SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order) T
Nine species on trees in various families, but
especially Myrtaceae, in India, and east and south-east Asia. In several
species only alate viviparae are known, but this may be because apterous
viviparae are well camouflaged on the stems and easily overlooked, whereas
the alatae develop on the leaves. One species (dineni) was described from an unidentified species of
Combretaceae. Accounts are available for Java (Noordam & Hille Ris
Lambers 1985) and India (A.K. Ghosh & Quednau 1990), and Quednau (2010)
keyed the world fauna and illustrated all available morphs. Taiwanaphis atrovirens
Noordam & Hille Ris Lambers
Alatae have head, pronotum and sides of abdominal segments 3-8
yellowish or brownish, mesonotum and central part of dorsal abdomen dark
green; BL 1.2-1.5 mm. Immature alatae are yellowish-white with a dorsal
greenish-black area. Apterous viviparae, other morphs and life cycle unknown.
On upper and lower sides of new leaves and developing shoots of Syzygium sp. in Java (Noordam &
Hille Ris Lambers 1985). Taiwanaphis atuberculata
Noordam & Hille Ris Lambers
Alatae are orange with grey sides to thorax and abdominal segments
1-5, banded antennae and black-bordered forewing veins; BL 1.5-1.7 mm. Other
morphs and life cycle unknown. On upper sides and undersides of leaves and
sometimes on developing shoots of Syzygium
lineatum in Java. Taiwanaphis decaspermi
Takahashi Apterae and alatae are
brownish-black to black; BL 1.2-1.6 mm. Alatae have thickly bordered forewing
veins. Described from Decaspermum
fruticosum in Taiwan, attacking undersides of young leaves and shoots.
Subsequently recorded from other Myrtaceae (Syzygium buxifolium, hancei) in China (Guangdong, Hainan, Hong
Kong), and Japan. Alate males were collected in Hong Kong in March (BMNH
collection, leg. Commonwealth Institute of Entomology). Taiwanaphis dineni Mandal,
Agarwala & Raychaudhuri Alatae
have thickly-bordered forewing veins, colour in life is unrecorded; BL
1.8-2.0 mm. On an unidentified species
of Combretaceae in Sikkim, India (Mandal et
al. 1979). Apterae are unknown, but an alate male and an ovipara were
collected in October (A.K. Ghosh & Quednau 1990). Taiwanaphis kalipadi
(Raychaudhuri & Ghosh) Colour in
life is not recorded; apterae are rather elongate oval, BL 1.2-1.5 mm. On Annona squamosa in West Bengal, India
(Raychaudhuri & Ghosh 1964). Other morphs and biology are unknown, but
possibly T. randiae, known only
from alatae, is this species. Taiwanaphis memecyloni
Noordam & Hille Ris Lambers
Alatae are yellow with black sides to thorax and grey-bordered
forewing veins; BL 1.0-1.5 mm. On upper surfaces of young leaves, with
immatures positioned close against the main vein, on Memecylon lilacinum (= laevigatum, myrsinoides) in Java (Noordam & Hille Ris Lambers 1985).
Other morphs and life cycle are unknown. Taiwanaphis montanicola
Noordam & Hille Ris Lambers
Apterae and alatae are brown, the alatae having black-bordered
forewing veins; BL 1.7-2.2mm. On the undersides of young leaves of Syzygium racemosum in Java (Noordam
& Hille Ris Lambers 1985). The life cycle is unknown. Taiwanaphis pseudocaudata
Noordam & Hille Ris Lambers
Alatae have head and prothorax yellow, mesothorax and abdomen orange
with grey sides; BL 1.1-1.6 mm. Along veins on upper and lower sides of young
leaves and shoots of Syzygium
syzygioides in Java (Noordam & Hille Ris Lambers 1985). Other morphs
and life cycle are unknown. Taiwanaphis randiae
Ghosh, Banerjee & Raychaudhuri Alatae
are yellowish with dark markings and heavily bordered forewing veins; BL
1.7-2.0mm. On upper and lower sides of terminal leaves of Randia sp. in India (Arunachal
Pradesh), attended by ants (A.K. Ghosh et
al. 1971d). Other morphs and life cycle are unknown; possibly a synonym
of T. kalipadi, which is only known
from apterae.
About
six palaearctic species mostly on Saxifragaceae or
ferns, perhaps related to Utamphorophora
but with dorsal spiculation of the head. Apterae usually have secondary
rhinaria extending in a row along ANT III. Taiwanomyzus alpicola (Hille Ris Lambers) (Fig.43m) Apterae are shining black with antennae
and femora mainly black, and dark siphunculi and cauda; BL 1.1-1.7 mm. On ferns, (Asplenium, Athyrium, Blechnum, Cystopteris, Dryopteris, Gymnocarpium,
Polypodium), particularly those
growing in shady situations (Müller 1987). In central and eastern Europe (Germany,
Poland, Switzerland, Slovakia, Czech Republic). Monoecious holocyclic with
oviparae and apterous males in September (original description, as Utamphorophora). Taiwanomyzus babai Sorin & Arakawa Colour of apterae in life is unrecorded;
BL c.2.3 mm. Alatae have secondary rhinaria distributed III 30-34, IV 5-7, V
0. On Osmunda japonica in Japan
(Sorin & Arakawa 2005). Taiwanomyzus chrysosplenii Miyazaki
Apterae are pale brown to reddish brown, with black antennae and cauda,
siphunculi also blackish but often with a paler middle section; 1.3-1.6 mm.
On Chrysosplenium flagelliferum, feeding on undersides of
leaves in a wet environment (Miyazaki 1971), in Japan. Taiwanomyzus filicis (Miyazaki)
(Fig.43l) Apterae are pale yellow with mainly pale
appendages; BL 1.2-1.7 mm. Alatae have secondary rhinaria distributed III
30-39, IV 4-6, V 0(-1). On ferns (Athyrium,
Dryopteris, Osmunda) in Japan. The life cycle is unknown; alate males were
collected on Athyrium in late
September (original description). Taiwanomyzus himalayensis (Chakrabarti & Banerjee) Colour of apterae in
life is unrecorded, presumably pale; BL 2.0-2.3 mm. Alatae have secondary
rhinaria distributed III 2-6, IV 0, V 0. On ferns (Asplenium, Polypodium)
in Uttar Pradesh, north-west India (original description, as Utamphorophora). Taiwanomyzus montanus (Takahashi)
Plate 24f (= T.
darjeelingensis M.R. Ghosh, R.C. Basu & Raychaudhuri) Apterae are yellow with extensive black
dorsal markings, and black antennae, siphunculi and cauda; BL 1.8-2.4 mm. On
Saxifragaceae (Astilbe, Saxifraga) and also (in Japan) on Hydrangeaceae
(Hydrangea, Schizophragma). In Taiwan, Japan, Korea, Philippines
(Calilung 1967, as Aulacorthum) and
West Bengal, India (M.R. Ghosh et al.,
1977, as T. darjeelingensis).
Seven species of narrow-bodied aphids on
bamboos, resembling Myzocallis but
with the clypeus bearing an anteriorly directed tubercle, and with the
antennal terminal process about equal in length to the base of ANT VI.
Usually all viviparae are alate. The genus is oriental in origin, but several
species are now widely distributed. Higuchi (1968) revised the species then
known, and accounts are available for Japan (Higuchi 1972), India (A.K. Ghosh
& Quednau 1990), China (Qiao & Zhang 2004b), Korea (Y. Lee & S.
Lee 2018), and for species introduced into Britain (Stroyan 1977). Quednau (2003) reviewed the
genus, provided keys to species and illustrations of all available morphs. A
specific parasitoid Trioxys remaudierei Starý & Rakhshani has been
described that is associated with introduced Takecallis spp. in France and Spain (Rakhshani et al. 2017). Takecallis affinis
L.K. Ghosh Alatae are blackish,
wax-covered in life; BL 2.1-2.5 mm. On undersides of leaves of Bambusa sp. and Arundinaria jaunsarensis (= Yushania
anceps) in Himachal Pradesh,
India. Apterous viviparae are unknown. Apterous oviparae (illustrated by
Quednau 2003) and alate males occur in October (L.K. Ghosh 1986). B.C. Das
& Chakrabarti (1989b) recorded a new parasitoid species, Trioxys takecallis, from T. himalayensis (= T. affinis) in Garwhal, western
Himalaya. Quednau (2003) described a closely related form from Arundinaria sp. (or Yushania niitakayamensis?) in Taiwan
with both apterous and alate viviparae as a subspecies, T. affinis niitakayamensis. Takecallis alba Lee Alatae are pale yellow, covered in white
wax; BL 2.1-2.5 mm. The wing veins have conspicuous black spots/triangles at
their distal ends. On undersides of leaves of Pseudosasa sp. and Sasa
spp. in South Korea (Y. Lee & S. Lee 2018). The life cycle is unknown. Takecallis arundicolens
(Clarke) Alatae are pale yellow or
greyish-yellow, wax-dusted, usually without any dark dorsal abdominal
markings, but with a black cauda and black-banded antennae (see
influentialpoints.com/Gallery); BL 1.8-2.8 mm. [An
unpigmented population with a pale cauda occurred at Wisley, UK, in 1967
(BMNH collection, leg. K.M. Harris).] On mature leaves of bamboos (Arundinaria, Bambusa, Phyllostachys,
Sasa) in China, Japan, Korea,
eastern Russia and introduced into Europe, Africa (Algeria) and USA
(California). DNA barcoding has revealed differences between specimens from
European and Asian populations which might be indicative of there being more
than one species under this name (Y. Lee et
al. 2017). The life cycle in east Asia is unknown; probably it is mostly
anholocyclic where introduced, although Leclant (1966) reported oviparae in
southern France, and Quednau (2003) illustrated an ovipara collected in
California. 2n = 18. Takecallis arundinariae
(Essig) (= takahashii Hsu; synonymy by Quednau 2003) Alatae are whitish, pale yellow or
greyish-yellow, with paired black elongate spots on abdominal tergite 1-7 and
a pale cauda (see
influentialpoints.com/Gallery); BL 1.7-2.4 mm. An apterous
morph has been described, but only from high altitudes in Taiwan. On
undersides of mature leaves of bamboos (especially Arundinaria and Phyllostachys,
but sometimes also Bambusa, Dendrocalamus, Pseudosasa and Sasa).
In India, China, Taiwan, Korea, Japan and introduced to England, Netherlands (Piron 2009), Hungary (Basky & Neményi
2014), Madeira
(Aguiar & Ilharco 1997), Australia
(Valenzuela et al. 2010a), New
Zealand, North America and Argentina. An ovipara collected on Sasa kurilensis in Korea in early
November was illustrated by Quednau (2003); it is anholocyclic where
introduced. Coffelt & Schultz (1990) studied its population dynamics in
USA. 2n == 18. Takecallis assumenta Qiao
& Zhang Alatae are yellow, with
greyish-blue antennae; BL 1.9-2.5 mm. On upper surfaces of leaves of Bambusa spp., causing yellowing. In Qinghai, China (Qiao & Zhang 2004b).
Other morphs and life cycle unknown. Takecallis sasae
(Matsumura) Alatae are green; BL c.
1.8 mm. On leaves of Phyllostachys,
Pleioblastus, Sasa and possibly also Bambusa,
in Japan (Higuchi 1968). Tao (1964) had T.
taiwanus in China and Taiwan according to Higuchi (1968). The life cycle
is unknown. Takecallis taiwana (Takahashi) Alatae are pale green with a pale cauda (see influentialpoints.com/Gallery) ; BL 1.4-2.3 mm. Immatures are
yellow-green with longitudinal rows of small brown spots. In young, still
unrolled leaves and on new shoots of bamboos (Arundinaria and Phyllostachys,
sometimes Bambusa, Sasa). In China, Taiwan, Japan and
introduced to Europe, South Africa, New Zealand, USA, and more recently to
South America (Argentina, Delfino 2001; Chile, Gonzales et al. 2000). Cottier's (1953) account under T. arundinariae should be referred to this species. Quednau
(2003) illustrated an ovipara collected in China in early December, but it is
probably anholocyclic where introduced. 2n = 16.
A relict nearctic genus with 4-5 species living without
host alternation in galls on Arctostaphylos.
A detailed diagnosis and review were provided by Remaudičre & Stroyan
(1984). Oviparae are alate. The male genitalia of an unidentified species of this genus were
described and illustrated by Wieczorek (2011). Miller
& Crespi (2003) and Miller et al.
(2015) studied the molecular phylogeny of the genus and the evolution of
inquilinism and host relationships. Their studies included several
undescribed species including two nomina nuda collected on other genera of
Ericaeae, T. morani from Arbutus arizonica (“Tamalia sp. A” in Miller et al. 2015) and T. cruzensis from Comarostaphylis
diversifolia (“Tamalia sp. B”)
in Miller et al. 2015). Tamalia
coweni (Cockerell) Plate 3h Apterae are dirty yellow to blackish, with
variably-developed dark green cross-bands both dorsally and ventrally (see
influentialpoints.com); BL 1.25-1.5 mm. In reddish leaf-galls on Arctostaphylos spp. in western North
America, across boreal Canada to Ontario, and south to Mexico. Monoecious
holocyclic, with alate oviparae and males produced from June to October. The
peculiarities of the life cycle were discussed by Remaudičre & Stroyan
(1984), and studied further by Miller & Avilés (2000). T.
keltoni Richards, described from Arctostaphylos
sp. in Mexico, is doubtfully distinct. 2n=6. Tamalia
dicksoni Remaudičre &
Stroyan Colour of apterae in life is
unknown; BL 1.4-2.1 mm. In leaf-galls on Arctostaphylos
pringlei in California, USA (Miller & Crespi 2003). Presumably monoecious
holocyclic; males were collected in late June (original description). Tamalia
inquilinus Miller Apterae are dark grey to brown or black;
BL 1.4-1.7 mm. Living as an inquiline in leaf-galls of T. coweni on Arctostaphylos
spp. (Miller & Sharkey 2000). On west coast of North America from
north California to Baja California Norte, Mexico. Alate oviparae and males
were collected in late July. Miller & Crespi (2003) compared the relative
rates of mtDNA evolution of T. coweni
and T. inquilinus, Miller (2004)
studied the ecology of the relationship between the two species, and Miller et al. (2015) analysed their
phylogenetic relationship. Tamalia
milleri Kanturski &
Wieczorek Colour of apterae in life
is unknown (abdomen has broad dark spinopleural cross-bands); BL 1.8-2.1 mm.
On Arctostaphylos columbiana in
western USA (California, Oregon). Apterous viviparae and alate oviparae were
collected in August (Kanturski & Wieczorek 2015a).
Two oriental species on Betulaceae, possibly
related to Euceraphis and Cepegillettea, characterized by the
small rounded secondary rhinaria, ANT PT a little shorter than BASE VI and
siphunculi with swollen bases and a single appended hair. Both apterous and
alate viviparae occur. Quednau (1973) gave an account of the genus. Taoia chuansiensis
(Tao) Apterae and alatae are yellowish-green, the alatae having darker legs
and antennae, and dark-tipped siphunculi; BL 2.9-3.1 mm. Living solitarily on
undersides of leaves of Alnus spp.
(cremastogyne, japonica) and Betula
spp. in China (Tao 1964, as Euceraphis
chuansiensis; Qiao et al.
2003). Sexual morphs and life cycle are unknown. Taoia indica
(Ghosh and Raychaudhuri) Apterae are pale green; BL 2.7-4.1 mm. Alatae have
darker antennae and legs, and dark-tipped siphunculi. On Alnus nepalensis and Betula
alnoides in India, Nepal (A.K. Ghosh & Quednau 1990), on Alnus nitida in Pakistan
(Naumann-Etienne & Remaudičre 1995), and on Alnus sp. in China (Qiao et
al. 2003). There are many Indian records from other genera, presumably
due to “casual visits”. Oviparae were collected in December and apterous (or
alatoid) males in April, July and December (R.C. Basu & Raychaudhuri
1980).
One species from
Ukraine related to Lipamyzodes but
with dorsal abdominal hairs arising
from tubercles and apterae with secondary rhinaria on ANT III. Tauricaphis
arabisi Mamontova Apterae are greenish with mainly
yellowish brown appendages; BL c. 1.2 mm. On Arabis caucasica in Ukraine. Monoecious holocyclic; oviparae were
collected in September (original description).
One species in China
with elongate body, short 4- or 5-segmented antennae, long thick dorsal hairs
and numerous short dorsal cylindrical processes. Tenuilongiaphis stata Zhang &
Zhong Apterae are pale green, with
two longitudinal rows of darker green spots; BL 1.7-2.0 mm. Sedentary in
habit, and only occurring in very small numbers, on Artemisia sp. in Gansu province, China. Believed to be
heteroecious holocyclic, as a fundatrix(?) described from an unidentified
shrub (G. Zhang 1999, as Sportaphis
sporta; see G. Zhang & Qiao 2002),
appears to be this species, and alatae and oviparae described as Ceruraphis hippophaetropis collected
on Hippophae rhamonoides (G. Zhang
1999) seem likely to be be the autumn forms. Specimens with 4-segmented
antennae found on Artemisia
lavandulaefolia in Shanxi province, were described as a subspecies, Tenuilongiaphis
stata ssp. shanxiensis (L.
Zhang et al. 2008).
About 30 species in which the apterae are usually very
globose and have
one-segmented tarsi and the alatae have a simple, unbranched media in the
forewing and usually ANT IV much shorter than ANT V. At least seven species
have host alternation between leaf galls on Ulmus and roots of Poaceae, and the lesser-known species are described
either from Ulmus or grass roots.
The elm galls are invariably stalked, pouch-like outgrowths on upper sides of
leaves. Tetraneura seems to be
predominantly oriental and eastern palaearctic but T. ulmi appears to be native to northwest Europe, T. caerulescens and T. africana are mainly around the
Mediterranean and in south-west Asia. T.
fusiformis and T. nigriabdominalis are
probably east Asian although populations of these species are now widespread.
Hille Ris Lambers (1970b) revised the world fauna and there are more recent
accounts from India (A.K. Ghosh 1984b), China (G. Zhang et al. 1991a, G. Zhang et
al. 1999c), the
UK (Blackman et al. 2019b), and
the Iberian peninsula (Nieto Nafría et
al. 2002a). Akimoto (1995) studied the interactions within the complex of
six species found on Ulmus japonica,
and Suzuki et al. (2009) compared
amino acid concentrations in their galls. The genus is in need of further
revision. The karyotype variation may be useful in this respect. (See also
Blackman & Eastop 2000, pp. 348-351. Tetraneura aequiunguis
Zhang & Zhang Only the fundatrix
is described, from an unidentified Ulmus
sp. in China (Zhang et al.
1991). Tetraneura africana van der Goot
Apterae are buff‑coloured to pale brownish with a dark brown
head and prothorax, often bluish due to waxy bloom, and with whitish wax
secreted posteriorly; BL 2.0-3.3 mm. In root‑feeding colonies on Cynodon dactylon, often well below
ground level, and also recorded from Sorghum
halepense, Calamagrostis spp., Phragmites australis, and rarely on
cereals. Recorded from Italy, Morocco, Egypt, Turkey, Ukraine, Iran, and east
to Kazakhstan and Pakistan. A smaller aphid collected on the roots of Calamagrostis epigeios in Poland may
also be this species (Hille Ris Lambers 1970b). Galls of T. africana on Ulmus
have not been identified, although Mordvilko (1935) described sexuparae
collected from Ulmus campestris in
Italy. Populations on Cynodon roots
are probably mainly or entirely anholocyclic. Tetraneura asymmachia Zhang
and Zhang Described from alatae (BL
c.2mm) obtained from galls on Ulmus
pumila and Ulmus sp. in
June-July in China (Zhang et. al.
1991). Appears closely related to T.
chinensis. Tetraneura basui Hille Ris Lambers Colour of
apterae in life is unrecorded; BL 1.5-1.8 mm. On roots of Poaceae (Capillipedium, Echinochloa, Eleusine,
Eragrostis, Oryza, Pagonantherum, Paspalum, Pennisetum, Polypogon, Setaria)
in north-east India (Raychaudhuri 1980). It is not known whether there is a
gall generation on Ulmus. Tetraneura brachytricha Zhang & Zhang Colour of apterae in life is not
recorded; BL c.2.7 mm. On roots of Phragmites
communis in Liaoning Province, China. Presumed to be heteroecious
holocyclic; alate sexuparae collected on Ulmus
in Xinjiang Province were assigned to this species, but the life cycle
needs conformation. Tetraneura caerulescens
Passerini Galls are stalked,
hazelnut- or walnut- sized (fig. 134G),
red when mature, on Ulmus spp. (glabra, laevis, procera, minor, montana) in southern and south-eastern Europe, north Africa and
south-west and central Asia. Alatae (BL 1.5-2.2 mm) emerge in late May-July
and migrate to found colonies on roots of Poaceae (Alopecurus, Cynodon, Echinochloa, Eragrostis, Festuca, Milium, Poa, Setaria). Apterae on grass roots
are orange-brown to brown, secreting bluish flocculent wax; BL 1.5-2.2 mm. Records
of secondary host populations in China (Zhang et al. 1985, 1991a) need further confirmation. Tetraneura capitata Zhang & Zhang Apterae are pale reddish brown; BL c.2.2
mm. On roots of Sorghum vulgare (= bicolor) in Hebei Province, China.
