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SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order) S
A
genus for two or three species related to Iziphya, but these are longer-bodied aphids
with siphunculi placed further back. Dorsal hairs are mostly fan-shaped, but
on abdominal tergite 8 there is consistently a pair of backwardly pointing
processes bearing long rod-shaped hairs. One species (S. tuberculata) is known only from alatae trapped in Korea
(Quednau & Lee 2001). Saltusaphis scirpus Theobald
Plate 4g (Fig.18i,k; Fig.19a,b) Apterae are long-bodied, greyish yellow to
greenish yellow, with dark markings tending to form longitudinal bands, the
dorsum being powdered with a very thin layer of greyish white wax; BL 2.3-2.5
mm. Alatae have broad dark dorsal
abdominal cross bars and dark-bordered wing veins with spots at their apices,
and antennae with 10-21 rhinaria on III. On various Cyperaceae (Carex, Cyperus, Scirpus). In
Europe (not UK), across Asia to east Siberia, widely distributed in Africa,
also now in Argentina (Ortego et al.
2006), and with a single record from USA (Ohio, Smith 1940). Monoecious
holocyclic; oviparae and apterous males occur in September-October. [Saltusaphis lasiocarpae (Ossiannilsson)
on Carex lasiocarpa in Sweden (Heie
1982), is a seasonal variant of S.
scirpus according to Quednau (2010), and Saltusaphis kienshuensis Shinji
, described from Carex sp. in South
Manchuria, cannot be separated from S.
scirpus on the basis of the published description.] 2n=10.
One
North American species on Juniperus
with uniquely specialized morphology and therefore of uncertain phylogenetic
position, but perhaps closest to Siphonatrophia.
In addition to features mentioned in the key to juniper aphids, marginal
abdominal tubercles are absent and the reduced tarsi have hairless first
segments. Sanbornia juniperi
Pergande ex Baker Plate 13e, f, fig. 53C
Apterae are green, with a dorsal coat of
flaky, transparent wax (Boudreaux, 1949); BL 1.2-1.4 mm. Alatae have pale
brownish-green head and thorax. On Juniperus
spp. (scopularum, virginiana) in USA, and also recorded from Brazil on Cupressus lusitanica (Sousa-Silva & Ilharco 2001b) and J. chinensis (Bortolotto et al. 2013). Monoecious holocyclic;
oviparae and alate males occur in USA in October-November. Trioxys cupressicola may be a
specialized parasitoid.
A
small oriental genus with Pyrus as
primary hosts, related to Dysaphis
but with long hairs on body and antennae. Accounts are available for Japan
(Miyazaki 1971) and China (G. Zhang
& Zhong 1983b). The name Sappaphis
was used in a broader sense from 1952 to 1965 to include species now in Dysaphis. Sappaphis albinae Zhang, Chen, Zhong & Li Apterae are white, BL c.1.6 mm. On roots
of Artemisia scoparia in Gansu
Province, China (G. Zhang 1999). Possibly these are early-summer
exules of S. sinipiricola. Sappaphis angelicograstis Zhang, Chen, Zhong & Li Apterae are golden yellow, BL c.1.5 mm.
On roots of Artemisia argyi in
Gansu Province, China (G. Zhang 1999). Also recorded from Angelica sinensis, but this is
unlikely to be a true host. These are possibly late summer apterae and
gynoparae of S. sinipiricola. Sappaphis
piri
Matsumura Plate 10f (Fig.12q) Apterae in spring on Pyrus are broadly oval, brownish or
yellowish-brown, covered with white woolly wax; BL 1.8-2.5 mm. Alatae have
dark dorsal abdominal cross-bars, more weakly developed on anterior tergites;
secondary rhinaria are distributed III 21-28, IV 4-9, V 0-1. On undersides of
leaves of Pyrus spp. in east Asia
(Japan, Korea, China, Taiwan, eastern Siberia). Heteroecious holocyclic,
migrating in May-June to roots of Artemisia
spp. Apterae on Artemisia are pale yellow to
orange-red; BL 1.7-2.1 mm. The return migration to pear occurs in
October. Miyazaki
(1971) provided a full redescription. Kim et al. (1986) studied seasonal
population fluctuations on pears in Korea, and Takada & Hashimoto (1985)
studied relations with parasitoids and ants on Artemisia. [G. Zhang et al. (in G. Zhang 1999) described two new
species from north-west
China, S. albinae (q.v.) and S. angelicograstis (q.v.), and also
listed records from Artemisia of
two species previously described from Pyrus,
S. dipirivora Zhang and S. sinipiricola Zhang
(q.v.). These will all run to S. piri in
the key to aphids on Artemisia. Further
work is needed to confirm whether all or any of them are distinct from S. piri.] 2n = 12. Sappaphis sinipiricola Zhang
Described from apterae and alatae collected on Pyrus in Henan Province, China, in May, and subsequently recorded
from Artemisia (G. Zhang et al.
