL 2.1-2.8 mm. On bark of Pinus taeda in south western USA, often associated with lesions of the rust Cronartium fusiforme, and usually in ant (Crematogaster) shelters (Tissot & Pepper 1967). Sexuales and life cycle are unknown. This species is possibly synonymous with C. rigidae.

Cinara xylophila Zhang & Zhang BL of alata c. 4.2 mm, aptera undescribed. On Pinus armandii in Sichuan, China (G. Zhang et al. 1993b).

Cinara zoarcbursara Knowlton Appearance in life is unknown; BL of aptera c. 3.5 mm. Only known from apterae collected on Abies concolor in Utah USA (Knowlton 1935). Biology is unknown.

Clethrobius Mordvilko

Calaphidinae: Calaphidini

Three or four palaearctic species of large (2.8-5.6 mm) hairy brown aphids living on twigs and young branches of Alnus and Betula species. All adult viviparae and males are alate, the ovipara being the only apterous morph. Accounts are available for Europe (Heie 1982, Stroyan 1977, Wood-Baker 1983), India (A.K. Ghosh & Quednau 1990) and China (Qiao et al. 2005b).

Clethrobius comes Walker Alate viviparae are dark brown (see influentialpoints.com/Gallery), forming clusters on branches and twigs of Betula spp., often where the new growth is dying back, or on twigs of Alnus spp. overhanging streams. The populations on Alnus are regarded by some authors as a separate species, C. giganteus Cholodkovsky, and attempts to transfer aphids from birch to alder have not succeeded (Wood-Baker 1983). No consistent morphological differences have been found, however, and both birch and alder populations share the same peculiar structurally heterozygous karyotype (2n=11; Blackman 1988). C. comes occurs throughout Europe and across Asia to China, Korea and Japan. Holocyclic, with sexuales in October-November. The male genitalia were described and illustrated by Wieczorek et al. (2011).

Clethrobius dryobius Chakrabarti & Raychaudhuri Brown aphids with many more hairs on the dorsal surface of the body, and the last rostral segments, than C. comes, and also with some hairs on the siphunculi. Holocyclic on Betula alnoides and B. utilis in north-east India. with sexuales in October-November (Chakrabarti 1988, Chakrabarti et al. 1988, A.K. Ghosh & Quednau 1990). It is also recorded from China (Qiao et al. 2005b).

Clethrobius vermai Ghosh & Quednau Colour in life is not recorded. Described from a single alata (BL 3.3 mm) from Betula sp. in Himachal Pradesh, India (A.K. Ghosh & Quednau 1990).

Clydesmithia Danielsson

Eriosomatinae: Pemphigini

A genus with one nearctic species related to Pachypappa and Gootiella. The fundatrices lack wax glands and have antennae with only 4 segments.

Clydesmithia canadensis Danielsson Gall on Populus (balsamifera, trichocarpa) consists of an elongate slightly wrinkled swelling on the distal half of the leaf lamina, 1.0-1.5 cm long, tinged with red, opening on underside of leaf. Apterae in galls are yellowish with a thin layer of wax; BL 1.5-2.7 mm. In north-west USA and Canada. Living without host alternation on Populus; first instars of the third generation in the galls migrate, possibly by falling from the leaves, to the roots of the host (Aoki et al. 1997a). Presumably sexuparae develop in the root-feeding colonies and move back to the trunk to produce sexual morphs as in other Eriosomatini. It now appears, however, that some populations may host-alternate, as Pike et al. (2012) reported populations associated with conifer roots and mosses in Alaska, Montana and Washington, and described the apterous exule. No alate morphs have been described. Further work is needed to resolve these discrepancies.

Clypeoaphis Soliman

Aphidinae: Macrosiphini

One or two species on Amaranthaceae, with a swollen clypeus like that of Brachyunguis feeding on this plant family, but the absence of marginal abdominal tubercles and placement of spiracles on anterior abdominal segments suggest that this genus belongs in Macrosiphini (Remaudière & Bahamondes 1987).

Clypeoaphis (?) stavropolensis Ivanoskaya Apterae are yellowish brown; BL c.1.9 mm. On Salsola sp. in west Siberia. [The generic position of this species is uncertain; it was described as having small marginal tubercles on prothorax and abdominal tergites 1 and 7, but other features (length of antennal terminal process, number of hairs on R IV+V and HT I, shape of siphunculi and cauda) indicate a closer affinity to Clypeoaphis suaedae than to Brachyunguis (Xerophilaphis).]

Clypeoaphis suaedae (Mimeur) Plate 15c (Fig.36d,i) Apterae are pale olive-green or yellowish green, covered with greyish white wax powder (see influential points/gallery); BL 1.2-1.4 mm. In small colonies or scattered on stems of Suaeda spp. in saltmarshes and similar habitats. Also recorded from Kochia scoparia and Salsola komarovii. Alatae are rare (Gimingham 1942). In Europe, North Africa, Middle East, Central Asia and Korea. Monoecious holocyclic, with oviparae and apterous males in September (BMNH collection, leg. H.L.G. Stroyan). A population from the Netherlands with shorter and blunter hairs on head and abdominal tergite 8 is regarded as a subspecies, C. suaedae ssp. suaedicola Hille Ris Lambers.

Codonopsimyzus Lee

Aphidinae: Macrosiphini

One species in east Asia perhaps related to Cryptomyzus but with very short dorsal hairs.

Codonopsimyzus sasammi Lee Apterae are milky white except for black antennal joints and tarsi; BL 1.6-2.0 mm. Alatae have antennae, legs and siphunculi black, secondary rhinaria distributed III 34-55, IV 14-26, V 5-16, and a black quadrate dorsal abdominal patch. On undersides of leaves of Codonopsis lanceolata in Korea (Lee 2002a). The life cycle is unknown.

Colopha Monell

Eriosomatinae: Eriosomatini

Six northern hemisphere species related to Kaltenbachiella, but alatae have a branched media in the fore wing and one oblique vein in the hind wing. Akimoto (1985) and Smith (1985) reviewed the genus. Three species are known to migrate from cockscomb-shaped galls on Ulmus to grasses or sedges, where they produce colonies with abundant wax on the aerial parts and/or roots. For an account of the two European species see Nieto Nafría et al. (2002a). Life cycle variation in this genus in relation to distribution was discussed by Sano & Akimoto (2005). A cladistic analysis by Sano & Akimoto (2011) including four Colopha species indicated that the genus is paraphyletic and basal to Kaltenbachiella and Tetraneura.