A population on Agropyron cristatum in Gansu Province was described as a
subspecies, T. capitata ssp. agropyricena
Zhang (in Zhang & Qiao 1997b). Tetraneura chinensis
Mordvilko Galls are apparently rather
large, sac-like, broader than high, green and slightly hairy (fig. 134F). Recorded
from Ulmus sp. in China (Mordvilko
1935, as T. hirsuta) and U. pumila in Mongolia (Holman &
Szelegiewicz 1974a). Alatae (BL 2.2-2.6 mm) were obtained from galls in
May-June. Secondary host populations on roots of Poaceae have
been identified by comparison of COI sequence (H. Zhang et al. 2008). Tetraneura fusiformis Matsumura (= T.
nigriabdominalis of many authors, incl. Hille Ris Lambers 1970b; = T.
hirsuta Baker) Apterae are greenish‑ or
brownish‑white; BL 1.5-2.5 mm. In colonies on roots of many genera and
species of Poaceae, (Agropyron, Axonopus, Cenchrus, Chloris, Cynodon, Dactyloctenium, Echinochloa,
Eleusine, Eragrostis, Oryza, Panicum, Paspalum, Pennisetum, Saccharum, Setaria, Sorghum),
their presence often indicated by a reddish‑purple discoloration of the
leaves. In Middle East, Africa, India, Nepal, Bangladesh, Pakistan, Sri
Lanka, Andaman Isles, Thailand, Japan, Korea, Indonesia, Malaysia, the
Philippines, New Britain, Australia, New Caledonia, Fiji, Tonga, Brazil, Argentina,
Columbia and Central America. Heteroecious
holocyclic in Japan, with a sexual
phase and galls on Ulmus japonica; the galls are upright
spindle-shaped, up to 12mm in height, petiolate at base, with numerous
granular protuberances and short white silky hairs, usually crimson when
mature. Alatae
(BL 1.7-2.3 mm) emerge in June-July through a lateral opening. T. fusiformis is probably entirely
anholocyclic outside the range of Ulmus
japonica, which seems to be its only primary host; its distribution suggests
that parthenogenetic overwintering is only possible in tropical and
subtropical conditions. The taxonomic confusion involving the application of
the name T. nigriabdominalis was
discussed by Eastop & Blackman (2005). All
African records of T. nigriabdominalis
as well as many of those from Asia (Blackman & Eastop 2000) should be
referred to T. fusiformis, although
past pest outbreaks on upland rice in Japan, and perhaps elsewhere in east
Asia, were of the true nigriabdominalis
(S. Akimoto, pers. comm.). Hille Ris Lambers (1970b, as T. nigriabdominalis) distinguished several root-feeding
populations on the basis of abdominal chaetotaxy, and erected a subspecies, bispina, for his material from Africa
and North America. However, most of the African material in the BMNH
collection does not conform to bispina,
and probably there are several distinct clonal populations distributed around
the world. Raychaudhuri (1980) distinguished two populations in north-east
India. 2n = 18 (gall generation); 2n = 17, 18, 19 and 20 (permanently
parthenogenetic populations). Tetraneura indica L.K. Ghosh Apterae are yellow; BL 1.9-2.3 mm. Alatae
are unknown. In ant-attended colonies on roots of Calamagrostis pilosula in Himachal Pradesh, north-west
India. According to Chakrabarti (2001)
this species produces leaf-fold galls on an Ulmus sp., but the primary host generations have apparently not
been described. Tetraneura javensis van der Goot
Apterae are whitish to yellowish‑white (see aphids
of Karnataka website); BL 1.5-2.5 mm. On roots of Poaceae, especially sugar cane (Saccharum officinale), and also
recorded from Capillipedium, Echinochloa, Eleusine, Neyraudia, Oryza, Panicum, Setaria and Sorghum. Anholocyclic populations of T. javensis occur on grass roots in Pakistan, India, Japan, Java and New Guinea, and
it is a pest of sugar-cane in India and through east and south-east Asia (Blackman
& Eastop 2006, p.1310). Probably T. javensis is heteroecious
holocyclic in Pakistan, where there are galls believed to be of this species
on Ulmus wallichiana (Hille Ris
Lambers 1970b, Naumann-Etienne & Remaudičre 1995). Alatae
from these galls had BL 1.8-2.0 mm and secondary rhinaria distributed ANT III
11-14, IV 3-5, V 4-7. The morphology of embryos inside these alatae
corresponded well with that of T.
javensis on grass roots (Hille Ris Lambers 1970b), but the life cycle
needs to be confirmed by host transfers. Setokuchi (1993) studied populations on sugar
cane in Japan. We have not attempted to include in the key four other species
described from India that were differentiated from T. javensis mainly on the basis of dorsal hair chaetotaxy, as
this can show much intraspecific variation; Tetraneura kalimpongensis Raychaudhuri,
Pal & M.R. Ghosh from roots of Saccharum
and Pennisetum in West Bengal, Tetraneura lambersi Chakrabarti
& Maity on an unidentified grass in Uttar Pradesh, Tetraneura multisetosa Raychaudhuri,
Pal & M.R. Ghosh from roots of Capillipedium
and Imperata in West Bengal,
and Tetraneura utpali Chakrabarti,
Maity & Bhattacharya from roots of Apluda
mutica in Uttar Pradesh. The group is clearly in need of further study. Tetraneura
nigriabdominalis (Sasaki) (= T.
akinire) Plate 5f Galls are stalked, hairy, elongate,
spindle- or pouch-shaped, 15-40 mm high, usually with a pointed apex (fig. 134E), usually
bicoloured green and rose-red when mature (see influentialpoints.com).
They occur on various Ulmus spp. (canescens, japonica, parvifolia, procera, pumila) in southern and south-east Europe (including Malta –
Mifsud et al. 2009), south-west
Asia (Iran, Georgia), Kazakhstan (Kadyrbekov 2014e), China, Korea and Japan.
It is also in eastern USA on U. minor
(Stoetzel 1987, as akinire), and in
western USA (California) on U. pumila
and hybrid elms (S. Dreistadt, pers.comm.). Galls on Ulmus sp. on high ground in Pakistan (Naumann-Etienne &
Remaudičre 1995, as akinire) are
probably also this species. Alatae (BL 1.4-2.3 mm) (fig. 117E)
leave the gall through lateral slits in May-July to found ant-attended
colonies on the roots of Poaceae (including Cynodon,
Digitaria, Echinochloa, Oryza, Saccharum, Setaria). Apterae on grass roots
are greenish‑ or brownish‑white; BL 1.5-2.5 mm. Their presence on
the roots of some hosts is indicated by a reddish-purple discoloration of the
leaves. This species is known particularly as
a pest of rice (Blackman
& Eastop 2000), but many of the references to nigriabdominalis,
e.g. in Africa, the Middle East,
India and south-east Asia, are referable to T. fusiformis. W. Lee et
al. (2012b) distinguished Japanese populations of T. nigriabdominalis (referred to as “Type A” in their paper) from
those of T. fusiformis (“Type B”)
by their differing mitochondrial COI sequences. Tanaka (1961) studied its
host preferences, although he may have had a mixture of T. nigriabdominalis and T.
fusiformis. Galli & Bonvicini-Pagliai (1998) studied its relations
with ants. Enoki
(1986) studied resistance to this aphid in U. japonica. The male genitalia were described and illustrated by Wieczorek (2012).
Hille
Ris Lambers' (1970b) descriptions of gall generations under this name (his
'Group I material') should be referred to T.
fusiformis (Eastop & Blackman 2005). Populations in China were described as a
subspecies, T. nigriabdominalis ssp. shanxiensis
Zhang & Zhang (as T.
akinire ssp.). 2n = 18 (in
gall generations); however Galli & Manicardi (1998) reported karyotype
variation from 2n =13 to 19. Tetraneura paiki
Hille Ris Lambers Described from a
single alate sexupara trapped in Korea (Hille Ris Lambers 1970b). Moritsu's
(1983) account and illustrations of an aphid under this name in Japan should
be referred to T. nigriabdominalis
(S. Akimoto, pers. comm.). Tetraneura polychaeta
Hille Ris Lambers Galls are conical,
bag-like (Chakrabarti et al. 1985),
on Ulmus laevigata (= villosa)
in Pakistan. Alatae (BL 2.0-3.2 mm) were collected from galls in April, but
probably emerge later in spring. A sexuparae trapped in Korea was also
tentatively assigned to this species (Hille Ris Lambers 1970b). Secondary
host generations are unknown. Tetraneura polychorema Zhang
Colour of apterae in life is not recorded; BL c. 2.1 mm. On roots of an undetermined grass species in Gansu
Province, China (Zhang & Qiao 1997). Tetraneura pumilae Lee,
Seo & Hwang Galls (undescribed)
are formed on Ulmus pumila.
Emigrant alatae are grey, BL 1.6-1.8 mm, emerging in early June (Lee et al. 1993). The secondary host is
unknown. T. iriensis Lee, Seo & Hwang was described from an
unidentified Ulmus sp. in the same
paper, with emigrant alatae of BL 1.2 mm, but the differences from T. pumilae are perhaps attributable to
smaller body size. Tetraneura radicicola
Strand Stalked pouch galls on Ulmus japonica in Japan are similar to
those of T. nigriabdominalis, but
hairless or with only sparse hairs (see http://www.agr.hokudai.ac.jp/systent/akimoto/gall/T.%20radic%20gall.jpeg).
Alatae (BL 1.7-2.5 mm) migrate in June-July to found colonies on roots of
Poaceae. Presumed anholocyclic populations of this species on grass roots are
recorded from Australia, Malaya, Philippines, Korea, Taiwan, India, Nepal and
Sri Lanka, although on its secondary hosts T. radicicola cannot be reliably distinguished from T. yezoensis (q.v.). Tong et
al. (2021) studied a mermithid nematode in Japan that parasitises alate
sexuparae. 2n
= 14. [Tetraneura persicina Zhang
& Zhang, described from alatae (BL c.2.6mm) from galls on Ulmus pumila and Ulmus sp. in
China (Zhang et al. 1991, Chen
& Qiao 2012), appears very similar to radicicola,
but the alatae have fewer dorsal cephalic hairs and their embryos have fewer
spinopleural hairs. See also T.
triangula.] Tetraneura sorini
Hille Ris Lambers Galls are stalked,
hairy, pouch-shaped, often somewhat globular, yellow and red (Moritsu 1983
and http://www.agr.hokudai.ac.jp/systent/akimoto/gall/T.%20sorini%20gall.jpeg).
On Ulmus spp. (laciniata, japonica, parvifolia) in Japan, Korea and China.
Usurpation of galls of their own and other Tetraneura spp. by first instars was studied by Akimoto &
Yamaguchi (1996). Alatae (BL 1.6-2.0 mm) emerge in June. Secondary host
generations are unknown. 2n = 16. Tetraneura triangula Zhang & Zhang Colour of apterae in life is not recorded;
BL c. 2.5 mm. On roots of various Poaceae (Echinochloa, Imperata, Phragmites, Sorghum) in Liaoning, Shanxi and Shandong Provinces, China. Described
from exules and sexuparae, which are similar to T. radicicola but much more hairy (W. Zhang & G. Zhang 1991).
Galls and spring generations are unknown, but possibly this species occurs in
Japan, on U. japonica (S. Akimoto,
pers. comm.). Tetraneura ulmi (L.) Elm‑Grass Root Aphid Galls
are stalked, approximately bean-shaped (fig. 134D),
smooth and shiny, reddish-green and/or yellow (see
influentialpoints.com/Gallery), on leaves of various Ulmus spp. (campestris, glabra, minor). Their histology was
investigated by Álvarez et al. (2013). Urban (2003a) made a
detailed study of its bionomics on elms. T.
ulmi
occurs commonly in Europe, across Asia to eastern Siberia (Pashchenko 1988a)
and northern Japan, and was introduced to North America. Alatae (BL 1.8-2.6
mm) (fig.
117E) emerge in June-July to colonize roots of Poaceae. Apterae on grass roots
(Plate 1d) are
pale orange yellow, yellowish white or reddish, with head, prothorax, and
appendages brown; the body lightly dusted with wax; BL 1.7-2.8 mm. They are
found on numerous species of Poaceae (especially Agropyron, Bromus, Dactylis, Deschampsia, Festuca, Holcus, Hordeum, Lolium, Poa, Zea), are invariably ant‑attended, and often in ants'
nests. Overwintering occurs commonly on grass roots or in ants' nests, root‑feeding colonies producing
rather few sexuparae in autumn. Ivens et
al. (2012a,b) used microsatellite DNA analysis to investigate clonal
diversity of root-feeding populations in the Netherlands. Zwölfer (1957) made
a detailed study of root-feeding populations, including comparison with the
closely related and perhaps synonymous Tetraneura longisetosa (Dahl), collected from roots of Brachypodium, Deschampsia and Festuca
in Germany. Tetraneura chui Zhang, described from Sorghum vulgare (= bicolor) in China, may also be a synonym. 2n (T. ulmi) =14 (gall generation); Galli & Manicardi reported 2n =14 to
16. Tetraneura ulmicema Zhang Known only from alatae emerging from galls
on Ulmus glabra in Ningxia, China;
BL c. 2.3 mm (G. Zhang & Qiao 1997a). The galls are smooth and hairless
(Chen & Qiao 2012). Tetraneura
yezoensis Matsumura
Stalked pouch galls on Ulmus
japonica and U. laciniata in
Japan resemble those of T. radicicola (see
http://www.agr.hokudai.ac.jp/systent/akimoto/gall/T.%20yezoensis.jpg).
Alatae (BL 1.8- 2.1 mm) migrate in late June-July to colonize roots of
various Poaceae (Echinochloa, Eleusine, Eragrostis, Imperata, Miscanthus, Oryza, Saccharum, Setaria, Triticum,
etc.).
Apterae on
grass roots are pale yellowish, brownish or pinkish white, with brown head
and prothorax. Anholocyclic populations on grass roots assigned
to this species also occur in Australia (Carver & Basu 1961, as T. heterohirsuta; Hille Ris Lambers
1970b). However
there are no really reliable morphological criteria for distinguishing adults
of T. yezoensis on grass roots from
those of T. radicicola, because the
adult chaetotaxy of this group is extraordinarily variable. Hille Ris Lambers
(1970b) noted that embryonic (and first instar) chaetotaxy is more stable
within species, but embryonic hair counts are almost impossible unless the
embryos are dissected out prior to mounting. Akimoto (1990) studied local adaptation to host
plant and possible host-race formation in T.
yezoensis. Suzuki et al. (2009)
found exceptionally high asparagine concentrations in its galls. 2n = 12 in
Japan (cf. T. radicicola); Chen
& Zhang (1985b) reported 2n == 18 from China, but probably had another
species. [T.
sikkimensis Raychaudhuri, Pal &
M.R. Ghosh, described from a small sample on Eragrostis nigra in Sikkim, India, is distinguished only by
characters of adult chaetotaxy and may be a synonym of T. yezoensis. Tetraneura
changaica, described from sexuparae only in Mongolia
(Szelegiewicz & Holman 1980), resembles T. yezoensis but has shorter R IV+V and different abdominal
chaetotaxy.]
One
species in China on Thalictrum,
perhaps related to Longicaudus but
with a short cauda bearing only 5-6 hairs. Thalictrophorus
thalictrophilus Zhang, Qiao & Zhao Apterae are white in life, BL c.1.9 mm. On
Thalictrum aquilegifolium in Shanxi
Province, China (Qiao et al. 2000).
Other morphs and life cycle are unknown.
About
12 species closely related and very similar morphologically to Pemphigus, but with some biological
differences, and retained here as a separate genus despite the reservations
of Aoki (1975), with Parathecabius
Börner as a subgenus. Spring generations form galls by folding the leaves of Populus. The fundatrix inhabits a
small gall set apart from that housing its progeny. Known secondary hosts are
Ranunculus, Lysimachia and Salix. Secondary hosts of
several species are unknown. Accounts are available for North
America (Smith 1974a), north-west Europe (Heie, 1980), the UK (Blackman et al. 2019b), Japan
(Aoki 1975, as Pemphigus) and China
(G. Zhang et al. 1995a, G. Zhang et
al. 1999c; their subgenus Oothecabius
Zhang is here synonymised with Epipemphigus).
[Note: The type species of Thelazocallis Zhang in Qiao & Zhang 2000c, T. ranunculicola, is evidently based
on exules of a Thecabius sp.; Thelazocallis Zhang 2000 is therefore
a new synonym of Thecabius Koch
1857.] Thecabius
affinis (Kaltenbach)
Plate 6d,
(fig. 3), (Fig.45a) Galls occur on Populus spp., in May-June, mainly on P. nigra and its varieties, and are formed
by folding of young leaves in half along mid-ribs; the outer, upper side of
the leaf becomes blistered and yellowish or reddish (see
influentialpoints.com/Gallery). These are secondary galls
containing only the alatiform progeny of the fundatrix, which inhabits a
small gall of its own formed by folding the edge of a leaf (fig. 131I).
Alatae (BL 2.2-3.1 mm, with green abdomen, fig. 86B)
leave the gall in late June-July to found colonies at stem bases and on
runners of Ranunculus spp. (fig. 3). Apterae on Ranunculus (Plate 2i)
are dirty yellowish white or yellowish green with shiny wax-wool (see
influentialpoints.com/Gallery); BL 2.1-2.9 mm. There may be rare occurrences on
other Ranunculaceae, e.g. Caltha
palustris (Barbagallo et al.
2011). Mordvilko
(1935) gave an account of the life cycle and described all morphs. Sexuparae
are produced from late August to fly back to poplars, but alate virginoparae
are also produced that colonise other Ranunculus
plants, and overwintering on buttercup roots is common. T. affinis occurs on a wide range of Populus spp. throughout Europe and across Asia to east Siberia,
Japan and Korea, and also in North America (see below). According to
Chakrabarti (1987, 2001), in northern India T. affinis induces leaf galls on Ranunculus hirtellus in spring, and migrates to form subterranean
colonies on Geranium spp. in late
summer and winter; the life cycle is still unclear, and warrants further
investigation, as this form would appear to be functioning as a distinct
biological species. Populations on east Asian Populus spp. are often treated as a separate species, T. orientalis Mordvilko, here regarded
as a synonym. T. populisuctus Zhang
& Zhong, described from sexuparae collected on P. cathayana in China (G. Zhang et al. 1995a) may also be a synonym. However, T. orientalis ssp. minensis Zhang, described from Ranunculus roots in China (G. Zhang et al. 1995a), differs from T. affinis by having 4 wax plates
instead of 2 on abdominal tergite 7, and could be a distinct species. The
karyotype of T. affinis is 2n=38 in
both England and Japan. [This chromosome number has been determined additionally for a sample
from British Columbia, Canada, leg. C.-k. Chan, and
COI barcode sequences have confirmed that
T. affinis is present in
North America (Skvarla et al.
2017). DNA data have also confirmed the existence of a separate North
American species, Thecabius populiconduplifolius (Cowen), which has 2n=28
(Harper & Macdonald 1966). T.
populiconduplifolius is virtually indistinguishable from T. affinis and has an identical
biology, but with native poplars as primary hosts (Palmer 1952).] Thecabius anemoni (Shinji)
Apterae are dark brick-brown, with white wax secreted from posterior
abdomen; BL unrecorded (“rather large”). On Anemone hupehensis (roots?) in Japan. The generic position is
uncertain. Thecabius (Parathecabius) auriculae (Murray) Apterae are white to dusky white or dirty
brownish, secreting wax-wool from posterior abdomen; BL 1.3-1.5 mm. Living
all-year-round on the roots of Primula spp.
in England, where it can be a pest of cultivated auriculas. It has also been
collected on Androsace sempervivoides and
Lysimachia (= Glaux) maritima. This
species is very similar to T.
lysimachiae, (see Stroyan 1964a), but unpublished data from more recently
collected specimens of apterae and alate sexuparae (the latter collected in
March and presumably redundant) has revealed some consistent differences (see
key to aphids on Lysimachia).
2n=16. Thecabius beijingensis Zhang Alatae from galls on Populus cathayana in China are yellow in life, BL c. 2.8 mm. They
appear very similar to emigrant alatae of T.
affinis except for the slightly higher rhinarial count. However, the
sausage-shaped mid-rib gall illustrated (G. Zhang et al. 1995a, fig. 2) looks more like that of an Epipemphigus, and is probably one
previously vacated by a species of that genus. Thecabius (Parathecabius) cerastii (Börner) Apterae with wax-wool; BL 1.4-1.6 mm. On
underground stems and roots of Cerastium uniflorum in Austria. The life
cycle is unknown. Thecabius (Parathecabius) gravicornis
(Patch) Galls are formed by folding
leaves downward along the mid-rib to form a yellowish-green, swollen pocket
which becomes filled with the developing emigrant alatae. The fundatrix
inhabits a separate small pocket on the upper side of a leaf lamina (Palmer
1952). On Populus balsamifera in
North America, widely distributed but apparently rather rare; also collected
on P. angustifolia in Utah (BMNH
collection, leg. R. Danielsson). Alatae (BL 1.7-2.8 mm, with olive-yellow
abdomen) migrate in July-August, presumably to an unknown secondary host. Thecabius (Parathecabius)
latisensorius Hori
Galls like those of T.
gravicornis are found on Populus
maximowiczii in Japan and Sakhalin, and on P. laurifolia and P.
talassica in Russia (Altai region) and Kazakhstan (Kadyrbekov 2017a).
Alatae (BL 3.0-3.5 mm, with dirty yellow abdomen, fig.
86A)
migrate in late July-September, presumably to an unknown secondary host (Hori
1938a; Aoki 1975, as Pemphigus
latisensorius). 2n=18 (Blackman 1986). Thecabius (Parathecabius) lysimachiae
Borner Galls containing developing
emigrant alatae are formed by folding and convoluting the leaf of Populus nigra (fig. 131J).
The fundatrix inhabits a separate oval pocket-gall on the upper side of the
leaf lamina, as in other Parathecabius.
Alatae (BL 2.3-2.6 mm, with dark, dirty greenish abdomen) migrate in late
June-July to roots and runners of Lysimachia
spp., especially L. nummularia.