1999a). S. albinae and S. angelicograstis (q.v.) may be seasonal forms of this species,
which is itself very close to S. piri.
[Gynoparae of S. sinipiricola from
China have ANT PT/BASE 1.5-2.7,
whereas gynoparae of S. piri from
Japan and Korea have ANT PT/BASE 1.4-2.1. Another sample of alatae from pear
in October, described as S. montana Zhang,
in G. Zhang
(1999) may also be gynoparae of S.
sinipiricola.] 2n=12.
One
species of east Asian origin related to Tinocallis
but with a squat body, a very different pattern of dorsal sclerotisation,
wings that are folded almost flat over the body in repose, and a host
association with Lythraceae. It was placed as a synonym of Tinocallis by Remaudière &
Remaudière (1997) but revalidated as a full genus by Quednau (2003). Sarucallis kahawaluokalani
(Kirkaldy) Alatae are broad-bodied,
pale yellow or yellow-green with dark brown markings (dark longitudinal
stripes on head and prothorax, dark brown pterothorax, transverse marks on
abdominal tergites 1 and 2 incorporating the paired tubercles and
distinctively marked forewings); BL 1.2-1.8 mm. Immatures are greenish-yellow
with pale brown dorsal spots. On the undersides of leaves of Lagerstroemia spp. and also recorded
from Lawsonia alba (Agarwala et al. 1989a). Widely distributed in
east and south-east Asia; introduced to Europe (Netherlands, France, Spain,
Italy, Croatia, Greece, Bulgaria, Turkey), Africa (Congo; Remaudière & Le
Ru 1995), USA (where it is widely distributed, including Hawaii), Central
America (Puerto Rico, San Salvador, Panama) and South America (Argentina, La
Rossa et al. 1997; Columbia, Kondo
& Cortés 2014). Oviparae and alate males occur in September-October.
Patti (1984) gave a concise general account with coloured photographs. The
ecology was studied in Italy by Patti et
al. (1984), and in North America by Alverson & Allen (1992). Mizell
& Knox (1993) studied susceptibility of numerous Lagerstroemia cultivars in Florida. An extensive search failed to
reveal any specific parasitoids of this aphid (Mizell et al. 2002). 2n = 6.
About 36 palaearctic and six nearctic species
resembling Rhopalosiphum but with tapering
siphunculi and media of forewing only once-branched. Probably about half of
the species live all year on Poaceae,
and most of the rest (subgenus Paraschizaphis)
are on Cyperaceae and Typhaceae, but a few mainly Asian species overwinter as
eggs on Pyrus or Malus. More than half the species are European, and the
others occur in Middle East, Central Asia, east Asia, Africa and North
America. The S. graminum group
comprises several closely-related and morphologically very similar species or
subspecies, some of which can only be reliably identified by their
associations with different grass genera or species, or by other differences
in biology. The
taxonomy and life cycles of the pear-feeding species also require further
study. Eastop (1961b) provided a key to the world fauna then known.