Colopha compressa (Koch) Plate 1a (Fig.20i) Galls are formed on the upper surfaces of leaves of Ulmus spp., usually U. laevis, often near the mid-rib; they are laterally compressed, pouch- or short cockscomb-shaped, yellowish, often tinged with red (fig. 134B). Alatae (BL 1.4-1.8 mm) (fig. 117D) emerge in July from an opening on the underside of the leaf and found colonies on roots of Carex and Eriophorum, sometimes in ants’ nests. Apterae on secondary hosts are dark yellowish, secreting flocculent wax; BL 0.9-1.5 mm. Sexuparae return to elm in September-October, and anholocyclic populations may also persist on roots of Cyperaceae through the winter months (Marchal 1933, Zwölfer 1957). Throughout Europe (including Iceland), east to Turkey, Iran, Ukraine and Kazakhstan (Kadyrbekov & Aoitzhanova 2005), and introduced into towns in Siberia (Heie 1980b). 2n=16.

Colopha graminis (Monell) Forming cockscomb-like galls on upper surfaces of leaves of Ulmus spp. (americana, rubra) on the leaf lamina between the veins (Hottes & Frison 1931 and see influential points/gallery). Alatae (BL 1.0-1.6 mm) emerge in June-July, migrating to found colonies on roots, stems and leaves of Poaceae, e.g. Aira caespitosa and Agrostis plumosa, from which it was originally described (Monell 1882), and Leersia virginica (Patch, 1910b). An attempted transfer to Eragrostis was unsuccessful (Patch 1910b, cf. ulmicola). Apterae on secondary hosts live in grey woolly wax (see influential points/gallery), and have BL 1.2-1.8 mm. Sexuparae develop to adult on the leaves of the secondary host, returning to elm in October (Patch 1910b). Widely distributed in North America. Some authors (e.g. Palmer 1952) have synonymised graminis with ulmicola, and apterae on secondary hosts cannot be separated, but Smith (1985) considered that there was sufficient evidence to regard the two species as distinct.

Colopha hispanica Nieto Nafría & Mier Durante Apterae are dirty yellowish grey to very pale greyish brown covered with white wax powder; BL 1.4-2.7 mm. Living endophytically within stems of Scirpoides holoschoenus (the only known truly endophytic aphid). In Spain and France (Nieto Nafría et al. 2002a). Apparently anholocyclic.

Colopha kansugei (Uye) (Fig.20g) Apterae in grey woolly wax; BL 1.2-1.6 mm. In conspicuous colonies on aerial parts of Carex spp. and unidentified grasses; in spring and summer it may also occur on roots (Akimoto 1985). In Japan, Ryuku Islands, Taiwan, China, and Nepal. Apparently mainly anholocyclic on Carex; sexual generations are unknown, although alate sexuparae are produced in autumn in Japan.

Colopha setaricola Sano & Akimoto Apterae are densely covered in white wax; BL 1.4-1.9 mm. On aerial parts of Setaria chondrachne in central Japan (Sano & Akimoto 2005). Alate sexuparae are produced in November, but no primary host generations have been found and it is believed to be entirely anholocyclic.

Colopha ulmicola (Fitch) Forming elongate cockscomb-like galls, pale green tinged with brown, apparently indistinguishable from those of C. graminis, on leaves of Ulmus spp. (americana, rubra, thomasii; fig. 134C). Alatae (BL 1.0-1.6 mm) emerge in June-July and fly to Eragrostis spp., giving rise to colonies with flocculent wax on the upper leaves and stems. Possibly other Poaceae are also colonised, including Panicum and Zea (but not Leersia? – see Patch 1910b). Apterae on secondary hosts are reddish brown, BL 1.2-1.8 mm. Sexuparae return to elm in September-November (Patch 1910b). It is apparently widely distributed in North America, but records may be confused with those of C. graminis.

Colophina Börner

Eriosomatinae: Eriosomatini

Five or six east Asian species related to and morphologically resembling Eriosoma but biologically distinct (Akimoto 1983), with particularly complex life cycles and polymorphism, including sterile first instar soldiers on the secondary host (Aoki 1977). Two species are known to migrate from globular galls on Zelkova to the aerial parts of Clematis, and galls of a third have been experimentally induced on Zelkova. Two are only known from Clematis. The Chinese species were reviewed by G. Zhang et al. (1999c).

Colophina arctica Zhang & Qiao Apterae are presumably covered in wax wool; BL 2.0-3.3 mm. On Clematis brevicaudata in Beijing City, China (G. Zhang & Qiao 1997b, G. Zhang et al. 1999c). Primary host is unknown.

Colophina arma Aoki The galls are globular, loosely closed, on the margins of leaves of Zelkova serrata in Japan (fig. 125B). Alatae (BL 2.2-2.4 mm) emerge in July and fly to Clematis stans. where dense colonies (including sterile 1st instar soldiers) are formed on stems or peduncles. Apterae on Clematis are dark brown covered in white wax-wool; BL 2.4-3.3 mm. They produce first instar larvae of three types, normal, “midget” and soldier (Aoki 1977). Sexuparae are produced in autumn and presumably return to Zelkova, but overwintering also occurs as midget 1st instar larvae on the secondary host; the midget larvae hibernate under bark of lignified stems, providing an alternative life cycle (Aoki 1980b). Differentiation of soldiers and factors affecting their production were studied by Ijichi et al. (2005a,b). 2n (female) =10, 2n (male) =8 (Blackman 1986).

Colophina clematicola (Shinji) Apterae are very small, dark brick-brown, secreting white wax-wool; BL 0.8-1.3 mm. On stems, leaves and peduncles of Clematis spp. in Japan and Korea, and also collected from Clematis sp. in New Zealand (BMNH collection, leg. C. Butcher). In Japan it has been found to colonise several non-native Clematis spp. (Aoki & Kurosu 2016). In autumn colonies become large and produce alate sexuparae, which presumably migrate to Zelkova, on which galls have been artifically induced after transfer of sexuparae collected the previous year on C. terniflora (Aoki & Kurosu 2000), although not yet found in nature. The galls were of leaf-roll type, the roll tinged with yellow. Fundatrices, apterae and emigrant alatae from the galls were described. However, overwintering of first and second instar nymphs also occurs on C. terniflora in Japan (Aoki et al. 1997b). Soldier-like nymphs were reported by Akimoto (1998a). 2n=20.

Colophina clematis (Shinji) The galls on Zelkova serrata in Japan appear identical to those of C. arma (see above). Alatae (BL c. 2.2 mm) (fig. 120A) emerge in July and fly to Clematis (apiifolia, gouriana, trichotoma), where dense colonies (including soldiers) are formed on the stems, usually in shade near the ground (Aoki 1977). They are also recorded from Korea and Taiwan. In Japan it a large colony was found on the non-native Clematis × jouiniana (Aoki & Kurosu 2016). Apterae on Clematis are dark brown covered in white wax-wool; BL 2.2-2.8 mm. Alate sexuparae return to Z. serrata in October, but first instar larvae also overwinter in bark crevices on the secondary host (Aoki 1980b). 2n(female)=11, 2n(male)=9 (Blackman 1986).