Other species, e.g. Lysimachia (= Glaux) maritima, may also be colonised. Apterae on Lysimachia
are brownish or greyish green, with wax; BL 1.1-1.8 mm. Alate sexuparae
return to poplars in September, but anholocyclic
overwintering on Lysimachia is also
common. In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007) and
central Asia, and recently (2000) recorded from New York State, USA (Foottit et al. 2006). T. luppovae (Narzikulov), described from Populus densa (= P.
talassica) in Tajikistan, may be a synonym. 2n=18. Thecabius populimonilis
(Riley) The galls on Populus spp. (angustifolia, balsamifera,
fremontii, trichocarpa) are rows of ovoid, bead-like cells, pale green or
with a reddish tinge, developed from upper side of leaf lamina on both sides
of mid-rib (see influentialpoints.com).
Each gall in spring contains one developing aptera or alata. The fundatrices
inhabit similar, solitary galls. This species is apparently facultatively
monoecious; Gillette (1913) suggested that the large alatae (BL 2.7-3.0mm)
produced in individual galls in June-July might recolonize cottonwoods, but
it seems more likely that they are emigrants and found colonies of apterous
exules in wax on Salix roots such
as those observed by Lange (1943) in California. Presumably alatae produced
in these colonies in October are sexuparae and return to Populus. However, alate sexuparae (BL 1.7-2.0mm) are also
produced in galls on Populus, up to
12 per gall, in August-October (Maxson & Knowlton 1929, Palmer 1952,
Harper 1959a). A more recent study by Aoki et al. (1996b) has confirmed most of these observations; they
found heteroecious and monoecious life cycles at different locations, but
their data on sex ratios indicate that some gene flow is probably occurring,
rather than that two separate species are involved. Pike et al. (2012) have also now confirmed by DNA barcoding that
colonies on willow roots (probably Salix
dodgeana) in Washington are T.
populimonilis. This species is
widely distributed in USA, western Canada and Mexico. [Pemphigus saliciradicis may be a synonym; see note under that
species.]
Four oak-feeding species, three in Europe and the
Mediterranean region, and one in North America. Remaudiere (1983b) provided a
key including characters of alatae as well as apterae, Ilharco (1966a) keyed
first instar nymphs, and Wieczorek et
al. (2013) provided key couplets to distinguish between the oviparae and
males of T. dryophila and T. suberi. Thelaxes californica
(Davidson) Appearance in life is not
recorded, but presumably much like T.
dryophila; BL of aptera 1.2-2.0mm. On Quercus
spp. in western North America, from British Columbia to Mexico. Records from
Turkey (e.g. Görür et al. 2011a) should probably be applied to T. suberi. Monoecious holocyclic; sexual
morphs aestivate in first instar (Davidson 1917, as ?dryophila),
adult oviparae occur in September-October (Palmer 1952). 2n = 12*. [Note:
there is an important error in the printed version of Key F to the aphids on Quercus in Blackman & Eastop
(1994, pp. 430-431); the first proposition in couplet 5 should refer to T. californica, the second to T. suberi.] Thelaxes dryophila
(Schrank) Apterae are broadly oval,
rather flattened, dark brownish-red to purplish-grey with a paler spinal
stripe (see
influentialpoints.com/Gallery); BL 1.1-2.3 mm. Alatae have dark
marginal sclerites and dorsal cross-bands on posterior abdominal segments.
Ant-attended colonies form at tips of shoots, extending onto stems, leaf
petioles and along mid-ribs on undersides of leaves. On Quercus spp. in Europe, the Mediterranean region and south-west
Asia. Monoecious holocyclic; wax-coated sexual morphs aestivate from June to
September on undersides of leaves in forks of veins (Polaszek 1986), and
under cynipoid marble galls. Lombard et
al. (1988) studied composition of the honeydew. 2n = 8. Thelaxes suberi
(del Guercio) Plate 4a, b Apterae vary in colour from pale green to
dark brown (see influential
points/Gallery); BL 0.9-1.5 mm. Living abundantly on young shoots, leaves
and developing acorns of numerous Quercus
spp. (but not robur) in England,
Wales (Baker 2009), southern Europe, Mediterranean region and southwest Asia.
Also recorded from Castanea sativa
(in Italy). Monoecious holocyclic, but the life cycle is unstudied; Polaszek
(1986) collected presumed sexuparae on Q.
cerris in England in November-December, and Wieczorek et al. (2013b) described single
specimens of the ovipara and small apterous male collected in Turkey in early
November. 2n = 8. [Note: there is an important error in Key F to the aphids
on Quercus in Blackman & Eastop
(1994, pp.430-431); the first proposition in couplet 5 should refer to T. californica, and the second to T. suberi.] Thelaxes valtadorosi
Remaudičre Apterae are light brown,
somewhat waxy; BL 1.0-1.9 mm. On Quercus
spp. in Greece (Remaudiere 1983b), and also recorded from Turkey (Şenol et al. 2014a, in abstract only). The life
cycle is unknown. 2n = 8.
About.30
maculate aphid species with a knobbed cauda living on Leguminosae/Fabaceae of
the tribes Trifoliae, Loteae, Galaeagneae and Coronillae. Half the species
are known only from south-east Europe and the Middle East, four extend into
north-west Europe, and four or five extend across Russia, one as far as
Japan. The world fauna was revised by Quednau (2003), who provided keys and
illustrations of all available morphs. He raised Bicaudella to the status of a full genus, and introduced a new
subgeneric classification that is followed here. Earlier accounts are
available for Europe (Pintera 1957a), Hungary (Szelegiewicz 1969b),
north-west Europe (Heie 1982), the Iberian peninsula (Nieto Nafrěa & Mier
Durante 1998) and the Middle East (Remaudičre 1989c). Therioaphis (Pterocallidium) aizenbergi Ivanoskaya
& Tomilova Apterae (and alatae)
are yellowish, covered dorsally with numerous large dark spots; BL c.1.4-1.7
mm. On Caragana arborescens in Siberia and Mongolia. Monoecious
holocyclic; all the morphs are included in the original description. Therioaphis alatina Hille Ris Lambers & van den Bosch Apterae are pale dirty greenish yellow
with dark brown markings; BL 1.3-2.2 mm. On Ononis spp. in France, Italy and Switzerland. Although all adult
viviparae have wings, these are often small and non-functional (original
description). Sexual morphs occur in October. Therioaphis
(Rhizoberlesia) arnaultae Remaudičre Apterae are
probably pale yellowish with variably developed segmental markings; BL c.1.7 mm. On Astragalus sp(p). in Iran, Turkey
and Lebanon. Oviparae and alate males were present in November (Remaudičre
1989c). Remaudičre & Talhouk (2000) provided further information on
morphology of apterae. Therioaphis astragali (Dzhibladze)
Apterae seem not to occur, alatae often being brachypterous; alatae
are yellowish with black dorsal abdominal spots, and dark spots at ends of
wing veins, BL 1.5-1.9 mm. On Astragalus
caucasicus in Georgia, and also recorded from Astragalus sp(p). in Azerbaijan, Iran and Turkey. Oviparae and
alate males occur in October (original description). Therioaphis (Pterocallidium) azerbaidjanica Remaudičre Apterae are probably yellowish,
BL 1.3-1.7 mm. On an Astragalus sp.
with long, pointed leaves in Iran (Tabriz). Alatae and sexual morphs are
unknown. Therioaphis beijingensis Zhang
Apterae are unknown, alatae are probably yellow with four rows of dark
dorsal abdominal spots (two spinal, two marginal), and dark siphunculi; BL c.
2.2 mm. On Caragana frutex in
China. Apart from the pigmentation this species seems close to T. tenerum. Therioaphis (Pterocallidium) bonjeaniae Hille Ris Lambers &
van den Bosch (Fig.40g) Apt. whitish to yellowish; BL 1.4-1.8 mm.
On undersides of leaflets of Lotus
(Dorycnium) spp. (hirsutus,
rectus) in France (Alpes
Maritimes), Spain and Portugal. The life cycle is unknown; parthenogenetic
reproduction continued into November (original description). Sousa Silva
& Ilharco (2016) provided a redescription and morphometric data for
aptera and alatae collected in Portugal. Therioaphis (Rhizoberlesia) brachytricha Hille Ris Lambers &
van den Bosch (Fig.40c) Apterae are very pale buff to pale
yellowish with dusky spots; BL 1.3-2.0 mm. On upper sides of leaves of young growth of Lotus spp., especially L.
corniculatus growing in warm, dry positions. It can also feed on Trifolium pratense (BMNH collection,
leg. H. Szelegiewicz, and see original description). Widely distributed in
Europe, east to Iran. Monoecious holocyclic with alate males. Therioaphis cana Zhang, Chen, Qiao & Zhong Only alatae are described; pale grey and
leaf green in life, BL unrecorded. On Medicago
sativa in Xinjiang province, China (G. Zhang 1999). This species could
not be included in the key to aphids on Medicago
on the basis of the published description. Therioaphis dorycnii (Pintera)
(Fig.40a) Apterous viviparae do not occur; BL of al.
c. 1.7-1.8 mm. On Lotus (Dorycnium) spp. in eastern Europe (Czech Republic, Hungary) and Iran.. A
record from Ukraine (Mamontova-Solukha 1963) may be referable to Th. litoralis. Pintera (1957a)
described the ovipara. Therioaphis hungarica Szelegiewicz
Apterae are yellowish to orange with dark brown spots; BL 1.4-1.9 mm.
Described from a reddish-flowered Ononis
sp. in Hungary, and later collected on Ononis spinosa in Czech Republic (BMNH collection, leg. J.
Holman). It was placed as a subspecies of Th.
ononidis in Quednau 2003, but reinstated as a full species by Eastop
& Blackman (2005). Therioaphis (Pterocallidium) kermanica Remaudičre Apterae are yellowish with black dorsal
spots, BL 1.5-2.0 mm. On Astragalus
pseudosquarrosus (= squarrosus) in
Iran. Therioaphis (Pterocallidium) khayami Remaudičre
(Fig.16f) Apterae are yellowish with extensive dark
dorsal markings, and also usually with some dark submarginal sclerites
ventrally on abdomen; BL 1.3-1.6 mm. On Astragalus
sp. in Iran, and Afghanistan (BMNH collection, leg. R. van den Bosch). Therioaphis kundurensis Quednau
Apterae are yellow to yellowish green; BL 1.4-1.9 mm. On undersides of
leaves of Caragana spp., attended
by ants (Quednau 2003). In east Siberia. Therioaphis
(Rhizoberlesia) langloisi Remaudičre & Leclant
Apterae are golden yellow with brown transverse bands; BL 1.4-1.65 mm.
In small colonies on undersides of older leaves of Coronilla glauca in the south of France
and Spain (Mediterranean region), where it is anholocyclic (original
description). Therioaphis
litoralis Hille Ris Lambers &
van den Bosch ( = Therioaphis hillerislambersi Szelegiewicz; synonymy by Quednau,
2003) (Fig.40e) Apterae are bright yellowish to ochreous
yellow; BL 1.4-1.8 mm. On Lotus and
related genera (Anthyllis, Dorycnium). In Europe; southern
France, Hungary, Czech Republic, Yugoslavia, Greece, Ukraine (Mamontova-Solukha
1963, as dorycnii) and Iran. An
ovipara was found in October (original description). Therioaphis
loti Hille Ris Lambers &
van den Bosch (Fig.40b) Apterae viviparae do not occur; BL of alata
1.4-1.7 mm. On Lotus (incl. Dorycnium) spp. in Israel, Iran and
Turkey. Therioaphis
(Pterocallidium) luteola (Börner) (Fig.53b) Apterae are pale yellow or whitish; BL
1.7-2.0 mm. On Trifolium pratense in
west, north, central and eastern Europe. Monoecious holocyclic with alate
males (Heie 1982). Therioaphis
natricis Hille Ris Lambers &
van den Bosch Colour of apterae in
life is unrecorded; BL 1.25-2.2 mm. On Ononis
spp. (especially natrix) in the Middle East (Israel, Lebanon,
Iran). 2n=16. Therioaphis
obscura Hille Ris Lambers &
van den Bosch (Fig.40d) Apterae are banded grey with black spots; BL
1.3-1.8 mm. On undersides of leaves of Lotus
and Dorycnium spp. in France,
Spain and Italy. There are also records from Ononis spp. Oviparae
and alate males occur in October-November (original description), but it is
also partially anholocyclic (Leclant 1966). Therioaphis
ononidis (Kaltenbach) Apterae are yellowish to orange with
stiff black hairs with tubercular bases arising from large brown spots (see influentialpoints.com/Gallery);
BL 1.8-2.2 mm. On reddish-flowered Ononis
spp. in Europe, the Middle East and Central Asia. Records from North
America and Japan, and from other genera of Leguminosae, are likely to be
misidentifications of T. trifolii
(Quednau 2003). Oviparae and alate males occur in October. 2n=16. Therioaphis
pteromaculata Quednau Only known from alatae, colour in life
unknown; BL 1.8-2.0 mm. On Astracantha
adscendens in Iran (Quednau 2003). Therioaphis
(Rhizoberlesia) riehmi (Börner) Alatae are yellow or orange with dark
abdominal markings, the alatoid nymphs being paler yellow with four rows of
dark dorsal spots (see influentialpoints.com); BL 2.0-2.7 mm. Apterae are very rare, and have
not been described, but one is illustrated by Quednau (2003). On undersides
of leaves of Melilotus spp., the
preferred hosts, but there are also records from species of Medicago, Trigonella and Trifolium.
In Europe, Algeria (Laamari et al.
2013), Middle East, India (Chowdhuri et
al. 1969), Kazakhstan (Kadyrbekov 2009a), China (1 alata, BMNH
collection. leg. L.A. Mound); introduced to North America, where it is
widespread, and South America (Argentina: Ortego et al. 2004; Chile: Nieto Nafría et al. 2018). Monoecious
holocyclic with oviparae and alate males in September (Hille Ris Lambers
& van den Bosch 1964). 2n=16. Therioaphis
(Pterocallidium) subalba Börner (Fig.53a) Apterae are pale yellow, whitish
anteriorly; BL 1.7-2.0 mm. On Trifolium spp. in northern, central and eastern Europe, and Turkey (Şenol et
al.
2014); there are also records from Medicago sativa and Dorycnium (= Lotus) sp., and from Melilotus
suaveolens in Korea (Quednau 2003). Therioaphis
tenera (Aizenberg) Apterous viviparae seem not to
occur. Alatae are yellowish, the
dorsal abdomen with variably developed dusky spots, and the siphunculi are
pale (see
influentialpoints.com); BL 1.7-2.2 mm.
On Caragana spp., living
dispersed on undersides of leaflets, mostly on lower branches. In Europe
(Finland, Germany, Hungary, Czech Republic, Russia, Ukraine), Central Asia
(Kadyrbekov 2017a) and introduced to Quebec, Canada (Quednau 2003). The
record from east Siberia is referable to Th.
kundurensis. A short-haired
population on C. frutex in Ukraine
was distinguished as a subspecies, T. tenera ssp. frutex Bozhko, and a related, more pigmented form is
described from China as T. beijingensis
(q.v.). Oviparae and alate males were described from Finland (Heikinheimo
1997); fundatrices are present in June and oviparae in late
August-September. 2n=6. Therioaphis
(Pterocallidium) trifolii (Monell) Yellow
Clover Aphid, Spotted Alfalfa Aphid, Spotted Clover Aphid Plate 4c (Fig.16g, Fig.40f,h) Apterae
are pale yellow, greenish white to almost white, rather shiny, with rows of
small light- or dark-brown pigmented raised spots (see aphids
of Karnataka website); BL
1.4-2.2 mm. On many plants of Leguminosae/Fabaceae in genera Astragalus, Lotus, Medicago, Melilotus, Onobrychis, Ononis and Trifolium. Originally palaearctic;
Europe, North Africa, Middle East, Central Asia, India, Pakistan, Japan
(Sorin 1990c), and China (Zhang et al.
1999, as T. cana). Introduced
to North and South America, South Africa, and Australia. Recognisable forms
with more specific host-plant relations occur within the species, and are
particularly evident where separate introductions have occurred to countries
outside Europe. Some of these are regarded as distinct forms or subspecies,
including the Spotted Alfalfa Aphid, T. trifolii ssp. maculata (Buckton), introduced to North and South America,
Australia and New Zealand (for further details see Blackman & Eastop 2000
and 2007). T. trifolii ssp. ventromaculata
Müller occurs on Astragalus
onobrychus in Austria, and is also recorded from Poland and Czech
Republic (Remaudičre 1989c). T. trifolii ssp. albae Bozhko was described from Melilotus albus in Ukraine, and seems close to the nominate
species (Quednau 2003). Monoecious holocyclic, with alate males, in cold
temperate climates of northern and central Europe and more northerly USA,
anholocyclic in warmer regions. 2n=16 (incl. ssp. maculata).
Most species originally described in Thoracaphis have been removed to other
genera, leaving the type species (arboris
van der Goot) and a few others, mostly little known, which can hardly be
said to form a homogeneous group. One species (kashifoliae) placed in a separate genus (Xenothoracaphis) in Blackman & Eastop (1994), is transferred
to Thoracaphis following Remaudičre
& Remaudičre (1997). Probably they were associated in the past with Distylium as primary host, as are
other Nipponaphidini, but now only known from parthenogenetic generations on
what were the original secondary hosts (Quercus,
Lindera). Thoracaphis arboris
van der Goot Apterae are
aleyrodiform, flattened, broadly oval, bluish-black with wax frosting on
sides of body only; BL 1.3-1.6 mm.
Immatures are dull greyish-green with rather more wax. On undersides
of leaves of Quercus sp(p). in
Java. Alatae collected in May and August (Noordam 1991). The life cycle is
unknown; presumably anholocyclic. Thoracaphis flava
Takahashi Apterae are aleyrodiform,
flattened, broadly oval, only slightly sclerotized, pale yellowish-brown; BL
c. 0.5 mm. On undersides of leaves of Quercus
sp. in Malaya (Takahashi 1950), and recently reported from Turkey (Şenol
et al.
2014a), although this requires further confirmation. Other morphs and life cycle
are unknown; presumed to be anholocyclic. This species clearly does not
really belong in Thoracaphis;
possibly it requires a new genus. Thoracaphis kashifoliae
(Uye) Apterae black, strongly sclerotized, covered with powder; BL c. 1.5 mm.
Alatae are black with blackish subcosta, pterostigma and veins in forewing.
On upper sides of leaves of Quercus
glauca and Q. acuta in Japan.
Anholocyclic, with alatae appearing in November (Takahashi 1958b; as Xenothoracaphis kashiwae). Kurosu et al. (2016) reported the occurrence
of “defensive nymphs”; first instars of the alate morph with fore- and
mid-legs enlarged to a varying extent, and observed to attack lepidopteran
larvae. Thoracaphis
kumaoni Chakrabarti & Debnath Apterae are aleyrodiform, flattened, light
to dark brown; BL 1.4-1.5 mm. On Quercus
incana in northwest Himalaya, India (Chakrabarti & Debnath 2011).
Other morphs and biology are unknown. Thoracaphis
linderae Shinji
Apterae are aleyrodiform, flattened, broadly oval, green or brown becoming
sooty black with age; BL 1.5-2.0 mm. On twigs of Lindera spp. in Japan (Shinji 1941). Alatae were collected in May
(Shinji 1926). The life cycle is uncertain; Shinji (1926) reported
overwintering as eggs on Lindera,
but this was not confirmed by Takahashi (1958a) in his redescription of the
species. Thoracaphis
quercifoliae A.K. Ghosh Apterae are flattened, subcircular, brown
to dark brown sclerotic; BL 1.4-1.5 mm. On ?leaves of Quercus dealbata in Meghalaya, India (A.K. Ghosh 1988). Alatae
(with secondary rhinaria distributed ANT III 20-22, IV 15-17, V 7-10) were
collected in late March. The life cycle is unknown. This species perhaps
belongs in Nipponaphis. An aphid
collected on a plant of Lauraceae in China (Fujian) and used in DNA studies
(Huang et al. 2012) was identified
as this species. Thoracaphis
sp. on Cinnamomum Apterae are aleyrodiform, flattened oval,
reddish-brown with creamy-white markings on prosoma and blackish abdominal
plate. On bark of stems of Cinnamomum
japonicum in Japan (Moritsu 1983; as ?cinnamomiae
Shinji). This appears to be an undescribed species.
About
eight long-bodied, sedge-feeding aphids related to Subsaltusaphis but with pointed dorsal hairs, and often with wax pores.
Empodial hairs are spatulate. Apterae often have rhinaria on ANT III, and
alatae usually have extensive dark dorsal cross bands, sometimes merging into
a solid patch. Quednau (2010) reviewed
the world fauna, with keys and illustrations of all available morphs; his
review made substantial changes to the nomenclature, giving Allaphis Mordvilko full generic status
(q.v.), making Neobacillaphis
Huculak a subgenus of Thripsaphis,
and sinking several names in synonymy. Heie (1982) reviewed the species in north-west
Europe, and Nieto Nafría & Mier Durante (1998) reviewed those on the
Iberian peninsula. Thripsaphis ballii (Gillette)
Plate 4i (Fig.19f) Apterae are pale greyish green with
extensive darker cross-bands and marginal areas, lightly wax-dusted; BL
1.9-2.5 mm. Alatae have 6-8 secondary
rhinaria on ANT III, and dorsal abdomen with broad dark transverse bars that
are sometimes partially fused across tergites 3-5. Described from Carex nebraskensis in western USA,
producing oviparae and apterous males on this plant in October-November
(Palmer 1952). Similar aphids also
occur on other Carex spp. in
eastern USA and Canada, Europe, Pakistan, east Siberia and China. Quednau
(2010) recognised four subspecies, describing one as new. Unfortunately none
of them have a clearly defined specific host relationship or geographical
distribution. The nominate subspecies T ballii ssp. ballii is recorded from North America and Pakistan. T.
ballii ssp. caespitosae Ossiannilsson, described from C. cespitosa in Sweden as paler, dirty
yellowish in life and producing its sexuales in September (original
description), and also recorded from Finland (Albrecht 2010) and Quebec,
Canada, is considered by Quednau to have a holarctic distribution on various Carex spp. T. ballii ssp. longisetis Richards
was described from Canada, and populations in east Siberia (Quednau &
Shaposhnikov 1988) with similarly long hairs on the head and abdominal
tergite 8 are also regarded as this subspecies. However there is intergrading
with the nominate subspecies (Quednau 2010), and these hairs show almost continuous length
variation in BMNH specimens. The fourth subspecies, T. ballii ssp. pennsylvanica Quednau from Carex sp. in Pennsylvanica, also has
longer hairs, but has apterae with extensive dorsal sclerotisation (Quednau
2010). 2n=8 (for ssp. pennsylvanica). Thripsaphis (Larvaphis) brevicornis (Ossiannilsson) Apterae are very slender, probably
yellowish with brown markings; BL 2.7-3.0 mm. Alatae are apparently unknown.
