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SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order) C Cinara abietihabitans to
murrayanae Cinara nepticula to zoarcbursara
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).
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 (Starı 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 blackish antennae and tibiae and dark
wing veins; 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; 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 Moritsu 1983, as Neocalaphis); 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.
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.
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).
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 Compositae and Polygonaceae. Some species are monoecious holocylic
on plants in Elaeagnaceae, Compositae 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 Compositae (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 Compositae (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, 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, 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.
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).
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).
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,
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; 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.
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.
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; 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; 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.
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 (Starı 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. 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. 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, 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.
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).
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.]
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.
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.
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, 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
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).
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.
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.
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).
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).
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
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.
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; 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, but apparently not P.
anserina) may also be colonised. 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.
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).]
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).
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.
Molecular studies also indicate that Chaitophorus
may have a closer relationship to the monocot-feeding Siphini than to the Acer-feeding genus Periphyllus (Wieczorek & Kajtoch
2011). 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 & Starı 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 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).
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. 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 a record from Turkey (Akyürek et al.
2011) requires
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), 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, 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 (Murmansk region; Stekolshchikov &
Buga 2018). 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 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). 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, 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) 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. 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;
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.]
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.]
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.
(=
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 Compositae, 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.).
One species in Europe and west
Asia on roots of Compositae, 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.
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.
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
(Starı 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.
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).
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).
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.
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).
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; Starı 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) 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 (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, 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), Siberia (Pashchenko 1988a)
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
Cu1a and Cu1b. 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). 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, 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). 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). 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 conformation. 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, 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). 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), Siberia (Pashchenko 1988a), 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).
Starı (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
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), 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, 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 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 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 |