Colophina monstrifica Aoki Apterae are presumably brown covered in white wax-wool; BL 3.6-5.4 mm. In large colonies on stems of Clematis uncinata (= C. floribunda (Hayata) Yamamoto) in mountainous regions of Taiwan (Aoki 1983). Large first instar pseudoscorpion-like soldiers are produced. Alate sexuparae are produced in autumn. The primary host has now been identified as Zelkova serrata; pale green globular galls were obtained by experimental transfer of eggs on Clematis twigs, and a few naturally formed galls were also found (Uematsu et al. 2021). Identity of the inhabitants of the galls was confirmed by mitochondrial DNA sequencing. Alatae from the galls have not been described.

Colophina zelkovae Lee, Seo & Hwang Emigrant alatae (BL 1.5-1.9 mm) were collected from pouch galls on leaves of Zelkova serrata in Korea, flying in June-July to an unknown secondary host (Lee et al. 1993). Possibly these are small specimens of C. clematis.

Coloradoa Wilson

Aphidinae: Macrosiphini

Despite its name this is an Old World genus, of about 30 species living monoeciously on Anthemidae (Asteraceae). Characteristic features are the spatulate body hairs shaped like partly opened fans, the stiletto-shaped R IV+V and the reduction and ventral displacement of the ocular tubercles, so that they are often not evident in dorsal view. There are reviews for UK and Ireland (Blackman 2010), north-west Europe (Heie 1992), central Europe (Heinze (1960) and Central Asia (Kadyrbekov 2004a, 2020). Kadyrbekov (2020) provided a key to apterae of all known species. As noted in the brief accounts of individual species below, further studies may reveal a number of synonymies.

Colorodoa abrotani (Koch) (Fig.12h) Apterae are greenish with siphunculi and cauda yellowish brown; BL 1.2-1.6 mm. On leaves of Artemisia abrotanum and also recorded from A. afra. In northern Europe (Germany, Sweden), also reported to occur in Iran (Rezwani 2010), and introduced to Canada (New Brunswick). Sexual morphs are apparently unknown.

Coloradoa absinthii (Lichtenstein) (Fig.12m) Apterae are greenish with tips of antennae and tarsi black; BL 1.7-2.0 mm. On undersides of lower leaves of Artemisia absinthium, also A. abrotanum. Widely distributed in Europe, Iran (Rezwani & Parvizi 1990), eastern Kazakstan (Kadyrbekov 2009a), and introduced to North America (New York, Manitoba, Washington). Oviparae occur in October, males are undescribed.

Coloradoa achilleae Hille Ris Lambers (Fig.5a) Apterae are pale green to greyish green or reddish (see influentialpoints.com/Gallery); BL 1.1-1.6 mm. On leaves of Achillea millefolium. Europe, eastward to Russia and Turkey (Jörg & Lampel 1988), Iran (Mokhtari et al. 2012), Kazakhstan (Kadyrbekov 2017a), and introduced to USA (North Carolina; Halbert et al. 2000). Oviparae and apterous males occur in late September-October.

Coloradoa ajaniae Kadyrbekov Apterae are pale greenish with red eyes, with appendages only darker at apices; BL 1.2-1.5 mm. On upper sides of leaves of Ajania fastigiata in Kazakhstan (Kadyrbekov 2020). Other morphs and biology are unknown.

Coloradoa angelicae (del Guercio) (= absinthiella Ossiannilsson) (Fig.12n) Apterae are green, with tips of antennae, legs and siphunculi, and entire cauda brownish; BL 1.2-1.8 mm. On Artemisia absinthium (the original host plant was probably misidentified), often occurring with C. absinthii. Throughout Europe and eastward to Pakistan, and introduced to North America (Montana, New York, California) and Argentina (Ortego 1998a).

Colorodoa artemisiae (del Guercio) (Fig.12p) Apterae are dirty green or reddish, with black tips of antennae and tarsi; BL 1.3-1.8. On Artemisia spp., most usually A. vulgaris, on the upper parts of the plant. Throughout Europe, Iran (Rezwani 2010), India (Bhutan), Nepal, and introduced to USA and Canada, and to Argentina (Ortego 1998a). Oviparae occur in September (Heie 1992), males are undescribed. 2n=16

Coloradoa bournieri Remaudière & Leclant Apterae are pale or dirty green, with dark apices to appendages (see influential points/gallery); BL 1.1-1.5 mm. On Santolina chamaecyparissius, S. viridis and Santolina sp. Europe (England, France, Spain, Italy), and Middle East (Israel, Iran, Lebanon; BMNH collection). It has recently been found in South America (Argentina: Mier Durante et al. 2011). Anholocyclic where studied (France, original description), and probably elsewhere. A Spanish population was described as a subspecies, C. bournieri ssp. iberica, but might be a variant clone. 2n=22.

Coloradoa brevipilosa Ivanoskaya (= C. huculaki Szelegiewicz; synonymy by Kadyrbekov 2020). Apterae have dark green head and thorax, and yellowish green to green abdomen, with dark apices to antennae and legs, green to brownish green siphunculi and pale yellowish green cauda; BL 1.3-1.6 mm. Described from Artemisia sieversiana in Siberia, on A. pontica in Hungary, and also in Poland (Wojciechowski et al. 2015, as C. huculaki) and China (BMNH collection). It is also recorded (from A. absinthium) in Kazakhstan (Kadyrbekov 2018, as C. huculaki). Sexual morphs are unknown. 2n = c.24 (based on immature specimen from China).

Coloradoa brevisiphon Bozhko Apterae are green with whitish legs, siphunculi and cauda: BL c.1.3 mm. Living well-concealed on flower stems of Artemisia procera (= abrotanum) in Ukraine (Bozhko 1976b), reported from the same host in south-west Russia (Kadyrbekov 2014g), and also from A. pontica in Kazakhstan (Kadyrbekov 2017a). Sexual morphs are unknown.

Coloradoa campestrella Ossiannilsson Apterae are pale yellow to green, with brown tips of antennae, tarsi, and (sometimes) siphunculi; BL 1.3-1.6 mm. On Artemisia spp. in Europe, Kazakhstan, Pakistan and also in Japan, China and Korea (as artemisiae ssp. artemisicola). Oviparae are described (Ossialnnilsson 1959), but not the male.