In leaf sheaths of Carex elata and C. lasiocarpa, often under water. In northern Europe (Sweden,
Finland, Germany). Oviparae and apterous males occur in September-October
(original description and Ossiannilsson 1959). The very short, 5-segmented antenna of this
species are distinctive, and warrant the separate subgenus. Thripsaphis caricis (Mordvilko) (= Thripsaphis
gelrica Hille Ris Lambers; synonymy by Quednau 2010) Apterae are dark greyish brown, clothed
with bluish grey wax powder; BL 2.5-2.8 mm.
Alatae have 25-31 rhinaria on ANT III, and dorsal abdomen with broad
dark bars separated between tergites. On Carex spp. (rostrata, vesicaria)
in northern and central Europe (Heie 1982, as caricicola). [Quednau
(2010) pointed out that the name caricicola Mordvilko 1921 should be
treated as a synonym of Allaphis producta (Gillette).] Thripsaphis polytricha Quednau Colour in life unknown,
BL of apterae. c.2.7 mm. Alatae are unknown. On Carex sp. in
Idaho, USA (Quednau 2010). Thripsaphis sensoriata Hille Ris Lambers Only known from one alata, collected away
from its host plant (presumed to be a Carex
sp.) in Greenland. Thripsaphis (Neobacillaphis) striata (Bozhko) Apterae are orange with a pair of broad
brown longitudinal pleural stripes; BL c.1.4 mm. On Carex distans in southern Russia (Ciscaucasia). Thripsaphis (Neobacillaphis) szelegiewiczi
Huculak Plate 4f Apterae are brown with darker pleural and
marginal longitudinal bands; BL 2.2-2.6 mm. Alatae have 9-13 rhinaria on ANT
III. On Carex spp. in northern
Poland. A population on Carex vulpina in
the Ukraine is regarded as a subspecies, T. (N.) szelegiewiczii ssp. carpaticae (Mamontova), although
possibly only a seasonal variant of T.(
N.) striata (Quednau 2010). Thripsaphis unciniae Quednau
Colour in life unknown, BL of aptera c.1.9-2.1 mm. Alatae have 4-8
secondary rhinaria on ANT III. On Uncinia sp. and Carex sp. in Chile.
Four oriental species very similar to Eucallipterus, but with larger, pale siphunculi
and more extensive forewing pigmentation. Accounts are available for Japan
(Higuchi 1972), China (Zhang & Zhong 1982c, Qiao & Zhang 2003a) and
east Siberia (Pashchenko 1988a; Quednau & Shaposhnikov 1988). Quednau
(2003) keyed the species and illustrated all available morphs. Tiliaphis
coreana Quednau
All viviparae are alate, greenish-yellow to yellow with brown-black
sides to head and prothorax and longitudinal stripes on mesothorax, becoming
more evident in late summer; BL 2.5-3.5 mm. Forewings have a deeply pigmented
costal margin and a black zig-zag line between distal ends of veins.
Immatures are green with 4 rows of black spots. In rows along veins on
undersides of leaves, or as nymphs on upper sides of leaves, of Tilia spp. in Korea, China and eastern
Siberia. Sexual morphs occur in September (Paik 1972, as Tiliaphis japonica; Quednau 1979, Quednau & Shaposhnikov
1988). 2n = 38 (Chen & Zhang 1985b). [A DNA sequence assigned to T. shinae (Genbank accession
no. GU978821) should be reassigned to T.
coreana (Y. Lee et al. 2017).] Tiliaphis pseudoshinae
Quednau All viviparae are alate,
whitish with brown-black markings on body and wings as in coreana, but
without longitudinal stripes on mesothorax and with clear base to forewing;
BL 2.0-2.6 mm. On Tilia sp. in
Korea and eastern Siberia. Sexual morphs occur in September (Quednau
1979,Quednau & Shaposhnikov 1988). Tiliaphis
shinae (Shinji)
All viviparae are alate, pale yellow to greenish-yellow with dark
brown markings on body and wings as in pseudoshinae;
BL 1.7-2.4mm. On undersides of leaves of Tilia
spp. in Japan, China, Korea and eastern Siberia. Sexual morphs occur in early
October (Takahashi 1961b, Higuchi 1972). 2n = 14 (Shinji 1931). [A DNA sequence assigned to T. shinae (Genbank accession
no. GU978821) should be reassigned to T.
coreana (Y. Lee et al. 2017).] Tiliaphis shinjii
Higuchi All viviparae are alate,
colour in life not recorded, dorsal dark markings as in coreana but with paired dark spinopleural patches on abdominal
tergites; BL 2.0-2.7 mm. On Tilia
spp. in Japan, Korea and eastern Siberia.
It was recently reported to occur in Turkey (Şenol et al. 2014b), but the identification is suspect because this report
refers to apterous rather than alate aphids. Sexual morphs
were found in September (Higuchi 1972, Quednau & Shaposhnikov 1988).
One North American species, differing from Prociphilus in having alatae with many
more secondary rhinaria on the antennae. Tiliphagus lycoposugus Smith (fig. 115A) Large leaf-nest galls are produced in
spring by clumping and cupping terminal leaves so as to resemble a large,
poorly-formed head of lettuce, 10-30 cm in diameter. On Tilia americana in eastern USA (Maryland, North Carolina).
Fundatrices inside galls are brown, globose, BL 4.1-5.3 mm. Heteroecious
holocyclic; emigrant alatae are reddish-brown, BL 2.6-3.3 mm, leaving gall in
June-August to found colonies on the tuberiferous
roots of Lycopus virginicus. Apterae on Lycopus
roots are white to pale tan or flesh-coloured, with wax; BL 2.6-2.8 mm. Return
migration of sexuparae to trunks of Tilia
occurs in October-November (Smith 1965).
About 20 mostly small species usually associated
with Ulmaceae, although species have also been described from Lythraceae (Lagerstroemia, Duabanga), and from Corylus,
Dalbergia and Sapindus. Woody Leguminosae/Fabaceae seem to be recorded quite
regularly as alternative or 'casual' hosts. All viviparae are alate and
usually have paired spinal and marginal tubercular processes. Some species
have conspicuous black markings on dorsal body and/or forewings. Problems
with the taxonomy of Tinocallis may
arise from seasonal variation; alatae in summer and autumn are often more
pigmented than in spring and may have fewer secondary rhinaria. For example,
specimens of Tinocallis platani collected
in May have 17-25 (usually 20-23) rhinaria on ANT III, whereas specimens
collected from the same trees in July-October have 7-21 (mostly 11-17)
rhinaria on III; (V.F. Eastop, unpublished data). Quednau (2001) reviewed the
world fauna, and regional accounts are available for Britain (Stroyan 1977),
Fennoscandia and Denmark (Heie 1982), Italy (Patti & Barbagallo 1998),
Europe (Kanturski et al. 2018),
India (Chakrabarti 1988, A.K. Ghosh & Quednau 1990), eastern Russia
(Pashchenko 1988a), China (Qiao et al.
2005b), Korea (Lee & Lee 2017) and Japan (Higuchi 1972). Quednau (2003)
further revised the genus, illustrating all available morphs, erecting two
new subgenera (Orientinocallis) and
(Eotinocallis), and restoring three
taxa treated as subgenera in Blackman & Eastop (1994), Pseudochromaphis, Quednaucallis and Sarucallis, to full generic rank. Tinocallis (Orientinocallis)
dalbergicola Quednau Alatae are blackish in life (J.H. Martin,
pers. comm.); BL 1.9-2.2 mm. Only known from Hong Kong, on Dalbergia hancei (Quednau 2001). The
sexual morphs are unknown. Tinocallis (Orientinocallis) distincta
Ghosh, Ghosh & Raychaudhuri
Alatae are pale (colour unrecorded), with dark antennae, tibiae,
dorsal abdominal tubercles, siphunculi and cauda; BL 1.8-2.2 mm. Described
from an unidentified leguminous plant in West Bengal (M.R. Ghosh et al. 1971); Chakrabarti (1988) lists
other records from West Bengal, mostly from undetermined plants, but
including one sample collected on Duabanga
sonneratioides. Oviparae and alate males occur in December-January, but
alate viviparae are also present during winter months, so apparently in West
Bengal there is only a partial holocycle. Tinocallis (Orientinocallis)
himalayensis Ghosh, Ghosh & Raychaudhuri Alatae are greyish or greenish, somewhat
waxy (J.H. Martin, pers. comm.); BL 1.2-2.0mm. Forewings usually have dark
spots at ends of veins and the branches of the media are brown-bordered
distally. Described fom an unidentified leguminous plant (A.K. Ghosh et al. 1971e); specimens were
subsequently collected in India (West Bengal, Sikkim) on Peltophorum fleragineum (Leguminosae) and Duabanga sonneratioides (Chakrabarti 1988). Also found on the
undersides of leaves of Lagerstroemia
macrocarpa (= speciosa) in
Thailand (Danielsson & Robinson 1978, as T. khonkaensis), and on Lagerstroemia
sp(p). in east and south-east Asia (Bangladesh, Thailand, Malaya,
Singapore) (BMNH collection, leg. various collectors). The life cycle is
unstudied; alate viviparae and apterous oviparae occur together in
December-January in both northern India and Thailand, indicating that there
is only a partial holocycle. Males are unrecorded. Tinocallis insularis
(Takahashi) Alatae are
greenish-yellow with black-banded antennae; BL c. 1.6-1.7 mm. On undersides of
leaves of Sapindus mukorossi in
Taiwan (Takahashi 1927a) and Hong Kong (Martin & Lau 2011), and on Sapindus sp. in Japan (Higuchi 1972).
The sexual morphs are unknown. Also recorded from Pterocarya stenoptera (Tao 1964), Aesculus chinensis (Tao 1999) and Magnolia sp. in Meghalaya, India (A.K. Ghosh & Raychaudhuri
1972b, as T. magnoliae), but these
may all be vagrants. Tinocallis latifoliae Lee
& Lee Alatae are very pale yellow
with small paired black sppots on ABD TERG 3-7, and appendages mainly pale except
for contrastingly black joints of ANT III-VI; BL 1.2-1.5 mm. On Zelkova serrata var. latifolia in mountain areas of South
Korea (Lee & Lee 2017). Closely related to T. zelkowae, its status as a separate species being supported by
DNA barcoding (Y. Lee et al. 2017). Tinocallis mushensis
(Takahashi) Alatae are pale yellow,
with three longitudinal thin brown stripes on head and pronotum; BL c. 2 mm.
On young leaves of Zelkova formosana
in Taiwan (Takahashi 1925). The sexual morphs are unknown. Tinocallis (Sappocallis) nevskyi
Remaudiere, Quednau & Heie Alatae
are pale yellow or whitish, with very little pigmentation in spring and
summer populations, except for a black spot at the apex of the hind femur (see
influentialpoints.com/Gallery); BL 1.4-2.1 mm. Head and
mesonotal lobes may be dark in autumn populations. On Ulmus spp. in central and south-west Asia (Afghanistan, Iran, Turkey,
Georgia, Kazakhstan, Tajikistan, Uzbekistan, Pakistan, northern India). In
recent times it has invaded and dispersed extensively in Europe; England
(Hopkins 1997), Wales (Baker 2009), Belgium, Netherlands, France, Denmark,
Germany, Sweden, Finland (Albrecht 2010), Poland, Hungary, Czech Republic,
Switzerland, Italy (Patti & Barbagallo 1998). It is also recorded from Zelkova carpinifolia (= crenata), although Remaudičre et al. (1988) thought that the
occurrence of sexual morphs on this plant in Iran was 'accidental'. Oviparae
and alate males occur in November. Remaudičre et al. (1988) discussed differences from T. saltans, with which it has been confused by some authors. Tinocallis (Sappocallis) nikkoensis
Higuchi Alatae are pale yellow with
pale or dusky antennae and legs; BL c. 1.5mm. On Corylus spp. (heterophylla,
sieboldiana) in Japan (Higuchi
1972). Biology and sexual morphs are unknown. [Paik's (1965) record of T. zelkowae on Corylus sieboldiana in Korea should perhaps be referred to this
species. T. sapporoensis Higuchi,
which Blackman & Eastop (1994) suggested to be a midsummer population of T. takachihoensis, was made a synonym
of T. nikkoensis by Quednau
(2001).] Tinocallis (Eotinocallis) platani
(Kaltenbach) Plate
l0b Alatae are yellow to
greenish-white with extensive black-brown markings on head, thorax, abdomen
and forewings, and black siphunculi; BL 2.0-2.2 mm. On undersides of leaves
of Ulmus spp., especially U. laevis. Throughout Europe, across
Asia to eastern Siberia (Pashchenko 1988a) and China (Qiao et al. 2005b), and introduced into
western North America, where it occurs on U.
americana. Oviparae and alate males occur on U. laevis in Europe in October. Olkowski et al. (1982) reported biocontrol with an introduced parasitoid
in California. Patti & Barbagallo (1998) provided a recent account with
colour photographs. Tinocallis (Sappocallis) saltans
(Nevsky) (figs 116B,D) Alatae are yellow to orange-yellow, with
brown head and thorax even in early summer, with a more extensive distal
black patch on the hind femur and more forewing pigmentation than in nevskyi; BL 1.4-2.2 mm (Nevsky 1929b, Quednau
1979). On Ulmus spp. in Spain
(Núńez-Pérez et al. 1991), Italy
(Patti & Barbagallo 1998), eastern Europe (Hungary, Romania) and central
and east Asia (eastern Iran, Tajikistan, Kazakhstan, Uzbekistan, Urals,
Afghanistan, Pakistan, Siberia, Korea and China). Also recorded from Zelkova serrata in Hungary (Ripka et al. 1998). Fundatrices collected in
April and oviparae and alate males collected in October were described by
Núńez-Pérez & Mier Durante (1993). Introduced into North America (Halbert
& Pike 1990), Argentina (La Rossa et
al. 1990) and Bolivia (cited by Naumann-Etienne & Remaudičre 1995).
[Some accounts under this name from north-west Europe (e.g. Heie 1982) and
south-west Asia (Richards, 1967) should be referred to T. nevskyi (Remaudičre et al.,
1988).] 2n = 16 (in China; Chen & Zhang 1985b). Tinocallis sophorae Zhang Colour of alatae in life is unrecorded; BL
c. 1.7 mm. Described from Sophora
japonica and Zelkova schneideriana
in Jiangsu, China (Zhang & Zhong 1980c). Tinocallis suzhouensis
Zhang Alatae are dusted all over with
white powder; BL c.2.2cm. On Zelkova
schneideriana in China (Zhang & Zhong 1980c). Sexual morphs and life
cycle are unknown. Tinocallis (Sappocallis)
takachihoensis Higuchi (fig. 51B) Alatae are pale yellow-green with shiny
black head and thorax, black distal section of hind femur and base of hind
tibia and black markings on the wings (see
influentialpoints.com/Gallery); BL 1.8-2.0 mm. Recorded from Ulmus spp. in Japan (Higuchi 1972),
China (Tao 1999) and eastern Siberia (Pashchenko 1988a, as T. ussuriensis), and also from Hemiptelea davidii in China (Zhang & Zhong 1980c, as T. hemipteleae). Introduced to Europe
and North Africa, where it is recorded from Ulmus spp. in southern France (Quednau & Shaposhnikov 1988),
Germany, Netherlands (Piron 2013), Denmark, Czech Republic and
Poland (Kanturski et al. 2018a),
Serbia ( et al.2018), Italy
(Patti & Barbagallo 1998), Andorra (Mier Durante & Pérez Hidalgo
2002), Spain and the Balearics (Pérez Hidalgo et al. 2013), Malta
(Mifsud et al. 2009), Algeria
(Laamari et al. 2013), Greece (Papapanagiotou et al. 2012) and Turkey (Görür
et al. 2011),
and also introduced to USA (Maryland, first record 1996; Foottit et al. 2006). In England it has
regularly been intercepted since 1997 on bonsai Zelkova serrata and Ulmus
spp. imported from China, but was more recently collected outdoors on Ulmus glabra in Berkshire (Döring
2007). Oviparae and alate males occur
in Sicily from mid-October (Patti & Barbagallo 1998) and an ovipara
collected in France was illustrated by Quednau (2003). [T. nevskyi ssp. lianchengensis
Zhang & Qiao, described from Ulmus
sp. in China (Qiao & Zhang 1998b) is a synonym (Quednau 2001).] 2n =
16*. Tinocallis (Sappocallis) ulmicola
(Matsumura) (fig. 116C) Alatae are greenish-white, with black head
and thorax, black apices of hind femora and bases of hind tibiae, and three
large black patches on each forewing; BL 1.4-1.6 mm. On the undersides of
leaves of Ulmus spp. in Japan,
China, Korea and eastern Siberia (Quednau & Shaposhnikov 1988). Life
cycle and sexual morphs are apparently unrecorded. 2n = 16. Tinocallis ulmifolii
(Monell) Alatae are pale yellow to
greenish, reddish green or dark grey-green with white-bordered dusky
longitudinal streaks on head and pronotum; abdomen with rows of white specks
and dusky spots around hair-bases (see influentialpoints.com
and aphid trek); BL
1.5-2.0mm. On the undersides of leaves of Ulmus
spp. (especially U. americana),
widely distributed in North America. Apterous oviparae and alate males occur
in September-October (Hottes & Frison 1931, Palmer 1952, as Myzocallis ulmifolii); two variants of
the oviparous morph were illustrated by Quednau (2003). Tinocallis ulmiparvifoliae
Matsumura (fig. 116A) Alatae are pale bluish-green, developing
paired longitudinal white wax stripes on head and pronotum, a single, spinal
white stripe on pterothorax and white wax spots on dorsal abdomen (Moritsu
1983). Tips of abdominal tubercles and distal ends of forewing veins are
frequently dark. BL 1.6-2.7 mm. On Ulmus
parvifolia in Japan, Korea, China and Taiwan. Introduced to Australia
(Zeck 1933, as Myzocallis viridis),
USA (California, Florida; see Foottit et
al. 2006), England (on bonsai U.
parvifolia and Zelkova serrata;
Stroyan 1977 and BMNH collection, leg. R. Hammon), Italy (Patti &
Barbagallo 1998) and Spain (Hidalgo & Nieto Nafría 2005). Oviparae have
been recorded from Korea (Paik 1972) and China (in November; illustrated by
Quednau 2003) but males are so far unrecorded. Hales (2020) studied populations
in Australia. 2n = 16*. Tinocallis viridis
(Takahashi) (fig. 119A) Alatae are green, with green eyes, banded
antennae, forewings with dark spots on pterostigma and at distal ends of
veins, dark abdominal tubercles and dusky brown siphunculi; BL 2.2-2.7 mm. On
Zelkova formosana in Taiwan, and on
Z. schneideriana in China (Zhang
& Zhong 1980c, as T. allozelkowae). Life cycle and sexual morphs are unknown.
[This species has been confused in the literature with ulmiparvifoliae.] Tinocallis (Eotinocallis) zelkovae
Dzhibladze (fig. 119B) Alatae have a dark brown head, thorax,
siphuncui and cauda, yellowish-green abdomen with extensive dark brown paired
patches or cross-bands and maculate forewings with extensive pigmentation
around veins; BL 1.4-1.8 mm. Described from Zelkova carpinifolia in Georgia and also collected on Zelkova sp. in Iran (BMNH collection,
leg. R. van den Bosch). Monoecious holocyclic, with apterous oviparae and
alate males (Dzhibladze 1957). Tinocallis
zelkowae (Takahashi)
(fig.
119C) Alatae are shining pale
yellow or greenish-yellow, with pale eyes, banded antennae, a fine brown
mid-dorsal line on head and pronotum, pale yellow legs and siphunculi and a
pair of small brown dorsal spots on each of abdominal tergites 3-7; BL
1.4-1.8 mm. On undersides of leaves of Zelkova
serrata and related species in Japan (Takahashi 1919b), Korea and China,
and introduced to England on bonsai Zelkova
(Stroyan 1979; as T. nirecola), and
to California, USA (Kono 1983). DNA barcoding studies (Y. Lee et al. 2017) revealed that specimens
from Z. serrata var. latifolia in mountain areas of Korea
were another, cryptic, species, now described as T. latifoliae (q.v.).
Records from Ulmus may
mostly be misidentifications of other species, although this aphid does seem to
be recorded frequently, as immature stages as well as adults, on other
plants; especially Betulaceae (Alnus,
Carpinus) and Leguminosae (Robinia, Glycine max and 'cultivated beans'). Monoecious holocyclic in
Japan, with alate males and apterous oviparae in October-November; Inouye
(1968) gives an account of the biology and descriptions of all morphs. 2n =
12*.