Pear-feeding species were keyed by Shaposhnikov (1952), and included in
regional accounts for India (A.K. Ghosh & L.K. Ghosh 2006); eastern
Siberia (Pashchenko 1988a) and China (Tao 1962). Ilharco (2002) reviewed the world fauna of Paraschizaphis (treating it as a full
genus), and Leclant (1967) gave an account of the “S. cyperi group”. Regional accounts are available for UK (Stroyan
1984), north-west Europe (Heie 1986), Iberian peninsula (Nieto Nafría et al. 2005a), China (Zhang et al. 1998), Japan (Miyazaki 1988)
and east Siberia (Pashchenko 1988a). Schizaphis (Paraschizaphis) acori (Shinji) Apterae are very dark green-black, with
reddish tinge around bases of siphunculi; BL 1.8-2.4 mm. Alatae have
secondary rhinaria distributed III 11-15, IV 2-8, V 0-1. On Acorus calamus in Japan; specimens
from Cyperus difformis in Japan may
also be this species. Monoecious holocyclic, with apterous and brachypterous
males (Miyazaki 1988). This species has been erroneously synonymised in
catalogues with S. rotundiventris.
2n=8. Schizaphis agrostis Hille Ris Lambers Apterae are grass-green to yellowish
green with darker green spinal stripe; BL 1.4-2.1 mm. Alatae have secondary
rhinaria distributed ANT III 4-10, IV 0-2. On uppersides of leaves of Agrostis spp., especially alba and canina; sometimes on Poa
annua, particularly on dry, sandy soil, sometimes ant-attended. In
northern and central Europe, and aphids identified as this species have been
collected on an Elymus sp. in
Kurgan city (south-west Siberia; Novgorodova & Stekolshchikov 2013). Perhaps introduced to USA, as molecular and
morphometric studies (Kati et al.
2013) indicate that US “biotype H” of S. graminum may be this species.
Monoecious holocyclic, with oviparae and alate males in October. It is a
member of the S. graminum group,
all of which are very closely related. Morphometric discrimination of alatae
of this species from S. holci and S. graminum were discussed by Kati et al. (2013). Schizaphis aurea Miyazaki
Apterae are bright yellow with antennal flagellum and siphunculi
black, legs banded with black, and cauda pale; BL 1.6-1.9 mm. On Pennisetum alopeculoides in Japan.
Monoecious holocyclic with oviparae and alate males in mid-November (Miyazaki
1988). Schizaphis borealis Tambs-Lyche
Apterae are pale, colour not recorded; BL 1.3-1.9 mm. Alatae have
secondary rhinaria distributed ANT III 4-6, IV 0-1. On Phleum pratense in Fennoscandia. Schizaphis (Paraschizaphis)
brachytarsus (Takahashi) Apterae are brown
with rather long black siphunculi; BL 1.4-2.3 mm. Alatae are unknown.
Described from a plant of family Cyperaceae in Japan, and since recorded from
Carex in Japan (neotype, described
by Ilharco 2002) and recorded from Scirpus
in east Siberia (Pashchenko 1988a). Schizaphis (Paraschizaphis) caricis (Schouteden) (Fig.18f) Apterae are shiny blackish brown to
blackish green; BL 1.3-2.4 mm. Alatae have secondary rhinaria distributed III
7-20, IV 2-10, V 0-5. On Carex and Scirpus, living in small compact
colonies on basal parts of leaves, and often in ant “shelters”. In Europe,
eastwards to Ukraine, and in east Siberia (Pashchenko 1988a, as caricicola) and USA (Ilharco 2002).
Monoecious holocyclic, with apterous males (Heie 1986). Schizaphis chaenometicola Zhang
Apterae are yellowish green, BL c.1.5 mm. On leaves of Chaenomeles
speciosa in China (Gansu Province). Other morphs have not been described
(Zhang et al. 1998). Schizaphis cuprea Miyazaki
Apterae are dull brown with black antennae and siphunculi, and
yellow-brown legs ringed with black; BL 1.5-1.8 mm. On Calamagrostis sp. in Japan. Monoecious holocyclic with oviparae
and alate males in late September-October (Miyazaki 1988). Schizaphis dubia Huculak
Apterae are green or greenish yellow; BL 1.6-2.1mm. Described from Calamagrostis canescens, and there are
also records from Agrostis stolonifera and
Bromus arvensis. Alatae have 5-8
secondary rhinaria on ANT III. Pettersson (1971c) studied host preferences.