Coloradoa campestris (Börner) (Fig.12i) Apterae are green with blackish brown siphunculi and cauda; BL 1.1-1.3 mm. On Artemisia spp. in Europe; Jorg & Lampel (1988) discussed its habitat and distribution. Also recorded from Kazakhstan (Kadyrbekov 2016d) and Pakistan (Naumenn-Etienne & Remaudière 1995). Oviparae are figured by Heinze (1960), the male is undescribed.

Coloradoa deserta Kadyrbekov Apterae are pale greenish with red eyes; BL c.1.1 mm. Alatae are undescribed. On Artemisia santolina in the Xinjiang-Uygur region of north-west China (Kadyrbekov 2004a).

Coloradoa heinzei (Börner) Apterae are green or reddish, with apical parts of antennae, legs and siphunculi dark; BL 0.8-1.6 mm. On upper parts of Seriphidium (= Artemisia) maritimum in Europe, and there are Asian records from many other Artemisia and Seriphidium species. Apterae collected on S. aucheri in Iran were very small (BL 0.8-0.9 mm) and had relatively short siphunculi (Mehrparvar & Rezwani 2007). In Europe, and eastward to Kazakhstan and Pakistan. Stroyan (1979) gave a full redescription including oviparae and apterous males collected in UK in September.

Coloradoa indica Verma Appearance in life is unknown; BL of aptera 0.65-0.9 mm. Described from an unidentified Artemisia sp. in Kashmir; later collected from A. sibirica (= Filifolium sibiricum) in Kashmir (BMNH collection, leg. N.D. Rishi), and also in China (leg. V.F.Eastop). Probably close to C. ponticae, which it resembles apart from its larger size and 6-segmented antennae. Sexual morphs are unknown.

Coloradoa inodorella Ossiannilsson Apterae are green, with appendages often dark distally; BL 1.3-1.7 mm. On leaves of Tripleurospermum inodorum in north-west and central Europe, eastward to Russia and Turkey. Also found on Matricaria discoidea. Oviparae and apterous males were described by Ossiannilsson (1959).

Coloradoa mesasiatica Kadyrbekov Apterae are pale greenish with red eyes; BL 1.0-1.5 mm. On upper sides of leaves of an Artemisia sp. in Kazakhstan (Kadyrbekov 2004a), and subsequently recorded from Seriphidium heptapotamicum (Kadyrbekov 2014d). Close to C. absinthii.

Coloradoa moralesi Remaudière & Leclant Apterae are pale green with apices of appendages darker; BL 1.1-1.6 mm. On Santolina spp. in Spain (Sierra de Guadarrama).

Coloradoa palmerae (Börner) Apterae are dirty green; BL 1.4-1.7 mm. On Artemisia alba, living inconspicuously on the margins of the finely divided leaves. Alatoid nymphs are strikingly brownish yellow. Hille Ris Lambers (1967) redescribed this aphid and discussed differences from C. absinthii. Found in France, Italy and Hungary. Sexual morphs are unknown.

Coloradoa paradoxa Szelegiewicz Apterae are pale green with antennae, legs and siphunculi brownish green, and cauda dirty green to yellowish; BL 1.2-1.5 mm. On unidentified Artemisia sp(p). in Turkey, Uzbekistan, Pakistan and Mongolia. Specimens in the BMNH collection from Turkey contain parasitoid larvae. Originally described from Mongolia as a subspecies of C. heinzei. C. paradoxa could be the fundatrix generation of a species that has 6-segmented antennae and longer body hairs in later generations.

Coloradoa ponticae (Börner) (Fig.12k) Appearance in life is unknown; BL 1.0-1.5 mm. On Artemisia pontica in Czech Republic (Heinze 1960), and also found on an unidentified Artemisia in China (BMNH collection, leg. V.F. Eastop). Probably close to C. indica (q.v.). 2n=16.

Coloradoa procerae (Bozhko) Apterae are green; BL c.1.1 mm. On Artemisia scoparia in Ukrainian steppes (Bozhko 1976b; original description from A. procera (= abrotanum) was presumably in error?). Possibly this is C. viridis.

Coloradoa rufomaculata (Wilson) Plate 15a (Fig.12o, Fig.52c) Apterae are pale green with pale brown head, with siphunculi distally dark and often with white patches at their bases (see aphids of Karnataka website); BL 1.0-1.7 mm. On stems and undersides of leaves of cultivated chrysanthemums (Dendranthema), Chrysanthemum coronarium, and sometimes on other compositae genera (Ambrosia, Artemisia, Pentzia, Siegesbeckia, Tanacetum). Throughout the world, occurring mainly in glasshouses in cold temperate regions. Probably anholocyclic almost everywhere, although oviparae are known from Australia and New Zealand. Hussain et al. (2015b) discussed morphological variation between populations from different parts of the world. 2n = 8 or 17 ( Panigrahy & Patnaik 1991, both samples from Chrysanthemum coronarium) or 18 (P.L. Das et al. 1985); it is very unlikely that 8 and 17/18 would be found in the same species, so two species are probably being confused under the one name.

Coloradoa santolinae Hille Ris Lambers (Fig.5b) Apterae are green, BL 1.2-1.4 mm. Described from Achillea santolina in Israel, and subsequently also recorded from this host in Iran and Afghanistan, from A. crithmifolia in Bulgaria (Holman 2009)and from Artemisia monosperma in Israel (BMNH collection). Able to transmit cucumber mosaic poytyvirus (strain 2 ; B. Raccah, pers. comm.). Sexual morphs unknown. 2n=20 (specimens from A. monosperma).

Coloradoa scopariae Szelegiewicz Apterae are yellowish green with brownish antennae and legs, brownish yellow siphunculi and dirty yellow to green cauda; BL 1.1-1.4 mm. On Artemisia scoparia in Mongolia. Sexual morphs are unknown.

Coloradoa submissa Doncaster (Fig.12j) Appearance of apterae in life is unknown; oviparae found in October in Denmark were dirty yellowish or greenish brown, wax powdered, with head, pronotum, siphunculi and cauda brown-black (Heie 1992). On Artemisia maritima in UK and Denmark, and also collected from Artemisia caerulesens ssp. gallica in France, on Artemisia sp. (cf. brevifolia) in Pakistan (Naumann-Etienne & Remaudière 1995), and from A. capillaris in Korea (BMNH collection, leg W.H. Paik). Oviparae and alate males in England in September (BMNH collection, leg. H.L.G. Stroyan).

Coloradoa tadzhica Narzikulov Apterae are green, sometimes tinged with yellowish-pink; BL c.1.3 mm. On Artemisia ferganensis in Tajikistan (Narzikulov 1958, as Neaphis artemisiae). Sexual morphs are unknown. Probably this is a synonym of C. heinzei (Kadyrbekov 2004a).