One species on Prunus in northern India and Nepal. A.K. Ghosh & Quednau (1990)
gave a full account. Tinocalloides montanus
Basu All viviparae are alate, with
brown head and thorax, pale yellow abdomen with two widely-separated rows of
black spino-pleural patches; BL 2.1-2.6 mm. On undersides of leaves of Prunus spp., especially P. cerasus, in northern India (West
Bengal, Himachal Pradesh) and Nepal. Holocyclic, with apterous oviparae and
alate males in December-January. Sexuales occur on P. persica as well as on P.
cerasus (Agarwala et al. 1984).
2n = 18 (Kurl 1981).
A
genus for 4-5 palaearctic species mostly on Artemisia, similar to Uroleucon
but with very long, robust siphunculi that lack polygonal reticulation.
Pashchenko (1988a) keyed the east Asian species, and Kadyrbekov (2002c) keyed
those of the western palaearctic region. The synonymies given here are
discussed by Eastop & Blackman (2005) Titanosiphon artemisiae (Koch) Plate 28i Apterae are very dark green to black; BL
1.7-2.5 mm. On thin stems of Artemisia spp.,
esp. campestris. Widely distributed
in continental Europe, and also reported to occur in Iran (Rezwani et al. 1994). This species has been
confused with T. minkiewiczi, so
that it is not possible to apply all locality records to the correct species.
Hille Ris Lambers’ (1947a) account applies to a mixture of the two species.
Probably it is monoecious holocyclic. Titanosiphon chondrillae Kadyrbekov
Apterae are pale brown, or with pale brown head and thorax and dark
green abdomen, with dark antennae, legs, siphunculi and cauda; BL 2.1-2.5 mm.
On stems and upper leaves of Chondrilla
sp. in east Kazakhstan (Kadyrbekov 2002c), but with a record also from Saussurea salsa (Kadyrbekov 2017a). Titanosiphon dracunculi Nevsky ( = Titanosiphon
baichengense Zhang) Apterae have
a shiny orange head and thorax, mainly green abdomen, and mainly black
antennae, legs and siphunculi; BL 2.3-3.2 mm.
On flower stems of Artemisia
dracunculus, occurring in large colonies attended by ants (Nevsky 1928a).
In south-west Siberia, Central and eastern Asia. Oviparae and apterous males
occur in September-October (Narzikulov & Umarov 1969). Titanosiphon minkiewiczi Judenko (= T.
benoisti Balachowsky, = T. kazakhstanicum Kadyrbekov) Apterae are very dark green to blackish;
BL 1.8-2.8 mm. Living singly or in small colonies on thin stems and
undersides of leaves of Artemisia spp.,
esp. campestris, in Europe and
Central Asia (recorded from France, Germany, Poland, Lithuania, Belarus, Kazakhstan).
Monoecious holocyclic, with oviparae and apterous males on A. campestris in September-October. Titanosiphon neoartemisiae (Takahashi)
(= T. bellicosum Nevsky, = T. zaisanicum Kadyrbekov; Eastop &
Blackman 2005) Apterae are pale green
to pale brown or reddish, with distal segments of antennae dusky, legs
basally pale with darker femoral apices, tibiae and tarsi, and siphunculi
dark with paler reddish bases; BL 1.7-2.8 mm. (However, specimens in the BMNH
collection have very variable pigmentation, with tibiae sometimes dark but
often quite pale.) Forming large
ant-attended colonies on stems of Artemisia and Seriphidium spp., in south-west, central and eastern Asia
(Narzikulov & Umarov 1969, as T.
bellicosum, Pashchenko 1988a).
Oviparae and alate males occur on A.
scoparia in October-November in Central Asia (Nevsky 1928a, as T. bellicosum), but in Taiwan it is
anholocyclic on A. capillaris (Takahashi
1927). Narzikulov & Umarov (1969) described specimens from Artemisia tenuisecta with particularly
long siphunculi (c.0.7 × BL) as a
subspecies, T. neoartemisiae ssp. hissaricum
(as T. bellicosum ssp. hissaricum). Specimens with similarly
long siphunculi were collected from A.
campestris in Spain (BMNH collection, leg. D. Hollis), and had apterae
with 11-13 secondary rhinaria on ANT III. 2n=8 (for specimens on A. dracunculus in Iran, leg.
V.F.Eastop).
NOTE: The name Toxoptera
has been applied for many years to species resembling Aphis but having a stridulatory apparatus consisting of
ventrolateral ridges on the abdomen and peg-like hairs on the hind tibia
(Eastop 1952). Phylogenetic studies using both molecular and morphometric
methods have now shown that this genus is paraphyletic, that all the species
probably belong within Aphis, and
that the three best known species (aurantii, citricidus and odinae) are not closely related (Kim & Lee 2008, Kim et al. 2010, Wang & Qiao 2009).
Blackman et al. (2011) transferred T. odinae to Aphis. We have followed Lagos et
al. (2014) and Aphid Species File
in placing Toxoptera as a subgenus
of Aphis, although it should
probably eventually be synonymised with Aphis,
further work being needed to establish the nearest relatives of each of the five or six remaining
species.
Medium-sized
to rather large pale brown or whitish aphids living mostly on roots of Compositae,
in association with ants. Many of the nominal species described from Europe
and Asia are little-known and not clearly distinguishable from the type
species (troglodytes). T.
rara is the only species in North America (formerly known there as T. oculata), and was probably introduced from Europe on Taraxacum roots. Subgenus Neotrama
differs from Trama in presence
of siphunculi and compound eyes, but
DNA studies (Normark 1999) show a close
relationship. Eastop (1953), Heinze (1962), Heie (1995), Nieto Nafría et al. (2002a), Blackman et al. (2019b) and Szelegiewicz (1978)
provided accounts of European species. The unusual extent of karyotype
variation in Trama was studied by
Blackman et al. (2000). Trama (Neotrama) afghanica Narzikulov Apterae are matt white; BL c. 3.1 mm. On
roots of Euphorbia sp. in
Afghanistan. Other morphs are unknown. Trama (Neotrama) antennata Mordvilko Colour of apterae in life is unknown,
probably whitish; BL c. 3.4-3.7 mm. On roots of Rumex sp. in Tajikistan. It was redescribed from the original
material by Szelegiewicz (1982). Trama (Neotrama) baroni (Hille Ris
Lambers) Appearance in life is
unknown, BL 4.2-4.9 mm. Described from roots of a Carduus sp. in Malta, in a nest of the ant Camponotus barbaricus, and apterae have also been collected in
Lebanon and Turkey (BMNH collection). Other morphs are unknown. Trama (Neotrama) bigrarta Zhang, Chen, Zhong
& Li, in Zhang Apterae are dark
yellow; BL unrecorded. On Sonchus
oleraceus in Gansu Province, China (Zhang, 1999). Perhaps it is a synonym
of T. caudata (or T. maritima). Trama (Neotrama) caudata (del Guercio) Apterae are whitish, pale yellow or
brownish; BL 2.5-3.3 mm. On roots of
plants in composite genera of tribe Lactuceae (Cichorium, Hypochaeris,
Picris, Lactuca, Leontodon, Sonchus, Taraxacum). Transfers between many of these genera are successful
(Eastop 1953), so some of the other names currently listed in Neotrama (e.g. formicella, narzykulovi) are
probably synonyms. In Europe, and also now
recorded from Turkey (Özdemir et al.
2005), Iran (see Momeni Shahraki et al. 2019) and Central Asia (Kadyrbekov 2017a).
Anholocyclic, no sexual morphs have yet been found. 2n=
9-12 (see Blackman et al. 2000). Trama centaureae Börner Apterae
are grey-green; BL c. 3 mm. On roots of Centaurea
spp. in northern and central Europe (Heinze 1962, Heie 1995). Apart from
its colour in life the characters distinguishing T. centaureae from T. troglodytes (short-based mesothoracic
furca and absence of a frontal suture) show intraspecific variation, and the
separate identity of this species needs further confirmation (Eastop 1953). Trama (Neotrama) euphorbiae Juchnevitch &
Kan Apterae are pale green or white;
BL c.2.5 mm. On roots of Euphorbia lamprocarpa
in Kazakhstan. Trama (Neotrama) kulinitschae Narzikulov Apterae are milky white; BL c.4.7 mm. On
roots of Ferula sp. in Central Asia
(southern Tajikistan). Trama (Neotrama) maritima (Eastop) Apterae are whitish, pale yellow or
brownish; BL 2.6-3.9 mm. In ant-attended colonies on roots of Picris echioides and Sonchus asper in coastal regions of
Europe (including the Azores and Madeira); also in Central Asia, if Protrama pamirica, described from Cichorium in Tajikistan (Narzikulov
1963), and also recorded from Kazakhstan (Kadyrbekov 2018a), is a synonym. It is apparently anholocyclic.
2n=10-14 (see Blackman et al.
2000). Trama (Neotrama) nigrarta Zhang, Chen, Zhong
& Li, in Zhang Apterae are dark
yellow. On Sonchus arvensis in
Gansu, China (Zhang 1999). Perhaps this is a synonym of T. caudata (or T. maritima). Trama orobanches Mamontova
Apterae have BL c. 3mm. On roots of Orobanche sp. in Ukraine (Mamontova 2012). Described as having
apterae similar to T. rara but with
a secondary rhinarium often present on ANT III (as well as those on IV-V),
shorter hairs and smaller eyes. Trama (Neotrama) penecaeca Stroyan Apterae are probably dirty greyish-white
with brown head, antennae, legs, siphuncular cones and sclerites of posterior
segments; BL 3.8-5.0 mm. On roots of Helianthus
tuberosus in India (Jammu, Kashmir). Possibly monoecious holocyclic; an
apterous male was described by Verma (1969). Trama rara Mordvilko Apterae are whitish
to olive-brown (see influentialpoints/Gallery);
BL 2.5-3.5 mm. In ant-attended colonies on roots of Taraxacum spp., and sometimes on certain other Compositae, e.g. Helianthus. In Europe, Iran (Rezwani
2001), Kazakhstan (Kadyrbekov 2014e), Siberia, Japan and North America.
2n=12-14. Trama (Neotrama) taraxaci Shinji Apterae are pale; BL c.1.5 mm. On (roots
of ?) Taraxacum officinale in
Japan. Possibly these are small or immature T. caudata, although that species is only certainly known from
Europe. Trama troglodytes von Heyden
Plate 3g Apterae are white or yellowish white (see influential
points/Gallery), or grey (older adults); BL 2.5-3.9 mm. They occur in
ant-attended colonies on the roots of numerous Compositae (incl. Achillea, Artemisia, Centaurea, Cichorium, Cirsium, Cynara, Helianthus, Lactuca, Lapsana, Sonchus, Taraxacum). In Europe, west Siberia, south-west and Central Asia,
and Japan. Various names have been applied to populations on different host plants,
but there is no evidence from host transfers, karyotype or DNA of any
specific host associations. Long thought to be entirely anholocyclic, but DNA
evidence (Normark, 1999) and the discovery of oviparae and blind, apterous
males in a population in southern England (Blackman et al. 2001) shows that sexual recombination occurs sometimes
(but probably rarely) in this species. 2n=14-23 (see Blackman et al. 2000).
A genus for two species in the Middle East with
emigrant alatae having 5 antennal segments and containing embryos with
remarkably long hind legs. The sexuparae are also distinctive because of
their large size, long rostrum and elongate hind tarsi. Barjadze et al. (2020) reported mitochondrial
and microsatellite DNA differences between the two species. Tramaforda koachi Barjadze,
Foottit & Maw Forming flat galls
on the margins of the leaflets of Pistacia
atlantica, similar to those of T.
wooli but without a serrated outer margin (see Fig.7 in Barjadze et al. 2020). The life cycle is
similar to that of T. wooli, with
migration of emigrant alatae in early October to an unknown secondary host.
Only known from northern Israel and the Golan Heights (Barjadze et al. 2020) This is “Fordini sp. A”
of Koach & Wool (1977) and Wool et
al. (1994). Tramaforda wooli Manheim Forming flat, green galls on Pistacia atlantica by folding the
margin of a leaflet towards the upper surface. The outer edge of the gall is
distinctly serrated (see Fig.7 in Barjadze et al. 2020). The number of aphids in each gall is remarkably
small, at most only 25 (Manheim 2007). Emigrant alatae leave the gall in
early October, flying to an unknown secondary host. Presumably there is a
two-year cycle as in other Fordini, but the secondary host generations are
undescribed, although they would be easily recognised by their long hind legs
with elongate second tarsal segments. Sexuparae were collected on trunks of P. atlantica and P. palaestina in late May to early June (Wool et al. 1994, as “Fordini sp. B”). Only
known from Israel, mostly in the south.
One
Asian species very like Liosomaphis,
but with tubercular spinal processes on the last two abdominal segments,
and the immatures have spinulose hind
tibiae. Tricaudatus polygoni (Narzikulov)
Plate 16b (Fig.42q) Apterae are whitish, pale yellow or
yellowish green; BL 1.6-1.7 mm. Alatae have a quadrate black dorsal abdominal
patch. On undersides of leaves of Polygonum
and Persicaria spp., or on leaves
and twigs of Spiraea. Other
recorded hosts (Berberis, Bidens, Prinsepia) require
confirmation. In Central Asia, India (Himachal Pradesh) and Japan.
Heteroecious holocyclic, with the sexual phase on Spiraea (Chakrabarti & Banerjee 1993a). 2n=8.
Seven east Asian species, at least six of them
associated with Acer. Related to Chaitophorus
and Periphyllus, but apterae have
fused head and pronotum, and long thick, hollow hairs around margin of body, and
their siphunculi lack polygonal reticulation. DNA data is available for only
one species (koyaensis) and
indicates a close relationship with Periphyllus
(Wieczorek et al. 2017b). Alatae
where known are very different from apterae, with head and pronotum
distinctly separated and dark dorsal abdominal sclerotisation. To judge from
the work of Sugimoto (2013) on T.
koyaensis (see below), there can be considerable seasonal intraspecific
variation in the distribution and shape of the dorsal hairs in this genus.
Chakrabarti & Mandal (1986) revised the genus and keyed the species then
known, Sorin (2002) provided a key to species, and Qiao et al. (2004a) reviewed the species in China. Trichaitophorus aceris
Takahashi Apterae are dark green with
the last two segments of the antennae dusky, tarsi dark and siphunculi black
(Takahashi 1937a); BL 1.4-1.8 mm. On young leaves of Acer spp.; originally described from A. rubescens in Taiwan. Aphids identified as aceris have since been recorded from various Acer spp. in north-west India and Nepal (Chakrabarti & Mandal
1986). However, there must be some doubt about the identity of the Indian
populations, as they are pale in life and have pale tarsi and siphunculi
(A.K. Ghosh, 1980). Alatae of Indian populations differ greatly from apterae,
having black-brown head and thorax, antennae and legs mainly dark, abdomen
with a large black patch on abdominal tergites 4-6 and separate sclerites on
other tergites, and Chaitophorus-like
siphunculi with polygonal reticulation (Quednau & Chakrabarti 1976, as Periphyllus pusillus). Chakrabarti et al. (2019) distinguished alatae
from an Acer sp. in Bhutan as a new
species, T. acerifolius Chakrabarti,
Das & Sarkar, but it needs to be confirmed that these are not the unknown
alatae of another species. The populations identified as T. aceris in northwest India are monoecious holocyclic; apterous
males and oviparae were described by Chakrabati & Mandal (1986),
collected on Acer sp. in
October-November. Trichaitophorus aenigmatosus
Pashchenko Apterae are pale green to
green; BL c.1.05 mm. On Acer
tegmentosum in eastern Siberia (Pashchenko 1988a). Other morphs and life
cycle are unknown. Trichaitophorus ginnalarus Qiao,
Zhang & Zhang Colour of apterae
in life is unrecorded; BL 1.3-1.6 mm. On Acer
ginnale in Heilongjiang, China (Qiao et
al. 2004a). Other morphs and life cycle are unknown. Trichaitophorus japonicus
Sorin Apterae are green, with
black-brown tarsi and pale siphunculi; BL c. 1.6 mm. On Acer tschonskii at high elevations in Japan, feeding on upper
sides of leaves along mid-rib and main veins (Sorin 1979b). Sugimoto (2020b)
provided some further description.of the syntype apterae. Other morphs and
life cycle are unknown. Trichaitophorus kominecola Sugimoto
Apterae are yellow in life; BL 0.9-1.8 mm. Monoccious holocyclic on Acer micranthum in Japan, colonising
undersides of leaves. Fundatrices were collected (on twigs) in late April,
and oviparae and apterous males were collected
in late October. Sugimoto (2020b) described and illustrated all morphs, finding
differences between summer and autumn forms of apterae and in the marginal
hairs of first instar larvae produced by different morphs. This species
appears to be specific to A. micranthum,
often in montane regions. Trichaitophorus koyaensis
Takahashi (= Periphyllus montanus Sorin, = Trichaitophorus
takahashii Sorin; synonymy by Sugimoto 2013) Apterae are yellow, yellowish green or
brownish green with pale antennae, legs and siphunculi; BL 0.9-1.6 mm. On Acer rufinerve in Japan (Takahashi
1961b), and also recorded from A.
crataegifolium and A capilllipes (Sugimoto
2013, 2020b). Alatae have black dorsal abdominal cross-bands. Sugimoto (2012)
has provided detailed descriptions and photomicrographs of all morphs
including fundatrices (collected in April), oviparae and apterous males
(October to early December). There are two forms of apterous viviparae; the
form occurring in late summer and autumn was described by Sorin (1979b) under
the name Periphyllus montanus.
There are also different forms of first instars (Sugimoto 2013), some of
which have lamellate marginal hairs similar to those of aestivating nymphs of
some species of Periphyllus (Sorin
2002, as T. takahashii). Trichaitophorus recurvispinosus
Hille Ris Lambers & Basu Apterae
are flattened, pale glassy white with tips of antennae and legs dusky brown;
BL 2.4-2.5 mm. Stated to live on undersides of rolled leaves of a plant that
was unidentified at the time the description was published (Hille Ris Lambers
& Basu 1966), but subsequently identified as Elaeocarpus sikkimensis (BMNH collection, leg. D. Hille Ris
Lambers). In West Bengal, India. Other morphs and life cycle are unknown. [The
host plant and leaf-rolling habit is unlikely for a chaitophorine and needs
additional confirmation. Probably the true hosts are Acer spp., as listed by Chakrabarti & Mandal (1986) under T. aceris. However, the size, colour
in life, length of antennae and shape of marginal hairs make it unlikely that
the synonymy with T. aceris
proposed by these authors is correct.]
About
12 palaearctic, mainly east Asian Myzus-like species with hairs on the siphunculi,
typically host alternating between Lonicera
and Polygonum. In Trichosiphonaphis s.str. (3 spp.) the
siphunculi have a distinct apical flange, whereas in subgenus Xenomyzus (9 spp.) the siphunculi are
flangeless Remaudičre et al. (1992b) provided a review of the genus and a key to
species. Trichosiphonaphis (Xenomyzus) alpestris (Hille Ris
Lambers) Apterae are brown, BL
1.7-2.2 mm. In ant-attended colonies on twigs of Lonicera xylosteum in alpine Europe (Austria, France,
Switzerland). Monoecious holocyclic, with oviparae and apterous males in
September (original description). Trichosiphonaphis (Xenomyzus) cornuta Miyazaki (Fig.42e) Apterae are yellow, BL 1.2-1.3 mm. Alatae
have secondary rhinaria distributed III 70-73, IV 33-41, V 18-30. On Polygonum thunbergii in Japan. Trichosiphonaphis (Xenomyzus) corticis (Aizenberg) (Fig.38k) Apterae are dirty greenish brown; BL
1.80-2.50 mm. In ant-attended colonies on twigs of Lonicera spp. in
north-eastern and central Europe (Russia, Byelorussia, Latvia, Finland,
Poland, Romania, Germany, Czech Republic, Austria, Switzerland), and also in
east Siberia (Pashchenko 1988a). Stekolshchikov & Buga (2006b) provided
descriptions of apterous males and oviparae collected on L. xylosteum in October. Trichosiphonaphis gerberae A.K. Ghosh & Raychaudhuri Apterae are pale yellowish; BL 1.9-2.2
mm. Alatae have secondary rhinaria distributed III 30-32, IV 15-17, V 10-11.
Described from Gerbera macrophylla,
an unlikely host, and later records are from Polygonum or Persicaria spp.
(Raychaudhuri 1980). The life cycle is unknown. Trichosiphonaphis lonicerae (Uye) (Fig.38h,i) Apterae are shiny greenish black or
blackish brown, with paler appendages; BL 2.0-2.2 mm. Alatae have secondary
rhinaria distributed III 32-55, IV 14-27, V 4-12. On twigs of Lonicera spp. in Japan, Korea and possibly also in Hong Kong (Martin & Lau
2011). The life cycle is unclear; it is found on Lonicera from April to November (Remaudičre et al. 1992b), but possibly there is at least a partial migration
to Polygonum. Trichosiphonaphis (Xenomyzus) polygoni (van der
Goot) (Fig.42d,f) Apterae are yellow to dark green,
sometimes almost black; BL 1.3-2.2 mm. Alatae have secondary rhinaria distributed III 20-41, IV 8-21, V 1-11. On Persicaria and Polygonum spp. in east Asia (east Siberia, Japan, China, Taiwan,
India, Pakistan, Java), and introduced to Africa (Burundi, Kenya; Remaudičre at al. 1994), USA (Smith & Denmark
1982), Central America (Panama, Remaudičre at
al. 1992; Costa Rica, Zamora Mejías et al.
2012), and Australia (Brumley 2020). Miyazaki (1971)
found it on the roots of the host. The life cycle is unknown. 2n=12 (Chen
& Zhang 1985a, as T. ishimikawae). Trichosiphonaphis (Xenomyzus) polygonifoliae (Shinji) Plate 20c (Fig.38j, Fig.42g) Apterae are greenish brown; BL 1.8-3.0 mm.