Coloradoa tanacetina (Walker) (Fig.52d,e) Apterae are yellowish green, with tips of antennae and tarsi dark (see influentialpoints.com/Gallery); BL 1.1-2.0 mm. On Tanacetum vulgare, feeding on edges of leaves, and also recorded from Dendranthema spp. (Ossiannilsson 1959) and Leucanthemum vulgare (BMNH collection, leg. H.L.G. Stroyan). In Europe, Central Asia (Kazakhstan; Kadyrbekov 2016d), and introduced to USA (Halbert et al. 2000 and aphidtrek.org) and Argentina (Ortego 1998a). Oviparae and apterous males occur in September.

Coloradoa taurica (Mamontova-Solukha) Apterae are brown; BL c.1.1-1.2 mm. On Artemisia and Seriphidium spp. in Crimea, south-west Russia and Kazakhstan, and also in Afghanistan (BMNH collection, leg. R. van den Bosch). Sexual morphs are unknown.

Coloradoa viridis (Nevsky) (= C. nodulosa Zhang; synonymy by Kadyrbekov 2020) (Fig.12l) Apterae are green; BL 1.0-1.6 mm. On Artemisia spp. in Iran, Central Asia, Mongolia (Szelegiewicz 1963) and China (BMNH collection, and Zhang & Zhong 1980a, as C. nodulosa). Sexual morphs are unknown. C. kondoi (Shinji) might be this species (as also might C. procerae, q.v.). 2n= 16.

Cornaphis Gillette

Eriosomatinae: Pemphigini

A genus for one North American species living without host alternation on Populus, characterised by a medial frontal projection that forms a distinct horn in apterae and a raised base for the median ocellus in alatae.

Cornaphis populi Gillette Galls on Populus angustifolia are crescent-shaped folds of the leaf-edge, pale or streaked with red. In western USA (Wyoming, Colorado, Utah). Apterae inhabit the same galls as the slate-grey fundatrices, and are straw-yellow with black head and four dark patches on the pronotum, dusted with wax. There is no host alternation; alate sexuparae (BL 2.0-2.3 mm) are produced in the galls in July (Gillette 1913, Palmer 1952).

Corylobium Mordvilko

Aphidinae: Macrosiphini

A genus for one Macrosiphum-like species with capitate hairs living on Corylus. Hille Ris Lambers (1947a) discusses its affinities.

Corylobium avellanae (Schrank) Plates 14 c,d Apterae are green, sometimes reddish, spindle-shaped (see influentialpoints.com/Gallery); BL 1.7-2.7 mm. On shoot apices, petioles and undersides of young leaves of Corylus avellana; dense colonies form particularly on suckers. In Europe eastward to Turkey and Ukraine, Iran, and introduced into Canada (British Columbia), where it was also found on C. cornuta and C. maxima (Forbes & Chan 1984). Monoecious holocyclic, with alate males. Natural enemies, especially the impact of a pathogen, were discussed by Viggiano in Cavallaro (1983), and Tremblay & Pennachio (1985) reported a parasitoid (Praon dorsale). See also Hille Ris Lambers (1947). 2n=10.

Cranaphis Takahashi

Calaphidinae: Panaphidini

A little-known oriental genus of two species perhaps most closely related to Tinocalloides (Quednau 2003), but specialised for feeding on bamboos (Arundinaria and related genera).

Cranaphis formosana (Takahashi) All viviparae are alate, long-bodied, yellow with black antennae and a broad medial dusky stripe on head and prothorax, and paired dusky-dark markings on most abdominal tergites (Takahashi 1924, Quednau 2003); BL 1.5-2.2 mm. On undersides of leaves of Yushania niitakayamensis in the central montains of Taiwan (Liao 1976), and also reported from mainland China (Qiao et al. 2005b). Sexuales are unknown; it is probably anholocyclic in Taiwan.

Cranaphis indica Chakrabarti & Raychaudhuri (1976) Described from a single alata and immatures collected on Arundinaria sp. in West Bengal, India. Quednau (2003) provided detailed drawings of the type specimens.

Cryptaphis Hille Ris Lambers

Aphidinae: Macrosiphini

This genus includes one palaearctic and one nearctic species living concealed on the basal parts of Poaceae, and resembling Metopolophium except for their long, capitate hairs. The other five Asian species currently placed in this genus from Lamiaceae and Geraniaceae are generally similar to the grass feeders but have spinulose heads, and are perhaps closer to Aulacorthum (Perillaphis).

Cryptaphis bromi Robinson Apterae are shining brown; BL 1.2-1.6 mm. Alatae have broad dark dorsal abdominal cross-bands. On Bromus inermis, feeding cryptically on brown stems at bases of plants. Canada (Manitoba). Monoecious holocyclic, with oviparae in November 2n=16.

Cryptaphis garwahlensis Bhattacharya, Mandal & Chakrabarti Colour of apterae in life is unrecorded; BL 2.1-2.4 mm. On Lamium album, causing curling and blistering of leaves. In Uttar Pradesh, India.

Cryptaphis geranicola (Shinji) Apterae are yellowish brown to blackish, BL 1.6-2.1 mm. Immatures are orange. Alatae have a solid dark dorsal abdominal patch (Miyazaki 1971). On undersides of leaves of Geranium spp., causing heavy leaf-curl. Monoecious holocyclic in Japan, and also recorded from Tajikistan (Narzikulov 1975), east Siberia (Pashchenko 1988a), Korea (Choi et al. 2011) and China (Su et al. 2011). 2n=14.

Cryptaphis menthae Takahashi Apterae are blackish with mainly dark appendages; BL c.2.1 mm. Alatae have broad dark abdominal cross-bands and large marginal sclerites. On Lamiaceae (Isodon, Mentha, Plectranthus), “crisping” leaves, in Japan, and also now recorded from Korea (Choi et al. 2011). Monoecious holocyclic; fundatrices were found on Isodon inflexus (Miyazaki 1971), and oviparae and alate males were found on the same host in October in Korea by Choi et al. (2011- but note that there are errors in the morphometric data tabulated by these authors).

Cryptaphis poae (Hardy) Plate 24c Apterae are very shiny, green mottled with red or yellowish with brown markings, to brownish black; BL 1.3-2.0 mm. Alatae have broad dark dorsal abdominal partly fused cross-bands. In small colonies at bases of various grasses (e.g. Festuca, Holcus, Poa) just below soil level, or under stones, usually in shady, damp situations. Widespread in Europe. Monoecious holocyclic, with oviparae and apterous males in October. 2n=20.

Cryptaphis rostrata Chakrabarti & Raychaudhuri Apterae are pale brown (in life?); BL 1.2-1.5 mm. Alatae have broad dark abdominal cross-bands and large dark marginal sclerites (BMNH collection, leg. S. Chakrabarti). Described from an unidentified labiate plant, and subsequently found on Isodon (= Plectranthus) rugosus in India (Himachal Pradesh) and Pakistan.

Cryptaphis salviae Bhattacharya & Dey Colour of aptera in life is unrecorded; BL 1.4-1.5 mm. Alatae have cross-bands which on ABD TERG 4-6 are merged into a solid patch. On Salvia leucantha in Uttar Pradesh, India (Bhattacharya & Dey 1996).

Cryptomyzus Oestlund

Aphidinae: Macrosiphini

About 20 palaearctic species associated with Ribes and/or Lamiaceae. Apterae have long antennae, swollen siphunculi, secondary rhinaria on ANT III (except fundatrices), and long capitate hairs on body and often on antennae and legs. ANT I usually has an angular projection at its apex. Alatae have a dark dorsal abdominal patch or cross-bands. Two species have been introduced into North America, presumably as eggs on Ribes. The species were often included in earlier accounts of Capitophorus. Taxonomic accounts are available for western Europe (Hille Ris Lambers 1953, Heie 1994, Blackman 2010), Kazakhstan (Kadyrbekov 1993) and Japan (Miyazaki 1971). Kadyrbekov (2021a) provided keys to apterous and alate females. Guldemond (1990, 1991a) reviewed the evolution of host plant relationships and life cycles of European species, and Guldemond & Eggers-Schumacher (1989) studied allozyme differences. Bašilova (2010a) carried out biosystematic studies of Cryptomyzus in Lithuania including multivariate morphometrics and analysis of DNA (COI and EF-1α) sequences, and Bašilova et al. (2008) reported on karyotypes of seven European species.

Cryptomyzus alatavicus Kadyrbekov Apterae are greenish-yellow; BL 1.5-2.1 mm. On Scutellaria sieversii (= transiliensis) and S. catharinae in Kazakhstan (Kadyrbekov 2017a).

Cryptomyzus alboapicalis (Theobald) Apterae are pale greenish, with a green spinal stripe, or yellowish with faint greenish cross-bands (see influentialpoints.com/Gallery); BL 1.6-2.9 mm. On undersides of leaves of Lamium album, occasionally on Ballota nigra, and able to reproduce in the laboratory on L. amplexicaule and L. purpureum, but not on L. maculatum (Guldemond 1991b). In northern Europe, across to west Siberia, Caucasus and Kazakhstan. Monoecious holocyclic with apterous males, or sometimes anholocyclic. 2n=12.

Cryptomyzus ballotae Hille Ris Lambers Apterae are pale green (see influential points/gallery) BL 1.7-2.1 mm. On Ballota nigra, with records also from Lamium album, Leonurus sp., Marrubium vulgare and Melittis melissophyllum. The specimens from Leonurus sp. in UK (BMNH collection, leg. V.F.Eastop) have shorter antennal, femoral and rostral hairs and could be a distinct taxon. In western, central and southern Europe, Iran, and possibly Pakistan, where it may be heteroecious holocyclic with a sexual phase on Ribes orientale (Naumann-Etienne & Remaudière 1995). It is also reported from South America (Argentina, Chile; Ortego 1998a). Probably anholocyclic in Europe, although oviparae and alate males have been obtained in the laboratory; oviparae being produced on Ribes alpinum, although not found on this species in nature (Guldemond 1991a). 2n=12.

Cryptomyzus behboudii Remaudière & Davatchi Apterae are white; BL 1.3-1.6 mm. On undersides of the basal, densely hairy leaves of Phlomis olivieri. In Iran and Turkey. Monoecious holocyclic with oviparae and alate males in September-October (Tuatay & Remaudière 1964).

Cryptomyzus elshotze Bhattacharya & Dey Colour of aptera in life is unrecorded; BL (fundatrix?) c.2.8 mm. Alatae are undescribed. Apterae (probably fundatrices) were collected in early May on an Elshotzia sp. in Garwhal Himalaya, Uttar Pradesh, India (Bhattacharya & Dey 2001).

Cryptomyzus galeopsidis (Kaltenbach) Apterae are pale greenish white, often with faint green spinal stripe (see influential points/gallery); BL 1.3-2.6 mm. On undersides of leaves of Ribes spp., most commonly R. nigrum. Heteroecious holocyclic, migrating to Lamium and Galeopsis, occasionally other Lamiaceae, and Veronica (Plantaginaceae), where they live dispersed on undersides of leaves. Colonies on Galeopsis can cause curling or rolling of edges of young leaves (Guldemond 1991a). In Europe (including Iceland and Faroes), west and east Siberia, Kazakhstan and North America. Populations monoecious holocyclic on Ribes in Europe have been distinguished as subspecies, G. galeopsidis citrinus Hille Ris Lambers which is lemon yellow and occurs on R. rubrum, and G. galeopsidis dickeri Hille Ris Lambers which is found only on R. nigrum. However, Bašilova (2010a) failed to obtain any obligatorily monoecious lineages on R. nigrum in her life cycle studies in Lithuania. Guldemond (1990, 1991b) studied the genetic relationships between life cycle category and host preferences in this species complex (see also discussion by Heie 1994). 2n=12.

Cryptomyzus heinzei Hille Ris Lambers Colour of apterae in life is unrecorded; BL 1.6-1.8 mm. Described from Clinopodium vulgare in Germany, but Guldemond (1991a) considered that its normal secondary host is Stachys officinalis, on which it has been found in Spain, Germany, Hungary and Czech Republic. Host alternation occurs to Ribes alpinum. It is closely related to and difficult to distinguish morphologically from C. ballotae (Guldemond & Eggers-Schumacher 1989).

Cryptomyzus karzhantavicus Kadyrbekov Apterae are white with green markings and red eyes; BL 1.4-1.9 mm. On undersides of leaves of Stachys betoniciflora in South Kazakhstan (West Tien Shan). Not attended by ants. Other morphs and life cycle are unknown (Kadyrbekov 2021a).

Cryptomyzus korschelti Börner Apterae are pale whitish green with some bright green mottling (see influentialpoints.com/Gallery), or (on Ribes) rose pink to salmon red; BL 1.5-2.5 mm. Heteroecious holocyclic, migrating from Ribes alpinum to undersides of leaves of Stachys silvatica, and also able to colonise certain Lamium and Galeopsis spp. under laboratory conditions, although it may not occur naturally on these plants. On Ribes the aphids live in cavities beneath red or yellow leaf blister galls, which can persist late into summer, and may eventually produce small apterous males (Guldemond 1990) . Widely distributed in Europe, and eastward at least to Central Asia (Narzikulov & Umarov 1969, Kadyrbekov 2005c, and probably also Nevsky 1929a – as C. ribis). It has also been reported from Argentina (see Ortego et al. 2004). 2n=12 (Bašilova et al. 2008).