In colonies on twigs of Lonicera spp.
in spring, migrating in June-July to roots of Persicaria and
Polygonum spp. (Hori 1938b, as Aulacorthum lonicerae). Spring
migrant alatae
have secondary rhinaria distributed III 16-46, IV 0-16, V 0. Gynoparae (with many
more secondary rhinaria distributed III 80-100, IV40-63, V 0-9) and males
return to Lonicera in October. The ovipara was illustrated by
Petrović-Obradović et al. (2010). Native to east Asia
(Japan, China, Korea, east Siberia), and introduced to Europe (France –
Remaudičre et al. 1992b, Italy – Coceano &
Petrović-Obradović 2006,, Serbia - Petrović-Obradović et
al. 2010, UK – Martin 2000) and Turkey (Ozdemir et al. 2005).
2n=12. Trichosiphonaphis polygoniformosana (Takahashi)
Plate 20d (Fig.42i) Apterae are dull blackish brown to black,
with dirty yellow siphunculi and cauda; BL 1.8-2.2 mm. On Polygonum and Persicaria spp. in China, Japan, Taiwan, Korea, Thailand and east
Siberia. Possibly with host alternation to Lonicera, but the life cycle is unclear. The original description
included apterae from L. japonica
in Taiwan, and apterae from this host in Japan were also mentioned, but these
specimens are not mentioned in later publications (e.g. Takahashi 1965c), and
may have been T. lonicerae Uye.
Specimens from Takahashi in the BMNH collection (leg. F.V. Theobald) labelled “Lonicera japonica, Japan” (no date) are another, very small,
apparently undescribed species. Trichosiphonaphis (Xenomyzus) tade (Shinji) (Fig.42h) Apterae are dark yellowish brown to muddy
green with dark antennae and siphunculi; BL c. 1.5-1.6 mm. On stalks of Polygonum thunbergii in Japan
(Miyazaki 1971, Moritsu 1983). 2n=12.
A
genus for one species in Taiwan with alate migrants from galls on Distylium having 3- or 4-segmented
antennae, the flagellum being undivided or indistinctly divided and bearing
numerous secondary rhinaria. Tripartita formosana Yeh Galls
on Distylium racemosum in Taiwan
are green, spindle-like, 9-19 mm long and 4-9 mm wide, arising from the upper
sides of leaves, with openings on the undersides, and with several galls on a
single leaf (Yeh & Ko 2017). Galls of this species were also found on Distyliopsis dunnii. Developing galls
contained fundatrices from October to May, producing alatae (BL 1.5-2.3 mm)
from February to June, which presumably migrated to an unknown secondary
host. First instars believed to be sterile soldiers were also observed,
walking on the surfaces of galls and leaves.
Two
Asian species related to Macrosiphoniella
but with a characteristic
caudal structure, and an apparent association with both Campanulaceae and Artemisia. Contrary to Lee et al. (2002c), this structure is a
modification of the actual cauda, rather than a process on abdominal tergite
8; the protruding structure beneath it is an extension of the anal plate. It is possible that this peculiar
cauda has some function in relation to ant attendance. Tshernovaia adenophorae Holman & Szelegiewicz Plate 28c (Fig.7a) Apterae are dull black with a paler
transverse bar anterior to base of cauda; BL 2.0-2.8 mm. In ant-attended
colonies on stems and undersides of leaves of Adenophora spp., and also found on Artemisia annua (Lee et al.
2002c). In Russia (Transbaikalia),
Mongolia and Korea.. Tshernovaia spirocaudicula Ivanoskaya
Apterae are yellowish, slightly waxy, with black antennae, legs,
siphunculi and cauda; BL c.2.2 mm. On Artemisia
sp., and also collected from stems of “bluebells” (Campanulaceae). In
Siberia.
One or two highly specialized conifer-feeding
species. Its relationship to other Hormaphidinae is uncertain, but a
cladistic analysis (Chen et al.
2014) indicates a sub-basal position in a clade with Hormaphidini. Only
apterous viviparae are known. Tsugaphis sorini
Takahashi Apterae are pale greenish,
including antennae and legs; BL c. 0.9 mm. On undersides of leaves of Tsuga sieboldii in Japan (Takahashi
1957). Other morphs, biology and life cycle are unknown. Tsugaphis sorini ssp. piceicola
Shaposhnikov & Gabrid (fig.
65B) Apterae are
pale green, translucent, older specimens becoming dark brownish-green; BL c.
0.9 mm. On Picea spp., especially pungens, schrenkiana, living in yellow depressions on upper and undersides
of needles, which eventually become twisted and brittle, and drop
prematurely. In Kyrgyzstan.
Anholocyclic, overwintering as hibernating apterae (Shaposhnikov & Gabrid
1987). Differences from sorini s.
str. are very small, and the two could be synonyms.
Two
palaearctic species associated with Ranunculaceae, resembling Myzus but with a distinctive constriction
at the base of the cauda. Alatae are without a dark dorsal abdominal patch,
and have numerous secondary rhinaria on ANT III, and also some in a row on IV
or IV-V. Tubaphis clematophila (Takahashi) (Fig. 23a,b; Fig.45e) Apterae are green, with apices of antennal
segments brownish; BL c.1.8-2.0 mm.
Alatae have secondary rhinaria distributed III 32-36, IV 11-14, V 2-9. On Clematis spp. and Ranunculus sp. in Japan, Korea, China (Su et al. 2011) and east Siberia, and it has also been collected on Clematis connata in Pakistan
(Naumann-Etienne & Remaudičre 1995). 2n=12. Tubaphis ranunculina (Walker) Plate 17f (Fig.45f,g) Apterae are yellowish with pale
appendages; BL 1.2-1.9 mm. Alatae have secondary rhinaria distributed III
23-38, IV 0-10, V 0-6. On undersides of leaves of Ranunculus spp. throughout Europe, and also recorded from western Siberia (Altai), India (West Bengal),
east Siberia (Kamchatka) and Japan (for detailed distribution see
Stekolshchikov & Buga 2014). Monoecious holocyclic, sexual morphs in
October (Blackman 2010); detailed descriptions of the ovipara and alate male are provided by
Stekolshchikov & Buga (2014).
About 14 species
in east and south-east Asia, typically with host alternation between often
elaborately-branched galls on Styrax,
and secondary hosts in Loranthaceae or Santalaceae. It is one of three genera
of Cerataphidini in which the typical intracellular aphid symbionts (Buchnera) have been replaced by
extracellular yeast-like symbionts, the others being Cerataphis (except C.
bambusifoliae) and Glyphinaphis (Fukatsu et al. 1994, Xu et al. 2018). Determinations of the
life cycles and/or symbiont types of several species (e.g. Aoki & Kurosu
1993, Kurosu et al. 1994, Aoki et al. 1995, Stern et al. 1997) have resulted in
transfers to this genus of species described from Styrax in the genus Astegopteryx,
and from Loranthaceae, originally placed in Rappardiella. A cladistic study (Chen et al. 2014) placed Tuberaphis
close to Cerataphis, and this is
confirmed by molecular work (e.g. Xu et
al. 2018). Noordam (1991) redescribed and keyed the species in
Java (as Rappardiella), and Jiang et al.
(2012a) and Qiao et al. (2018)
reviewed and keyed the Chinese species.
Fukatsu
et al. (1994) compared symbionts
and gall structure with other Cerataphidini. Aoki & Kurosu (2010)
reviewed life cycles, gall formation and soldier behaviour. Tuberaphis
breviseta A.K. Ghosh
(= Neotuberaphis indica of
Blackman & Eastop 1994) Apterae
are dark brown, BL 1.3-1.5 mm, living in white wax in marginal leaf-folds on Eugenia sp. in Nagaland, India. Alatae
have a pale brown abdomen and dark wing veins (Raha & Raychaudhuri 1981,
as Neotuberaphis indica). A.K.
Ghosh (1988) provided a redescription of type specimens. The life cycle is
unknown. Tuberaphis cerina (Noordam)
Apterae are yellowish white tinged with reddish brown, with a dull
transparent coat of wax; BL 1.1-1.3 mm. On Dendrophthoë pentandra in Java, living pressed close to surfaces
of rolled leaves, attended by ants. Tuberaphis
coreana Takahashi
(= taiwana of Blackman &
Eastop 1994) (Fig.60b) Galls are large (about 14cm in
diameter), pale greenish-yellow and coral-like, with small circular distal
openings, and arise from the stem of Styrax
formosanum in Taiwan (fig. 124G).
Alate emigrants (BL c.1.75 mm) were collected in August (Takahashi 1934b).
The secondary hosts, assuming the synonymy with T. taiwana is correct, are Viscum
spp., on which apterae of the presumed secondary host generations have
been collected in Japan and Korea. The colour of these apterae in life is
unrecorded, but presumably they have a wax fringe; BL 1.7-2.0 mm. The
synonymy is fairly well-supported by molecular data (Stern et al. 1997) but secondary host
populations are not known from Taiwan, and some further confirmation seems
necessary (see Aoki & Kurosu 1993, as taiwana). [Tao (1969, as Astegopteryx)
synonymised this species with A.
vandermeermohri (Hille Ris Lambers), but this is untenable because the
gall described for that species is quite different, and there are also
differences in the soldier morph and first instar exule (Aoki & Kurosu
1993).] Tuberaphis
cymigalla (Qiao & Zhang) Galls are green, branched, arising from
twigs, with many openings, on Distylium
racemosum in Fujian, China (Qiao & Zhang 1998a, as Rappardiella). Only a single aptera
from the gall is described, and the host association is an unlikely one
requiring further confirmation of the generic placement of the aphid, and its
host. Tuberaphis dendrotrophe Qiao, Jiang & Chen (in Jiang et
al. 2012a) Apterae are broadly
oval, brown, with margin of body covered with white wax; BL 1.5-1.8 mm.
Alatae are undescribed. On Dendrotrophe
frutescens in Hong Kong (Tuberaphis
sp. in Martin & Lau 2011). Jiang et
al. (2012a) provided descriptions of adult apterae and all larval instars
(but most of the measurements in Table 1 of their paper are about 33% too
high). Tuberaphis
leeuweni (Takahashi) Galls are formed from flower buds of Styrax subpaniculatus in Sumatra. They are slender, green, pod-like, downy,
5-8 cm long, with a distal opening (fig. 124D). Alatae, BL about 1.6 mm, were collected
from them in June by Takahashi (1936,
as Astegopteryx), and in September
by Aoki et al. (1995) The September-collected
alatae were found to be sexuparae, indicating an absence of host alternation.
Soldier-like second instars occur in the galls (Aoki et al. 1995). Tuberaphis loranthi (van der Goot)
(Fig.39a-d) Apterae are orange-brown, with a narrow wax
fringe, and a segmentally-divided dusting of mealy wax; BL 1.5-2.1 mm. On old
leaves and in dense colonies on woody stems of mistletoes (Scurrula spp.; records from Loranthus should probably also be
referred to this host genus). In Taiwan, India (A.K. Ghosh 1988), Java
(Noordam 1991) and Papua-New Guinea (BMNH collection, leg. E.J. Brough). The
life cycle is unknown. Tuberaphis macrosoleni Noordam
Apterae are brown, with antennae and legs paler, and with a dorsal
layer of flaky wax, becoming thicker and woolly on margins; BL 1.3-1.6 mm. In
rolled-leaf galls on Macrosolen
cochinchinensis in Java (Noordam 1991). Tuberaphis owadai Kurosu & Aoki Galls on Styrax tonkinensis in northern Vietnam are silvery grey,
coral-like, 9-24 cm across, similar to those of T. coreana and T. sumatrana
on other Styrax spp. (Kurosu
& Aoki 2003). Colony sizes in the galls were estimated at 50,000 to
180,000 individuals. Alatae, BL 1.5-1.9 mm, produced in the galls in
September, were found to be sexuparae, so the species lives on Styrax without host alternation.
Numerous second instar sterile soldiers were present in the galls. Jiang et al. (2012a) provided a
redescription including immature stages from galls on Styrax japonicus in
Yunnan, China. Tuberaphis scurrulae Noordam (Fig.39e) Apterae are velvety black or dull brownish
black, with a flat narrow marginal wax fringe; BL 1.7-2.0 mm. On twigs of Scurrula sp. in Java (Noordam 1991). Tuberaphis styraci (Matsumura) (= Hamiltonaphis styraci of Blackman & Eastop 1994) Large, coral-like galls are formed on the
twigs of Styrax obassia in Japan.
The galls take two years to develop (Aoki & Kurosu 1990, as Astegopteryx). Colonies in the galls
may reach a size of 15,000 to 20,000 individuals, of which more than half may be sterile second
instar soldiers. The soldiers not only defend the colony but also clean the
gall, by pushing out honeydew droplets and exuviae (Aoki & Kurosu 1989,
as Astegopteryx). Shibao et al. (2004, 2016) studied the
controlling mechanism for soldier production, and Kutsukake et al. (2004) identified a
soldier-specific venomous protease presumably used in colony defense. There
is no host alternation. Alate sexuparae (BL c. 1.4 mm) appear in late July or
August of the second year and deposit yellowish sexuales on the undersides of
the Styrax leaves. Oviparae lay
overwintering eggs probably in fissures in the bark (Aoki & Kurosu 2010). Tuberaphis sumatrana (Hille Ris Lambers) Galls are coral-like, silvery grey, about
10 cm in diameter when fully developed, with small openings on distal lobes,
arising from the stem via a short thick branching tube (fig. 124F). They are found on Styrax spp. (serrulatus
var. mollissimus, subdenticulatum) in Sumatra. Alate emigrants, BL about 1.5-1.6 mm, were
collected in February (Hille Ris Lambers 1931a, as Astegopteryx). Tuberaphis takenouchii
(Takahashi) The 'broccoli-head-like'
gall of this species on Styrax japonicus (fig. 124I)
was redescribed by Aoki & Usuba (1989; as Aleurodaphis), who found that it contained sterile second instar
soldiers. Migration occurs to Loranthaceae, including Scurrula sp. This host
alternation and synonymy was proposed by Kurosu et al. (1994) and supported by the molecular studies of Stern et al. (1997). For a description of
the alate emigrant see Takahashi (1934a, as Astegopteryx). Apterae on Scurrula are dark
grape-coloured, usually with little wax, BL 1.2-1.4 mm, and inhabit rolled
leaves (Noordam 1991, as Rappardiella
plicator). In Taiwan, Sulawesi and
Java. To
judge from a paratype slide in the BMNH collection, T. loranthicola A.K.
Ghosh 1988, originally described from Taxillus
sutchuenensis in China (Tseng & Tao 1938, as Astegopteryx loranthi) is probably a synonym. Tuberaphis viscisucta Zhang
Colour of apterae in life is unrecorded, not wax-powdered; BL c. 1.3
mm. On Viscum album in Yunnan,
China, causing downward curling of edges of leaves. This is likely to be a synonym of T. takenouchii. Tuberaphis xinglongensis (Zhang) Apterae were described from an unidentified
plant in Hainan, China (G. Zhang & Zhong 1982b, as Astegopteryx). Alatae have numerous secondary rhinaria,
distributed ANT III 33-39, IV 13-17, V BASE 7-10. This species was
subsequently recorded from an unidentified species of palm, and then from Cocos nucifera. Its generic placement
in Tuberaphis by Jiang et al. (2012a) was confirmed by Chen et al. (2014), although the host
association is an anomalous one for a member of this genus.
About 60 species of oak- and chestnut-feeding
aphids with one or more tubercular abdominal spinal processes. All viviparae
and males are alate. The genus includes some well-defined subgenera of
limited distribution (Tuberculoides
in western palaearctic, Orientuberculoides,
Nippocallis and Acanthocallis
in east Asia, Pacificallis in
western nearctic, and Toltecallis
in Mexico). Richards (1968c) and Hille Ris Lambers (1974) reviewed the world
fauna, and Quednau (1999) revised the subgeneric classification and provided
a key to species and illustrations of all available morphs. Quednau (2003,
pp. 294-6) keyed the currently known oviparae of four subgenera. Regional
accounts are available for Britain (Stroyan 1977), north-west Europe (Heie
1982), Iberian peninsula (Nieto Nafría & Mier Durante 1998), India
(Chakrabarti 1988; A.K. Ghosh & Quednau 1990), China (Qiao et al. 2005b), eastern Siberia
(Pashchenko 1988a) and Japan (Higuchi 1970). W. Zhang & G. Zhang (1991)
did a cladistic analysis of 23 species in China. Yao (2011) constructed
phylogenetic trees based on mtDNA sequences for 20 species of Tuberculatus in Japan, including some
undescribed species, and showed that wing-loading was greater in ant-attended
species. Y. Lee et al. (2017) found variation in DNA barcoding sequences
indicating the existence of cryptic species in several subgenera. Quednau
(1992) keyed the species of subgenus Pacificallis,
and Qiao & Zhang (2002a) reviewed Chinese species of the subgenus Nippocallis. Tuberculatus
(Acanthocallis) acuminatus Zhang, Zhang & Zhong Alatae are whitish with dark head and
siphunculi; BL c. 1.3 mm. On Quercus
acutissima in China (G. Zhang et al. 1990b). Sexual morphs unknown. Tuberculatus (Tuberculoides)
albosiphonatus Hille Ris Lambers Alatae are pale green, with black-banded
antennae, a sharply-bordered dark brown spot on the pterostigma, and pale
siphunculi; BL 1.1-1.7 mm (Hille Ris Lambers 1974a). On Quercus infectoria (incl. var. boissieri) in Iraq (Kurdistan) and Israel (BMNH collection, leg.
V.F. Eastop). Sexual morphs are unknown. Tuberculatus (Tuberculoides) annulatus
(Hartig) Alatae are yellowish, greyish-green or pink to purple in summer,
with black-banded antennae, black tarsi, and dark distal two thirds or more
of siphunculi (see
influentialpoints.com/Gallery); BL 1.7-2.2 mm. On undersides
of leaves of Quercus spp.,
especially Q. robur, less commonly Q. petraea. Throughout Europe, Iran
(Rezwani et al. 1994), Siberia
(Pashchenko, 1988b), Kazakhstan (Kadyrbekov 2017a), north-west China (Qiao et al. 2005b) and introduced on
European oaks to Australia, New Zealand, North and South America. Alate males
and apterous oviparae occur in October (northern hemisphere). Rohitha &
Penman (1984) studied development under laboratory conditions, Kennedy
(1986b) studied the role of tarsi in host selection, Heimbach (1986) studied
population dynamics in relation to honeydew production, and Dahlsten et al. (1990) reported specific
parasitoids in Germany. [DNA barcode data now indicate that two species may
be confused under the name T. annulatus,
differing in colour, COI sequence and morphological features, and both
present in both Europe and North America (Skvarla et al. 2017; Y. Lee et al.
2017).] 2n = 14. Tuberculatus (Tuberculoides) borealis
Krzywiec Alatae are pale blue-green,
yellow mottled with green, or yellow (see
influentialpoints.com/Gallery), with blackish-banded antennae
and siphunculi only apically dark (rarely over more than distal half); BL
1.9-2.3 mm. On Quercus robur, more
rarely on Q. petraea or robur × petraea hybrids, in Europe (UK,
Belgium, Denmark, Norway, Sweden, Italy, Poland, western Russia, Turkey),
east to Iran (BMNH collection, leg. S.H. Hodjat), and introduced to North
America (British Columbia; Quednau 1999). Sexual morphs occur in
October-November (Krzywiec 1971, Quednau 1999). Tuberculatus (Pacificallis)
californicus (Baker) Alatae are pale
yellowish-green (Baker 1917a) with banded antennae and brown-black spots at
bases of all tibiae; BL 1.7-2.5 mm. On Quercus
lobata in western USA (California, Colorado, Oregon, Utah). Sexual morphs
occur in November (BMNH collection, leg. R. van den Bosch). Tuberculatus
(Orientuberculoides) capitatus (Essig and
Kuwana) Alatae are pale green,
greenish-yellow or whitish-yellow, antennae ringed with black, legs pale; BL
2.4-2.6 mm. Under leaves of Quercus
spp. in east Asia (China, Taiwan, Korea, Japan). Ant-attended (Yao 2011).
Oviparae and males were illustrated by Moritsu (1983). A single alata trapped
in Korea was described as a subspecies, T. capitatus intermedius (Hille Ris Lambers 1974a); Quednau
(1979) redescribed this form from specimens collected in Korea on Castanea crenata, and (Quednau 1999)
also recorded it from Q. variabilis.
2n = 14. [Note that the illustrations of T. capitatus by Quednau (1999, p. 231,
figs 872-887) should be substituted by those in Quednau (2003, p. 301, figs
2451-2464).] Tuberculatus (Nippocallis)
castanocallis (Zhang & Zhong) Alatae squat-bodied with thickly-bordered
wing veins, femora dark distally, conspicuous large dark marginal abdominal
tubercles and dark siphunculi; BL c. 2.2 mm.
On upper sides of the leaves of Castanea
bungeana and C. seguinii in
China (Liaoning, Hebei, Shandong, Zheijiang, Hunan, Guangxi, Yunnan; Qiao
& Zhang 2002a, Qiao et al.
2005b). Tuberculatus (Nippocallis) cereus (Zhang
& Zhong) Alatae are squat-bodied
with very thickly-bordered forewing veins, the pigment extending between Cu1b
and the branches of the media, and dark marginal abdominal tubercles and
siphunculi; BL c. 1.9 mm. On Castanea
bungeana in Shandong, China (Qiao & Zhang 2002a). Tuberculatus
(Nippocallis) ceroerythros Qiao & Zhang Alatae are pale red in life, coated with
white powder, with thickly-bordered wing veins; BL c. 2.3 mm. On Castanea bungeana and C. seguinii in China (Guangxi, Jiangxi;
Qiao & Zhang 2002a, Qiao et al.