Cryptomyzus leonuri Bozhko Apterae are whitish with a greenish tinge; BL c. 1.8 mm. Monoecious holocyclic on Leonurus cardiaca, and also recorded from L. quinquelobatus (Buga & Stekolshchikov 2012). Probably specific to Leonurus, although Lamium spp. may be colonised under laboratory conditions (Bašilova 2010a). In eastern Europe (Poland, Lithuania, Russia, Belarus, Ukraine). Bašilova (2010b) redescribed the apterous vivipara and also provided descriptions of alate viviparae, oviparae and alate males, and Bašilova & Rakauskas (2012b) discussed morphological discrimination from the very similar C. alboapicalis. 2n=12 (Bašilova et al. 2008).

Cryptomyzus (Alataumyzus) malkovskii Kadyrbekov Apterae are yellow-green with antennae and legs mainly brown, and brown siphunculi; BL 2.1-2.6 mm. Heteroecious holocyclic, migrating from Ribes spp. to Stachyopsis lamiiflora in Kazakhstan (Kadyrbekov 2002e).

Cryptomyzus maudamanti Guldemond Apterae are pale yellow to pale green, sometimes with a faint green spinal stripe (see influentialpoints.com/Gallery); BL 1.1-2.3 mm. Heteroecious holocyclic, migrating from Ribes rubrum to Lamium galeobdolon. Other Lamium spp. and Galeopsis tetrahit were colonised with reduced fecundity in the laboratory (Guldemond 1991a). In England, Netherlands, Germany, Lithuania (Bašilova 2010a), Czech Republic and Italy (Barbagallo & Patti, 1998). 2n=12 (Bašilova et al. 2008).

Cryptomyzus (Phlomimyzus) multipilosus Kadyrbekov Apterae are pale green; BL 1.6-2.2 mm. On Lamium album and Leonurus turkestanicus in Kazakhstan (Kadyrbekov 2000), and subsequently recorded from a primary host, Ribes meyeri (Kadyrbekov 2002e).

Cryptomyzus ribis (L.) Plate 22a (Fig.50a) Apterae are pale green to yellow-green or whitish, often with a green medial stripe (see influentialpoints.com/Gallery); BL 1.2-2.6 mm. On Ribes spp., especially R. rubrum, living in concavities on undersides of leaves formed by brownish or purplish red blister galls. Heteroecious holocyclic, migrating to leaves and stems of Stachys spp., with occasional records from other Lamiaceae. Colonies also sometimes persist on R. rubrum through the summer and these may eventually produce apterous males (Hille Ris Lambers 1953). In Europe, Central Asia, east Siberia, Korea, Japan, China and North America. [However, both the host range and the distribution are uncertain due to confusion with subsequently described species, particularly in Central Asia (C. korschelti) and the Far East (C. taoi).] 2n=12.

Cryptomyzus sairamugamicus Kadyrbekov Apterae are white with green markings and reddish eyes; BL 1.3-1.6 mm. On undersides of leaves of Phlomis salicifolia in South Kazakhstan (West Tien Shan). Not attended by ants. The life cycle is unknown (Kadyrbekov 2021a).

Cryptomyzus (Ampullosiphon) stachydis (Heikinheimo) (Fig.50b) Apterae are pale yellowish, whitish or pale green, with a spinal row of green spots; BL 2.7-3.1 mm. On young leaves of Ribes spp. in spring in northern Europe, causing yellowish vein-banding. It has recently been reported from western Siberia (Altai Republic; Stekolshchikov & Novgorodova 2013). Heteroecious holocyclic, migrating to undersides of leaves of Lamiaceae (Galeopsis, Lamium, Stachys). Stenseth (1971) studied its life cycle in Norway. 2n=12.

Cryptomyzus (Phlomimyzus) tadzhikistanica Narzikulov & Daniyarova Apterae are green or pale green, antennae and legs dusky; BL c.2.1 mm. In small colonies on undersides of leaves of Phlomis canescens in Tajikistan.

Cryptomyzus taoi Hille Ris Lambers Colour of apterae in life unrecorded, probably pale; BL c.2.0-2.3 mm. Described from China on Marrubium supinum, and subsequently found on Lamium album var. barbatum (Miyazaki 1971), L. amplexicaule, Leonurus sibiricus, Leucas aspera and Phlomis bracteosa (Naumann-Etienne & Remaudière 1995). In Pakistan, India, China, Mongolia, Japan, Korea and east Siberia. Heteroecious holocyclic, with Ribes spp. (burejense, fasciculatum var. chinense) as primary hosts (Lee et al. 2002c). 2n=12.

Cryptomyzus transiliensis Kadyrbekov Apterae are pale green; BL 2.0-2.2 mm. On Stachyopsis spp. in Kazakhstan, and also now recorded from Phlomis oreophila and from the probable primary host Ribes meyeri Kadyrbekov 2017a).

Cryptomyzus ulmeri (Börner) Apterae are mid to darkish green with faint cross-bands on dorsal abdomen; BL 1.0-1.5 mm. On undersides of leaves of Lamium maculatum, and probably specific to this plant, although it can feed on L. amplexicaule and L. purpureum in the laboratory (Guldemond 1991a). Monoecious holocyclic, with alate males. In western, central and northern Europe. 2n=12 (Bašilova et al. 2008).

Cryptosiphum Buckton

Aphidinae: Macrosiphini

A genus for 8 or 9 rather small, broadly oval aphids with short appendages galling leaves of Artemisia, a habit that has resulted in reduction of appendages and other characters, so that their taxonomic affinities are uncertain, and they have been placed in either Aphidini or Macrosiphini by different authors. While normally reliable characters for Aphidini such as marginal tubercles on abdominal tergites 1 and 7 are lacking, C. artemisiae has a very Aphis-like 2n=8 karyotype. However, the latest revision, by Kadyrbekov (2002b), places Cryptosiphum in Macrosiphini, and this has now been confirmed by molecular studies (Kim & Lee 2008a). The genus extends from western Europe to east Asia, and from Siberia to southern India and Burma, with most species in northern Asia. Heie (1986) gave a brief generic diagnosis, and Ivanoskaya (1960) keyed the species of former USSR.