2005b). Sexual morphs and life cycle are unknown. Tuberculatus (Pacificallis)
chrysolepidis Quednau Colour of alatae in life is unknown; BL
1.9-2.9 mm. On Quercus chrysolepis
in California, USA (Quednau 1992). Sexual morphs and life cycle are unknown. Tuberculatus (Pacificallis) columbiae
Richards Colour in life was not
observed; BL 2.0-2.5 mm. On Quercus
garryana, recorded from Oregon (BMNH collection, leg. D. Hille Ris
Lambers) and British Columbia (Richards 1965). Sexual morphs have not been
described. Tuberculatus cornutus
Richards Colour in life is not
recorded; BL c. 1.9mm. On an unidentified Quercus
sp. in Turkey. Sexual morphs have not been described, but the ovipara,
observed in October (Richards 1969a), is illustrated by Quednau (1999). Tuberculatus (Tuberculoides) eggleri
Börner Alatae are pale green or
yellow-green with orange thorax, banded antennae and siphunculi pale, or only
dark apically; BL 1.7-2.6 mm. On upper or undersides of leaves of Quercus spp. (especially pubescens; also recorded from aegilops, canariensis, cerris, etruscus,
frainetto, ilex, mirbeckii, petraea). In southern, central and eastern
Europe, North Africa (Algeria), and eastward to Turkey and Moldavia.
Monoecious holocyclic on Q. pubescens,
but it may overwinter as viviparae in mild winters. Lampel (1974) discussed
seasonal variation and differences from T.
borealis. [T. africanus Hille
Ris Lambers, described from Quercus
mirbeckii (canariensis) in
Algeria (Hille Ris Lambers 1974a), is the early summer form of T. eggleri (Quednau 1999).] Tuberculatus (Tuberculoides) etruscus
Barbagallo & Binazzi Alatae are
yellowish-green, with head and thorax slightly brownish, and siphunculi
darker towards apices; BL 1.5-2.9 mm. On Quercus
frainetto in Italy (Barbagallo & Binazzi 1991). Sexual morphs and
life cycle are unknown. Tuberculatus (Orientuberculoides)
fangi (Tseng & Tao) Alatae are pale green with banded
antennae; BL 1.8-2.9 mm. On undersides of leaves of Quercus spp. (acutissima,
variabilis) in China, Korea and Japan (Hille Ris Lambers 1974a, Quednau
1999). Sexual morphs have not been recorded. Tuberculatus (Orientuberculoides)
fuscotuberculatus Zhang, Zhang & Zhong Alatae are yellowish green to green with
banded antennae, dark longitudinal stripes on tibiae, tubercles on abdominal
tergites 1-2 or 1-3 often dark and arising from dark sclerites, and
siphunculi dark at least distally; BL 2.6-3.2 mm. On Quercus acutissima in China (G. Zhang et al. 1990b), and also recorded from Japan, and Korea, where two
vagrant oviparae were collected in November (Quednau & Lee 2001;
described and illustrated by Quednau 2003, p. 291). Tuberculatus
(Toltecallis) garciamartelli Remaudičre & Quednau Alatae are whitish, with banded antennae
and a pair of irregular pale green submarginal stripes on the abdomen; BL
1.4-1.9 mm. On Quercus microphylla
in Mexico (Remaudičre & Quednau 1983). Sexual morphs are unknown. Tuberculatus (Acanthocallis)
grisipunctatus Zhang, Zhang & Zhong Alatae are green in life; BL 1.7-2.3 mm.
Recorded from various Quercus spp.
in China (Zhang et al. 1990b,
Quednau 1999), although according to Qiao et
al. (2005b) it occurs only on Q.
dentata. Sexual morphs are unknown. Tuberculatus (Orientuberculoides)
higuchii Hille Ris Lambers Alatae are pale yellow (Quednau &
Shaposhnikov 1988); BL 1.8-2.7 mm. On Quercus
spp. (dentata, mongolica, serrata) in
Japan, Korea, China and eastern Siberia. In Japan, populations with thicker
and blunter abdominal spinal processes and shorter PT were described by Hille
Ris Lambers (1974) as a subspecies, T. higuchii breviunguis, which
might be seasonal variants. It is not clear whether “higuchii-a” and “higuchii-b”
of Yao (2011) correspond to these two named taxa. Hoever, DNA barcoding
studies have now indicated that the name T.
higuchii may be being applied to as many as four cryptic species, with
minor morphological differences betwen them (Y. Lee et al. 2017). Sexual morphs are unknown. Tuberculatus (Acanthotuberculatus)
indicus L.K. Ghosh
Alatae are reddish to greenish-yellow with pale eyes, banded antennae,
mainly dark hind femora and bases of hind tibiae and long black spinal
processes on abdominal tergites 2 and 3, contrasting with long pale processes
on thorax and abdominal tergite 1 (see Moritsu 1983; as T. fulviabdominalis); BL 2.0-3.2 mm. On Quercus spp. in India, Korea, China and Japan, attended by ants
(Yao 2011). The life cycle is variable; alate viviparae were collected in
December-January in Manipur, India (Chakrabarti 1988), but oviparae and males were collected in South
Korea in October-November (Quednau & Lee 2001). The ovipara is
illustrated by Quednau (1999). This is the species referred to in Japanese
and Korean literature (e.g. Higuchi 1972, Paik 1972) as T. fulviabdominalis (Shinji) - see Hille Ris Lambers (1974).
However, DNA barcoding studies have now indicated that the name T. indicus may be being applied to
more than one species (Y. Lee et al.
2017). Tuberculatus (Tuberculoides) inferus
Barbagallo Alatae are yellowish
green, lightly dusted with wax, with banded antennae and dark-tipped spinal
processes on abdominal tergite 3; BL 1.3-2.2 mm. Described from Quercus pubescens (sensu lato) in Sicily, inhabiting only
the undersides of the most basal leaves on young plants (Barbagallo 1990).
Also known from Quercus sp. in
Turkey (BMNH collection, leg. D. Hille Ris Lambers). Life cycle and sexual
morphs are unknown. Tuberculatus (Acanthotuberculatus)
japonicus Higuchi
Alatae are yellow to green or brown, with pale eyes, mesothorax
yellowish-green or brown dorsally and dark brown ventrally, and dark
green-black spinal processes on ABD TERG 2 and 3 (Quednau & Shaposhnikov
1988); BL 2.5-2.9 mm. On Quercus
spp. (dentata, mongolica), attended by ants, in Korea, Japan and eastern
Siberia, and also recorded from China (Qiao et al. 2005b). Sexual morphs were collected on Q. dentata (possibly the preferred
host) in Korea in early November (Quednau & Lee 2001). Tuberculatus (Orientuberculoides)
kashiwae (Matsumura)
Alatae are pale yellow to pale greenish-yellow, sometimes
yellowish-white or nearly white (Quednau & Shaposhnikov 1988); BL 1.4-1.9
mm. On undersides of leaves of Quercus
spp. (dentata, mongolica) in Japan, China, Korea and eastern
Siberia. DNA barcoding studies (Y. Lee et
al. 2017) indicate that there may be two cryptic species under this name,
the distribution and host specificity of which needs to be clarified.
Oviparae and a male were collected in Japan in the second half of September
(Quednau 2003), and Paik (1972) records oviparae in Korea in early November.
2n = 14. Tuberculatus (Pacificallis) kiowanicus
Hottes Alatae are pale yellow,
yellowish green or green, with head and thorax light brown, orange or
pinkish, often dusky on sides; BL 1.6-2.6 mm. On undersides of leaves of Quercus gambelii and Q. gunnisonii in Colorado and Utah,
USA. Sexual morphs were collected on Q.
gambelii in October (BMNH collection, leg. G.F. Knowlton) and the male
was described by Hottes (1949b, as Myzocallis
tonkawa). Palmer's (1952) account under the name M. maureri applies to this species. Tuberculatus (Orientuberculoides)
konaracola (Shinji)
Alatae are pale green or pale yellow-green to almost white, with
banded antennae, brown marginal stripes on pronotum, dark Cu1a and
Cu1b in forewing, dark tibiae, brown-black spinal processes on
abdominal tergite 3 and dark distal halves of siphunculi; BL 1.7-2.8 mm. On Quercus spp. (serrata, dentata) in Japan (Shinji 1941); redescribed by Hille
Ris Lambers (1974) from alatae trapped in Korea. Holocyclic in Japan; Shinji
(1941) described the sexual morphs, and Quednau (1999) illustrated an ovipara
from Q. dentata in Japan in
November. Tuberculatus (Orientuberculoides)
kunugi (Shinji)
According to the original description, alatae are green with black
eyes, banded antennae and with long spinal processes on thorax and abdominal
tergites 1-3 that are all concolorous with body (Shinji, 1924); BL not
recorded. On Quercus acutissima and
Q. serrata in Japan (Higuchi &
Miyazaki 1969); no specimens are known, and it could not be included in the
key. Tuberculatus
(Nippocallis) kuricola (Matsumura) Alatae are rather squat-bodied, pale green
(Essig & Kuwana 1918) or red (R.L. Blackman, unpublished observations),
covered with white wax; BL 1.2-1.9 mm.
Forewings are patterned with very broad bands of fuscous that follow
the veins. Brachypterae occur commonly
(Takahashi 1924). On undersides of
leaves, along the mid-ribs, of Castanea
spp., especially C. crenata,
attended by ants (Yao 2011). Records
from Quercus require
confirmation. In Japan, China, Taiwan
and Korea, and introduced to Brazil (Bergmann et al. 2002), Spain (Manilla et
al. 2001) and Madeira (Ilharco 1984). DNA analysis (Y. Lee et al. 2017) has indicated that the
name might be being applied to two species with minor morphological
differences; it is not known whether one of these could be M. kuricola var. cantonensis of Takahashi (1936a). Monoecious holocyclic in Japan;
sexuales occur in October-November (Takahashi 1924, Higuchi 1972). Sakata (1999) studied interactions between T. kuricola, Lachnus tropicalis and the ant Lasius niger. 2n=14 (Shibata’s record of 2n=12 in both male and
female cells must be erroneous.) Tuberculatus
(Toltecallis) leptosiphon Quednau Colour of alata in life unknown; BL c.1.7
mm. On Quercus microphylla in
Mexico (Quednau 1999). Sexual morphs are unknown. Tuberculatus
(Acanthocallis) macrotuberculatus (Essig &
Kuwana) Alatae are dark green or
mid-brown with blackish cross-bands on abdominal tergites 1 and 2 bearing
dark (sometimes pale-tipped) processes and forewing veins dark-bordered; BL
2.0-3.2 mm. Immatures are pale green, sometimes darker posteriorly, often
with a whitish spinal band. On upper
and undersides of leaves of Quercus
dentata, attended by ants. In
Japan, and probably more widely distributed, as previously synonymised
and confused in the literature with T. quercicola (e.g. Quednau 1999).
Watanabe et al. (2015) described
two forms of this species from central and eastern Hokkaido. 2n=16 (Blackman
1986, as T. quercicola). Tuberculatus maculipennis
Hille Ris Lambers Alatae are greyish
due to wax dusting, with pale antennae and legs, dark head and thorax,
blackish spinal and marginal processes, and maculate forewings; BL 1.0-2.0mm.
On Quercus spp. (infectoria, ?calliprinos, ?robur) in
south-west Asia (Turkey, Lebanon, Iraq) and also recorded from Q. pubescens in Italy (Patti &
Barbagallo 1997). Sexual morphs were collected in Turkey in October-November
(Hille Ris Lambers, 1974). Tuberculatus (Nippocallis)
margituberculatus (Zhang & Zhong) Alatae are squat-bodied with dark head and
thorax, wing-veins very thickly bordered with pigment extending between Cu1a
and branches of media (but leaving basal parts of these veins unbordered),
very conspicuous large black marginal tubercles on abdominal tergite 4, and
dark siphunculi; BL c. 1.9 mm. On Castanea
bungeana in China (see Qiao & Zhang 2002a), and in Hong Kong it was
found on C. mollissima (Martin
& Lau 2011). Records from Quercus
spp. (Qiao et al. 2005b) require
further confirmation. Quednau (1999) illustrated an ovipara collected on Castanea sp. in October. Tuberculatus (Pacificallis) maureri
(Swain) Alatae are light green to
apple green with red eyes, dusky-dark tibiae and forewing with a
conspicuously dark-bordered Cu1b vein; BL 1.4-2.2 mm. On Quercus agrifolia and Q. kellogii in California, USA. Sexual
morphs were found in early November (BMNH collection, leg. D. Hille Ris
Lambers). Tuberculatus (Tuberculoides) maximus
Hille Ris Lambers Colour in life is
unknown, probably pale, with banded antennae, forewings with dark triangular
spots at distal ends of Cu1a and branches of media, and a rather
dark cauda; BL 2.0-3.1 mm. On Quercus spp.
(macranthera, persica) in southwest Asia (Iran, Turkey, Armenia), and there is
also a record from Greece (Tsitsipis et
al. 2007). Hille Ris Lambers (1974) discussed seasonal and geographic
variation; sexual morphs tentatively ascribed to this species were collected
on Q. macranthera in Iran in
November (an ovipara is illustrated by Quednau 1999). Tuberculatus (Toltecallis) mexicanus
Remaudičre & Quednau Alatae are
very pale green to whitish with a pair of dark lateral stripes on prothorax
and a pair of darker green lateral spots on each of abdominal tergites 1, 3
and 6; BL 1.3-2.4 mm. On Quercus
spp. (peduncularis, rugosa) in Mexico (Remaudičre &
Quednau 1983), and subsequently recorded from Panama (Quirós et al. 2009). Apparently anholocyclic. Tuberculatus (Tuberculoides) moerickei
Hille Ris Lambers Plate
9b Alatae are pale yellow with
banded antennae and distally dark siphunculi; BL 1.4-2.6mm. On Quercus spp. (especially infectoria) in Mediterranean (Cyprus)
and southwest Asia (Iraq, Israel, Lebanon, Turkey). A sample from Q. ?pedunculata (= robur) in Turkey was described as a subspecies, T. moerickei galatensis (Hille Ris
Lambers, 1974), but was considered to be merely a variant population by
Quednau (1999). 2n = 14. Tuberculatus (Tuberculoides) naganoe
(Shinji) According to original
description, alatae are light green with red eyes, banded antennae and black
tibiae, and with pale finger-like spinal processes on head, thorax and each
abdominal tergite; BL c. 1.6mm. On Quercus
spp. in Japan (Shinji 1941). No specimens are known and it could not be
included in the key on the basis of the available information. It was
considered a nomen dubium by
Remaudičre & Remaudičre (1997). Tuberculatus (Tuberculoides) neglectus
Krzywiec Alatae are pale yellow, with
darker yellow mesothorax, eyes usually whitish yellow, antennae banded,
siphunculi dark except at base (see influentialpoints.com/Gallery);
BL 1.4-2.2 mm. On undersides of leaves of Quercus
petraea, more rarely on Q. robur
or hybrids between the two. In northern and north-west Europe (Britain,
Ireland, Sweden, Denmark, Germany, northern Poland). Sexual morphs occur in
November (Krzywiec 1965; Heie 1982). Tuberculatus (Orientuberculoides)
nervatus Chakrabarti & Raychaudhuri Colour of alata in life unrecorded;
antennae banded, thorax probably dark with postscutum black, forewing veins
slightly bordered with fuscous, the long, finger-like spinal processes on
abdominal tergite 1-3 are dusky to dark, and siphunculi are dark towards
apices. BL c. 3.6mm. On Quercus griffithii (see Quednau 1999) in India
(Meghalaya, Manipur, Nagaland; Chakrabarti & Raychaudhuri 1976,
Chakrabarti 1988). Life cycle and sexual morphs are unknown. Tuberculatus (Acanthocallis)
nigrosiphonaceus (Zhang & Zhang) Colour of alatae in life unknown, but with
dark head and thorax, dusky wings with dark veins, dark transverse bars on
abdomen and dark siphunculi; BL c. 2.3 mm. On Quercus liaotungensis in China (Beijing; W. Zhang & G. Zhang
1994, as Myzocallis), and oviparae
were collected in Japan in late September on Quercus mongolica var. grosseserrata
(described and illustrated by Quednau 2003, p. 292-3). Quednau (1999)
transferred this species to Tuberculatus
(Acanthocallis). Tuberculatus (Orientuberculoides)
paiki Hille Ris Lambers Alatae are pale greenish-yellow (Quednau
& Shaposhnikov 1988), in midsummer the tibiae and finger-like processes
on abdominal tergite 3 may be dark (Quednau 1999); BL 2.4-3.2 mm. On
undersides of leaves of Quercus spp.
(dentata, mongolica) in India (Meghalaya), Bhutan (Chakrabarti & Das
2014), Korea, China, eastern
Siberia and Japan (Yao & Katagiri 2011).
A.K. Ghosh & Quednau (1990) provided a redescription. Sexual morphs and
life cycle are unknown. Tuberculatus (Tuberculoides)
pallescens Hille Ris Lambers Alatae are bright green, with banded
antennae including short blackish-brown sections near middle of ANT III; BL
2.5-2.5 mm (Hille Ris Lambers 1974a). On Quercus
spp. (calliprinos, infectoria, ?robur) in south-west Asia (Iraq, Lebanon, Turkey), and also
recorded from Q. pubescens in
Sicily (Barbagallo & Stroyan 1982). Sexual morphs were collected on Q. calliprinos in Turkey in November. Tuberculatus (Pacificallis) pallidus
(Davidson) Alatae are pale green; BL
1.5-2.2 mm. On leaves of a wide range of Quercus
spp. (including non-native oaks) in western USA. Partly anholocyclic; alatae
were collected in December-January in southern California. Separation from T. quercifolii, which is holocyclic on
native Californian oaks is difficult and needs further investigation. Tuberculatus (Orientuberculoides)
paranaracola Hille Ris Lambers Alatae are greenish yellow or yellowish
green to pale green (with 'dark green reflecting whitish'), with banded
antennae and dark tibiae (Quednau & Shaposhnikov 1988); BL 1.6-2.3 mm. On
undersides of leaves of Quercus
mongolica (incl. var. grosseserrata)
in Japan, China, Korea and eastern Siberia. Hille Ris Lambers (1974)
distinguished a subspecies, T. paranaracola hemitrichus, from
yellow traps in Korea, and this subspecies is also recorded from Q. acutissima and Quercus sp. in China (Chen & Zhang 1985b, Quednau 1999). The
life cycle and sexual morphs are unknown. 2n = 14 (for ssp. hemitrichus). Tuberculatus (Pacificallis) pasaniae
(Davidson) Alatae are pale green
mottled with darker green and with 'many delicate pruinose markings'
(Davidson 1915); eyes dark red, antennae ringed with black, spinal process
whitish; BL 2.0-2.6 mm. On undersides of leaves of Lithocarpus densiflora in California and Oregon, USA. An ovipara
collected on Lithocarpus in
November was illustrated by Quednau (1999), but the occurrence of alate
viviparae in November to mid-February indicates at least partial anholocycly. Tuberculatus (Pacificallis) passalus
Quednau Colour of alatae in life is
unknown; BL 1.3-2.2 mm. On Quercus
dumosa in California USA. Oviparae were collected in late
September-December, but it is apparently partially anholocyclic, as alate
viviparae occur in December and March (Quednau 1992). Tuberculatus (Acanthotuberculatus)
pilosulus Quednau
Colour of alatae in life is unknown; BL 1.6-2.5 mm. On Quercus serrata and Quercus sp. in Japan, attended by ants
(Yao 2011). Oviparae were collected in the second half of November (Quednau
1996). Tuberculatus (Nippotuberculatus)
pilosus (Takahashi)
Alatae are yellowish brown with blackish eyes, with forewing veins
having broad brown borders; BL 1.9-2.8 mm. Described from Quercus sp. in Taiwan (Takahashi 1929)
and redescribed from Japan on Q.
phillyraeoides (Higuchi 1970), but there are some discrepancies between
the two descriptions which warrant further investigation. Yao (2011)
identified a population on Lithocarpus
edulis in Japan as this species. Sexual morphs and life cycle are
unknown. Tuberculatus querceus
(Kaltenbach) Alatae are dirty
greenish to straw-coloured, covered with fine powdery white wax, with a dark
bifurcate spinal process (fig. 94A), a dark patch
near end of hind femur and dark siphunculi; BL 1.4-2.4 mm. On undersides of
leaves of Quercus robur in Europe
and east to Iran, Turkey and the Caucasus (Heie, 1982), Central Asia
(Kadyrbekov 2017a), and introduced to British Columbia, Canada (Foottit et al. 1999), and to South America
(Argentina, Chile; Caballero et al.
2000, Ortego et al. 2006). DNA
barcoding has raised the possibility that specimens from Europe (Poland) and
Canada may be different cryptic species (Y. Lee et al. 2017). Sexuales occur in October. Tuberculatus
(Acanthocallis) quercicola (Matsumura) Alatae are green with brown head and
thorax, banded antennae, dark hind femora, dusky wings with bordered wing
veins, and dark bars on abdominal tergites 1 and 2; BL 1.8-2.7 mm. On upper
and undersides of leaves of Quercus
mongolica (incl. ssp. crispula), attended by ants. Records
from other Quercus spp. may be
incidental; those from Q. dentata
should probably be referred to T.
macrotuberculatus (Watanabe et al.