The taxonomy is difficult and, as with another xerophyte-associated genus with short appendages, Brachyunguis, the separate identity of several of the nominal species is suspect. The antennae of alatae are considerably longer than those of apterae, but apterous specimens occur with antennae of intermediate length. The short appendages do not lie flat in slide preparations, and this magnifies errors of measurement. Populations in central and western Europe are easily separated (couplet 51 of Key A of Artemisia) into two host-specific species (artemisiae and brevipilosum), but apterae of this group from Asia are commonly intermediate in the characters that discriminate European apterae, although alatae consistently have the lower number of secondary rhinaria that is characteristic of C. brevipilosum.

Cryptosiphum artemisiae Buckton Plate 14g Apterae are dark red or brown to almost black, powdered with greyish wax; BL 1.1-1.9 mm. Alatae are also wax-powdered in life and have secondary rhinaria distributed ANT III 8-30, IV 0-5, V 0-2. It lives in globular red or yellowish leaf galls (see influentialpoints.com/Gallery) formed on various Artemisia spp., especially A. vulgaris, throughout the palaearctic region. Monoecious holocyclic; oviparae (with conspicuously swollen hind tibiae) and alate males occur in September-October in England (Blackman 2010) and October-early November in Japan (Sugimoto & Seino 2011). Several populations from China with ANT PT/BASE 1.05-1.30 (as opposed to 0.45 in fundatrix and 0.7-1.0 in summer apterae of C. artemisiae s. str.) have been described as a subspecies, C. artemisiae ssp. linanense Zhang, but are probably a distinct species, possibly C. sieversianae (q.v.). Chinese populations (of C. artemisiae s. str.) were heavily parasitised. C. artemisiae is notable for having a specific syrphid predator, Triglyphus primus. 2n=8.

Cryptosiphum astrachanicae Ivanoskaya Apterae are broadly oval, BL c. 1.0-1.1 mm. In leaf galls on Artemisia and Seriphidium spp. in southern Russia, and Kazakhstan (Kadyrbekov 2002b, 2009b).

Cryptosiphum brevipilosum Börner Apterae are broadly oval, brownish, wax-powdered, BL 1.1-1.3 mm. Distorting the shoots of Artemisia campestris in central and eastern Europe (Germany, Poland, Hungary, Ukraine), eastward to Kazakhstan (Kadyrbekov 2002b). C. atriplicivorum Zhang, Chen, Zhong & Li on Artemisia sp. in China (G. Zhang 1999) is closely related or synonymous.

Cryptosiphum dracunculum Kadyrbekov Apterae are brownish, thickly powdered with white wax; BL 1.1-1.5 mm. In leaf galls on Artemisia dracunculus in Kazakhstan (Kadyrbekov 2002b).

Cryptosiphum innokentyi Ivanoskaya Apterae are broadly oval, pale green, thickly wax-powdered; BL c. 1.3 mm. In cigar-shaped leaf galls on Artemisia frigida and another Artemisia sp. in Russia (Siberia: Kuznetskiy Alatau, Tuva) and Kazakhstan (Kadyrbekov 2009b).

Cryptosiphum mordvilkoi Ivanoskaya Apterae are brownish, with thick white wax film; BL 0.9-1.2 mm. In oblong galls on Artemisia sp(p). in southern Urals and north Kazakhstan (Kadyrbekov 2002b).

Cryptosiphum sieversianae Ivanoskaya Apterae are yellowish green or brownish with thick white wax film; BL c.1.4 mm. In leaf galls on Artemisia sieversiana in southern Russia (Kulundinskaya) and east Kazakhstan (Kadyrbekov 2002b). C. artemisiae ssp. linanense, recorded from several Artemisia spp. in China, could be large specimens of this aphid.

Crypturaphis Silvestri

Calaphidinae: Calaphidini

A genus for one Alnus-feeding species with peculiar projections of the head and prothorax.

Crypturaphis grassii Silvestri (fig. 27B,C) Apterae are brownish, dorso-ventrally flattened (see influentialpoints.com/Gallery); BL 2.3-3.2 mm. Living dispersed along veins on both upper and lower leaf surfaces of Alnus cordata, and apparently specific to this host. Immature stages are green. In Italy and Corsica, and now established on the same host in various locations in southern England (Harrington 1998, Campbell 2007) and Wales (Baker 2009b). Alatae, the most commonly collected morph in summer, have black head and thoracic lobes, paler prothorax, and an ill-defined and variably developed brown patch across abdominal tergites 5-6; BL 2.2-3.0 mm. Oviparae and males occur in October-November in England and Italy (BMNH collection), but anholocyclic overwintering can also occur. Patti (1983) provided some biological information.

Ctenocallis Klodnitsky

Calaphidinae: Panaphidini

A genus for three species on Leguminosae/Fabaceae with body of aptera bearing long finger-like marginal backwardly-directed processes. These processes are also present in a much reduced form in alatae. Quednau (2003) reviewed the genus and illustrated all the species. Halaj & Osiadacz (2014) reviewed host plants and distributional data.

Ctenocallis dobrovljanskyi Klodnitsky Apterae are straw-coloured with brown dorsal sclerites; BL c. 1.5 mm. On Cytisus spp., Genista pilosa and Lembotropis nigricans, colonising young growth, leaves and leaf petioles. Recorded from Belarus, Ukraine, west Kazakhstan, Romania, Hungary, Czech Republic, Serbia, Austria, France and northern Italy. Monoecious holocyclic; oviparae and small alate males were collected in late August-September in Italy (BMNH collection, leg. H.L.G. Stroyan).

Ctenocallis israelica Hille Ris Lambers Apterae are yellowish with brown to blackish brown dorsal sclerites; BL 1.5-1.7 mm. On leaves of Calicotome spp. in the Mediterranean area (France, Spain, Italy and Israel), and also now recorded from Hungary (Halaj & Osiadacz 2014). Broza et al. (1992) noted that attacks by this aphid killed seedling plants after forest fires. 2n=16.

Ctenocallis setosa (Kaltenbach) Plate 4b Apterae are yellowish with brown dorsal sclerites; BL 1.5-1.6 mm. Feeding inconspicuously, flattened against the midribs, on the upper sides of leaves of Cytisus spp. (scoparius, villosus). Widely distributed in Europe (including UK) and North America (North Carolina, Ontario, north-west Pacific region). Oviparae and alate males occur in September. 2n=18.

Cyrtomophorodon Zhang & Qiao

Aphidinae: Macrosiphini

One species collected on a fern in China, with dorsal side of head smooth and first tarsal segments bearing only 2 hairs, perhaps belonging in Micromyzella.

Cyrtomophorodon cyrtomophitum Qiao & Zhang Apterae are yellowish-white in life ( presumably with a distinctive pattern of dark dorsal sclerotisation); BL c.1.9 mm. On Cyrtomium falcatum in Shanxi Province, China (Qiao et al. 2000). Other morphs and life cycle unknown.