2015). In Japan, Korea, China and eastern Siberia. Quednau & Shaposhnikov
(1988) noted some differences in life between populations on Q. dentata and Q. mongolica (but their aphids on Q. dentata were probably T.
macrotuberculatus, q.v.). Yao & Akimoto (2002), Yao
(2011) and Yao & Katagiri (2011) studied the
effects of ant attendance on composition of the honeydew and wing
development. Yao (2012) described the
life cycle and studied seasonal changes in the flight apparatus in
male and alate females. 2n = 16 (Chen & Zhang 1985b). [Also
note: 1. T. pappus Zhang, Zhang & Zhong
1990 is probably a synonym; oviparae collected in Japan on Q. mongolica var. grosseserrata (= ssp. crispula)
in late September, described under that name by Quednau (2003, p. 293-4), and
illustrated along with a male (2003, p. 299), should probably be referred to T. quercicola. 2. DNA barcoding of specimens from Korea has
revealed that aphids from Q. aliena identified
as T. quercicola may be another
distinct, cryptic, host-specific species (Y. Lee et al. 2017)]. Tuberculatus (Pacificallis) quercifolii
(Davidson) Alatae are pale green with
olive green head and mesothorax (Davidson, 1919); BL 1.2-2.4 mm. On leaves of
Quercus douglasii in California,
USA. Apparently holocyclic; fundatrices occur in April (BMNH collection, leg.
D. Hille Ris Lambers), and an ovipara collected in early December was
illustrated by Quednau (1999). Hille Ris Lambers (1974) probably had a
mixture of T. quercifolii and T. pallidus (see Quednau 1992). Tuberculatus (Orientuberculoides)
querciformosanus (Takahashi) Alatae are yellow or yellowish white with
yellow eyes and antennae ringed with black; BL 1.5-2.5 mm. On Quercus spp. (aliena, dentata, mongolica,
variabilis) in Japan, China,
Taiwan, Korea and eastern Siberia. Sexual morphs and life cycle are
unrecorded. Tuberculatus (Acanthotuberculatus)
radisectuae Zhang, Zhang & Zhong Alatae are brown and have wings dusky with
dark veins, black distal parts of hind femora, and tubercles on abdominal
tergites 2 and 3 black and arising from dark transverse bars (but those on
abdominal tergite 1 pale); BL c. 2.6 mm. On Quercus fabri in China (Zhang et
al. 1990b, as T. japonicus ssp.
radisectuae). 2n = 14 (Chen &
Zhang 1985b). Tuberculatus (Tuberculoides)
remaudierei Nieto Nafría Alatae are pale green with red eyes; BL 1.8-3.0
mm. On Quercus pyreniaca in Spain,
and there is also a record from Greece (Tsitsipis et al. 2007). Sexual morphs occur in October (Nieto Nafría &
Mier Durante 1978). Tuberculatus (Toltecallis) spiculatus
Richards Alatae are white to very
pale green; BL 1.9-2.4 mm. On Quercus
rugosa and Q. microphylla in
Mexico, and also recorded from Q.
arizonica in Arizona, USA (Quednau 1999). A subspecies, T. spiculatus rebecae, was described
from Q. rugosa in Mexico
(Remaudičre & Quednau 1983). Sexual morphs are unknown. Tuberculatus (Arakawana) stigmatus
(Matsumura) Alatae are shining dark
brown to bluish-black with banded antennae, pale fore legs and black hind
legs (see Moritsu 1983); BL 2.0-2.6 mm. Immatures are pinkish blue or pinkish
brown, with longitudinal rows of black spots. On Quercus spp. in Japan, Korea, China and eastern Siberia. DNA
analysis has indicated that the name may be being applied to two species with
minor morphological differences (Y. Lee et
al. 2017). Sexual morphs occur in October in Korea (Paik 1972). Tuberculatus (Orientuberculoides)
yokoyamai Takahashi
Alatae are pale greenish yellow or pale yellow with whitish eyes and
banded antennae (see Moritsu 1983); BL 1.6-2.4mm. On Quercus spp. (crispula,
dentata, mongolica, serrata) in Japan, China, Korea and
eastern Siberia. DNA barcoding has indicated that the name may be being
applied to two species with minor morphological differences (Y. Lee et al. 2017). An ovipara was collected
on Q. dentata in South Korea in
November (Quednau & Lee 2001; described and illustrated by Quednau 2003,
p. 292). 2n=14.
One
or two species associated with Hydrangea,
having long dorsal hair-bearing processes, and eyes without distinct ocular
tubercles. The life cycle is unknown;
possibly there is a host alternation to ferns, as in the related genus
Macromyzus. Tuberoaphis hydrangeae Tseng & Tao
Apterae are yellow, with apices of appendages dark brown; BL c.1.1-1.2
mm. On leaves of Hydrangea aspera in
Szechuan, China. Tuberoaphis hydrangeae ssp. digitata Hille Ris
Lambers & A.N. Basu Plate 22d (Fig.32a) Apterae are pale glassy white to pale
yellowish green, with apices of appendages darker; BL 1.4-1.7 mm. On
undersides of leaves of Hydrangea
robusta in West Bengal, India, producing large numbers of alatae in late
May and probably migrating to an unknown secondary host (original
description). Matsumuraja indica A.K.
Ghosh, M.R. Ghosh & Raychaudhuri, described from (?) Rubus sp., is a synonym, but Rubus
is an unlikely secondary host.
Fifteen or more species of Myzus-like aphids in east Asia, often with hair-bearing
siphunculi, galling the leaves of Prunus
in spring. Where the life cycle is known there is host alternation to
Anthemidae (Artemisia, Chrysanthemum), where they are small aphids on growing points
or undersides of leaves (Tuberocephalus
s.str.) or on subterranean shoots or roots (subg. Trichosiphoniella). The generations on secondary hosts
look very different from the spring forms on Prunus, and this has contributed to the taxonomic confusion in the
group. Accounts are available from Japan (Miyazaki 1971), China (Su et al. 2010) and eastern Russia
(Pashchenko 1988a). Moritsu & Tokumoto (1972) gave an account of Japanese
Tuberocephalus (under Myzus) on flowering cherries in which they
included several unnamed species. Sorin & Remaudičre (1998) reviewed the
genus and provided new information about morphs and life cycles. Tuberocephalus artemisiae Shinji Spring colonies are in large, bag-shaped
galls produced on the upper sides of the leaves of Prunus donarium “var. spontanea
subvar. speciosa” (= P. jamasukara
var. speciosa?) and P. yedoensis. The galls are yellowish
green becoming bright red on upper side when mature. Fundatrices are dark
green, BL c. 2.2 mm, and all their progeny are alatae (BL c.1.6 mm, with
greenish-yellow abdomen without a central patch), which migrate to Artemisia spp. Apterae on Artemisia are pale yellow to dull
yellow, body rather elongate, with pale appendages; BL 1.1-1.5 mm. They form
colonies on the undersides of the leaves, heavy infestations curling leaves
and causing plants to become sooty with mould (Miyazaki 1971). Gynoparae
and males return to Prunus in
October-November (Sorin & Remaudičre 1998). It is recorded from Japan,
China and east Siberia. Tuberocephalus (Trichosiphoniella)
higansakurae (Monzen) Spring colonies in galls caused by upward
rolling and thickening of edges of distal parts of leaves of cherry trees,
the sausage-shaped leaf-roll becoming yellowish-green or red. Dark green
fundatrices give rise to reddish-yellow apterae (BL 1.5-1.9 mm) and alatae
(BL c. 1.8-1.9 mm). Alatae have a dusky yellow abdomen with a dark central
dorsal patch and fly to an unknown secondary host. Greenish-white apterae of
a Tuberocephalus that could be this
species were collected on Gnaphalium
(BMNH collection, leg. R.L. Blackman), but this needs experimental
verification. On Prunus spp.
(cherries) in Japan, Korea and China; however, it seems that Zhang &
Zhong (1976) in China may have had T.
misakurae. Monzen (1929) recorded alatae (gynoparae) returning to P. subhirtella in October-November,
but did not describe the sexual morphs. Tuberocephalus (Trichosiphoniella)
jinxiensis Zhang & Zhong Described from apterae only (BL 1.4-1.8 mm),
collected on Prunus humilis in
China in June (Zhang & Zhong 1976). Barrel-shaped galls are formed on the
undersides of the leaves along lateral veins. The life cycle is unknown. It
seems very similar to T. higansakurae,
but with shorter siphunculi. Tuberocephalus lazikouensis Zhang, Chen, Zhong & Li, in Zhang Apterae are green; BL c.1.7 mm. On
undersides of leaves of Artemisia
argyri in Gansu province, China (Zhang 1999), and subsequently in Shaanxi
and Yunnan provinces (Su et al.
2010). Other morphs and life cycle are unknown. Tuberocephalus (Trichosiphoniella)
liaoningensis Zhang & Zhong Galls on Prunus tomentosa in Japan are formed from curled leaves and are
multi-coloured yellow and green with some reddish purple when mature (Sorin
& Remaudičre 1998). Fundatrices are deep blackish green, producing
blackish-brown apterae (BL 1.7-1.9 mm), and emigrant alatae that migrate to Artemisia vulgaris var. indica (in Japan). The colonies on Artemisia are found on subterranean young
shoots.
This species was originally described from China (Zhang & Zhong 1976),
where it is a common species on Prunus
pseudocerasus, with heavy infestations curling and withering young leaves
in spring, and it is also recorded from P. salicina (G. Zhang
1999) and Amygdalus
triloba (Su
et al. 2010). 2n=12 (Chen &
Zhang 1985a). Tuberocephalus (Trichosiphoniella)
misakurae Moritsu & Hamasaki Spring colonies live in bag- and
sausage-shaped galls formed by rolling and thickening of edges of distal
parts of leaves of cherry trees, as in the closely-related T. higansakurae. It is recorded from Prunus pauciflora, P.
pseudocerasus and P. takenakae
in Japan (Moritsu & Hamasaki 1983) and from P. pseudocerasus in China (BMNH collection, leg. V.F. Eastop).
The fundatrix is deep green or green; the colour of its apterous progeny is
not recorded, BL 1.5-2.0 mm. Alatae have a yellowish-green abdomen with a
dark central dorsal patch and migrate in May to form colonies on the thin
lateral roots of Chrysanthemum. Apterae on the
secondary host are greenish brown, BL 0.9-1.2 mm. Gynoparae,
males and oviparae were collected in Japan in October-early December (Sorin
& Remaudičre 1998). 2n=12. Tuberocephalus (Trichosiphoniella)
momonis (Matsumura) Spring colonies roll the edges of the
leaves of Prunus persica and turn
them red. Fundatrices are dark green, broadly oval, with BL0 2.1-2.2 mm (Su et al. 2010). Apterous fundatrigeniae
are yellowish-brown to dark brown with a greenish tinge (Matsumura 1917); BL
1.4-2.0 mm. Alatae have a pale yellowish-brown abdomen without a distinct
dorsal patch and are produced in late May to June (Su et al. 2010 and BMNH collection, leg. V.F. Eastop), but colonies
persist on peach into summer and autumn, so it seems likely that there is no
host alternation. Su et al. (2010)
described apterae collected in September, which were dark green with BL
1.4-1.8 mm, and also provided descriptions of alate gynoparae, alate males
and oviparae collected in October. In Japan, China, Taiwan, Korea and
Mongolia. This species has been confused in the literature with T. higansakurae, T. misakurae and T. sakurae.
The true momonis seems to be
specific to peach. 2n=12 (Chen & Zhang 1985a). Tuberocephalus naumanni
Sorin & Remaudičre Mature galls formed on Prunus cornuta are shiny,
thick-walled, pale green to white, 3-4 mm wide and 20-25 mm long. They
contain up to three generations of pale to bright green aphids, all the third
generation being emigrant alatae (BL 1.8-2.1 mm). Migration occurs in May to
an unknown secondary host. Only known from Pakistan (Sorin & Remaudičre
1998). Tuberocephalus pakistanicus
Remaudičre & Sorin Mature
galls on Prunus cornuta are
sausage-shaped, papery, pitted, 2.5-3.0 mm wide and 15-30 mm long, dark green
to red with reddish speckles and spots sometimes coalesced into a red patch
on the upper side of the gall. All the
progeny of the fundatrix are reddish-brown emigrant alatae, BL 1.6-2.1
mm, which emerge in early May and fly to an unknown secondary host (Sorin
& Remaudičre 1998). Gynoparae have been trapped in October
(Naumann-Etienne & Remaudičre 1995). It is only known from Pakistan. Tuberocephalus
(Trichosiphoniella) sakurae (Matsumura) Plate 17g Spring
colonies cause terminal leaves of Prunus
(Cerasus) spp. to become
extensively curled, twisted and yellowed or reddened (see Moritsu 1983).
Apterae on Prunus are deep green to
black (fundatrices paler), with black siphunculi; BL 1.5-1.8 mm. Immatures
are bright greenish yellow. Recorded from P.
cerasus, P. jamasakura, P. lannesiana, P. maximowiczii, P.
sachalinensis, P. serrulata, P.
triloba and P. yedoensis, and
from Japan, China, Korea and eastern Siberia (including Kuril Islands; as Sorbaphis kurilensis Ivanoskaya-Shubina
1966). It has recently been recorded from P.
serrulata in USA (New York, Connecticut; Skvarla et al. 2017). Alatae are without a distinct dorsal abdominal
patch. They migrate to the roots and subterranean young shoots of Artemisia spp. (Sorin & Remaudičre
1998). Apterae
on the secondary host are deep bluish green to black, BL c.1.6-1.7 mm, with
immatures greenish yellow. Alate males and gynoparae are produced
in October in Japan and were described by Sorin & Remaudičre (1998).
[N.B. Tao's (1966) sakurae was
probably Myzus yamatonis, and
Miyazaki's (1971) description is partly of misakurae.] Tuberocephalus sasakii
(Matsumura) Conspicuous, brownish, elongate pouch-like galls are produced in
Japan on the upper sides of leaves of Prunus
spp. between two veins (see Moritsu 1983). Progeny of the fundatrix are all
alate, with bright yellow abdomen; BL 1.2-1.8 mm. Alatae are without a
distinct dorsal abdominal patch. They leave the gall in June through an
opening on the underside of the leaf and fly to found colonies on Artemisia spp. (Monzen 1929, Sorin & Remaudičre 1998). Apterae on Artemisia are whitish or greenish,
dorso-ventrally flattened, BL 0.8-1.3 mm, and live adpressed to the
undersides of leaves The return migration occurs in
October (Monzen 1929). In Japan, Korea, the Kuril islands (Pashchenko 1988a),
Taiwan, China and India. It
is possibly anholocyclic in West
Bengal as it is common on Artemisia but not known to occur there on Prunus
(Ghosh 1973). The record from Java (Noordam 2004) applies to T. tsengi (q.v.). Tuberocephalus (Trichosiphoniella)
tianmushanensis Zhang
(= T. higansakurae ssp. hainnevilleae Remaudičre & Sorin; synonymy
by Su et al. 2010) Galls on Prunus are of two types; bag-shaped on the edge of tender leaves
formed by the fundatrices, and leaf-rolls similar to those described for the
nominal form, bright reddish yellow when mature and inhabited by their
apterous progeny. Fundatrices are matt brown, BL 1.7-2.0 mm, giving rise to
very dark apterae (BL 1.5-1.9 mm) and alatae (1.6-2.0 mm). Emigrant alatae
have a black dorsal abdominal patch. On Prunus
spp. (cherry trees) in China and Japan; also found in France on seedlings of P. subhirtella var. pendula imported from Japan
(Remaudičre & Sorin 1993, as T.
higansakurae ssp. hainnevilleae),
and it has possibly become established on the same host in northern Italy
(Pellizzari & Frigimelica 2014). Emigrant alatae colonised subterranean
young shoots of Artemisia in
transfer experiments in Japan (Sorin & Remaudičre 1998, as T. higansakurae ssp. hainnevilleae). Gynoparae, oviparae
and males were collected on Prunus
in mid- to late November. A redescription of the type specimens (from
Zhejiang province, China) was provided by Su et al. (2010), identifying the host as P. pseudocerasus. Tuberocephalus tsengi Tao Apterae are yellowish
green with dark red eyes, black tips to antennae, legs and siphunculi, and a
pale cauda (see
influentialpoints.com/Gallery); BL 1.2-1.9 mm.
On young growth of Artemisia spp. in China, Indonesia and West
Malaysia, and recently found (on A.
vulgaris) in southern England (Blackman et al. 2020). Presumably there is a primary host generation
forming galls somewhere on Prunus,
but this has still to be recognised, and its full distribution is uncertain
because of confusion with T sasakii,
with which it was erroneously synonymised. [In Japan there is an undescribed Tuberocephalus species on Gnaphalium sp., with shape of antennal
tubercles and ANT I, reduced eyes and sculpturing of dorsal cuticle very like
T. tsengi (BMNH collection, leg. R.
Blackman, RLB 2171). The apterae are greenish white in life, and compared
with T. tsengi it has shorter
antennae, longer R IV+V and shorter cauda.]
Tuberocephalus usubai Sorin The gall is undescribed and only the
emigrant alata is known, BL c.2 mm. On Prunus
buergeriana in Japan (Sorin 2011). The alata is very similar to that of T. naumanni, but has more rhinaria on
ANT III (7-12 compared with 3-8 in naumanni)
and a much longer R IV+V (c. 1.2 × HT II, compared with 0.7-0.8 × HT II in naumanni). Tuberocephalus (Trichosiphoniella)
uwamizusakurae Sorin & Remaudičre Fundatrices are dark green in life,
inhabiting elongate pouch-like galls along the margins of the upper sides of
the leaves of Prunus grayana in
Japan. When mature the galls are brownish green and about 10 mm wide and
50-60 mm long. Spring migrant alatae (BL 1.7-2.0 mm) are produced in the
second generation and migrate in April-May to found colonies on the curled
lateral roots of Artemisia vulgaris var. indica. Apterae on Artemisia
roots are probably dark green to black, BL 1.0-1.4 mm. An
ovipara was collected on P. grayana
in late November (Sorin & Remaudičre 1998).
Three species with a single large conical spinal
process or tubercle on abdominal tergite 4. A.K. Ghosh (1982b) provided an
account of the two species then known. It seems probable that the presence of
a dorsal tubercle is not a reliable character on which to base a genus, and
that some other Asian Salix-feeding
species currently placed in Lachnus (salicis, tatakaensis) may be more closely related to T. salignus than are the two species of Tuberolachniella. Tuberolachnus (Tuberolachniella)
macrotuberculatus Yang, Qiao & Zhang Apterae have a blackish-brown head and
yellowish-brown thorax and abdomen, the abdomen bearing a large bladder-like
tubercle; BL 5.1-5.3 mm. On Eriobotrya
japonica in Sichuan, China (Yang et
al. 2005a). Other morphs and biology are unknown. Tuberolachnus salignus
(Gmelin) Plate
15c, d Apterae are mid-brown to
dark brown with rows of dark spots and a large dark brown tubercle in the
centre of the dorsum (see influentialpoints.com/Gallery); BL 5.0-5.8 mm. Alatae have
the forewing membrane unpigmented. On stems and branches of numerous Salix spp. and also very occasionally
recorded from Populus. Large
colonies build up in late summer and attract numerous honeydew-feeding
insects. Virtually cosmopolitan in distribution, wherever willows grow or are
planted; only recently (2013) reported to occur in New Zealand ( et al. 2014) and Australia (2014; &2018).
Apparently anholocyclic everywhere; no sexual morphs are known. Mittler
(1957, 1958a, b) used this species in classic studies of aphid feeding and
nutrition, and Hargreaves & Llewellyn (1978) studied its ecological
energetics. Effects of temperature on development and fecundity on various
willow species were studied by Collins & Leather (2001) and Özder et al. (2007), and selection between
willow varieties was studied by Aradottir et
al. (2009). Genetic diversity of Chinese populations was studied by F.
Fang et al. (2017). In Japan, T. salignus is parasitised by a
possibly specific parasitoid, Pauesia
salignae, which might indicate that this is where it is endemic. 2n = 20. Tuberolachnus (Tuberolachniella)
sclerata Hille Ris Lambers & Basu Apterae are blackish with a conspicuous
reddish-brown to black broad-based tubercle on the dorsum; BL 4.0-5.2 mm. In
large colonies on undersides of leaves, petioles and young shoots of Eriobotrya petiolata in West Bengal
(at 2100 m); on leaves they form rows along the mid-rib and main veins (Hille
Ris Lambers & Basu, 1966). Specimens of this or a very similar species
have also been collected on E. japonica
in Indonesia (BMNH collection, leg. D. Noordam). Alatae and other morphs are
undescribed, and the life cycle is unknown.
A genus for one large Asian species with 5
hairs on first tarsal segments, clavate siphunculi and numerous rhinaria on
antennal segment IV of alatae. Tumoranuraphis indica (Chakrabarti & Maity) Apterae of BL about 4.6-4.9 mm, pale
greyish covered with powdery wax, with brown, clavate siphunculi (fig. 88C). Alatae have black head
and thorax and blackish-brown dorsal, abdominal markings. Fundatrices are greenish with dark dorsal
spots. It feeds along the mid-ribs on
upper sides of leaves of Prunus cornuta
in late May-June, causing the leaf laminae to fold upwards and come together
above the aphids. In Uttar Pradesh,
India (Chakrabarti & Maity 1984), and recorded from north-west China on P.
pseudocerasus (G. Zhang 1999, as T. cerasicola). Heteroecious holocyclic; alatae leave Prunus in late June to fly to an
unknown secondary host (Medda et al. 1986). Oviparae were collected in China (Eastop
& Blackman 2005).
Two
species on Asteraceae: Cardueae described from the desert zone of Kazakhstan,
related to Macrosiphoniella but
with an elongate R IV+V, a well-developed median frontal tubercle and 5 hairs
on first tarsal segments (Kadyrbekov 2002a). The two species are very similar
and the reported differences might be due to nutrition; Goodarzifar et al. (2016) provide comparative
data. Turanoleucon mitjaevi Kadyrbekov
Apterae are black, BL 2.2-3.2 mm. On stems of Cousinia
spp. in deserts of south and east Kazakhstan. Turanoleucon
jashenkoi Kadyrbekov Apterae are
black, BL 2.3-4.0 mm. On stems of Echinops spp. in deserts of south and east
Kazakhstan, and recorded also from Iran by Goodarzifar et al. (2016), who provide a redescription.
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