In northern Europe (Finland, Poland, Sweden). A record from Turkey (Şenol et al. 2014b) requires further confirmation. Monoecious holocyclic with alate males (original
description). Pettersson (1971b) provided morphometric data and illustrated
four morphs. Schizaphis (Paraschizaphis) eastopi (van Harten &
Ilharco) Apterae are black, BL
1.4-2.0 mm. Alatae have secondary rhinaria distributed III 7-10, IV 1-4, V
0-1. On Typha capensis in southern
Africa (Angola, Mozambique). Records of this species in the literature from
other countries in Africa, and from Italy (Sicily) and Pakistan, should be
referred to S. rosazevedoi (Ilharco
2002). Schizaphis gracilis Richards Apterae are undescribed, and
appearance of alatae in life is unknown; BL of alata 1.4-1.8 mm. On Carex maritima in Northwest
Territories, Canada. Schizaphis graminum (Rondani)
'Greenbug' Plate 9b Apterae have head and prothorax
yellowish or greenish straw‑coloured, rest of thorax and abdomen
yellowish green to bluish green with a darker spinal stripe (see influential
points/Gallery); BL 1.3‑2.1 mm. Alatae have secondary rhinaria
distributed ANT III 4-10, IV 0-4, V 0-1. On leaves of very many genera and
species of Poaceae, often causing yellowing and other phytotoxic effects. It
is sometimes attended by ants (Orlob 1963). Records from grasses in western
Europe are now thought to apply to other closely-related species or
subspecies with more specific host associations (agrostis, borealis, dubia, holci, jaroslavi, phlei, thunebergi), all of which are very difficult to tell apart (e.g.
Pettersson 1971b). S. graminum is
of palaearctic origin, but now widely distributed; southern Europe, Middle
East, Central Asia, Africa, India, Nepal, Pakistan, Thailand, Korea, Taiwan,
Japan, and North, Central and South America. Two alatae trapped in UK had
identical mitochondrial and nuclear DNA sequence to the sorghum-adapted form
in the USA (Kati et al. 2013), but
there are no records from field crops in northern Europe. Records from
Australia and Philippines are referable to S. hypersiphonata. A population without marginal tubercles on an
unidentified grass (possibly Agrostis)
in Iceland was described as a subspecies, S. graminum ssp. gigjai Stroyan. The North American “biotypes” have been shown by mtDNA
analysis to fit into three clades with different host relationships, the
divergence of which predates modern agriculture (Shufran et al. 2000), and this is supported by analysis of nuclear
sequences (Shufran 2011). This indicates that these were three separate
introductions, and that palaearctic S.
graminum may be a complex of still-unrecognised incipient species or
subspecies (Blackman & Eastop 2007).
S. graminum is monoecious
holocyclic, with alate males, in cold temperate climates, e.g. in northern
USA, where overwintering in the egg stage occurs predominantly on Poa pratensis (although the
sorghum-adapted form is able to produce sexuales at more southerly latitudes,
indicating that it probably originated from further south in Europe or Asia).
S. graminum is anholocyclic
wherever winter conditions permit. 2n=8. Schizaphis hierochlophaga Zhang & Chen Apterae are yellowish green; BL c. 1.6
mm. Alatae have secondary rhinaria distributed III 15-16, IV c.9, V 6-7. On Cymbopogon sp.(spp?) in Beijing, China
(Zhang et al. 1998). Schizaphis holci Hille Ris Lambers Apterae are greenish yellow or pale
yellow, with dark-tipped siphunculi; BL 1.6-1.9 mm. Alatae have secondary
rhinaria distributed ANT III 7-8, IV 0. On leaves of Holcus spp., usually attended by ants; infested leaves turn brown
and die. In Europe (UK, Netherlands, France, Germany, Austria, Czech
Republic). Monoecious holocyclic with apterous males. Kati et al. (2013) studied host specificity
and sexual morph production in the laboratory, and also discussed
morphological discriminants between this species, S. agrostis and S. graminum. Schizaphis hypersiphonata A.N. Basu
Apterae are yellowish‑green, BL 1.1-1.7 mm. On various Poaceae (Brachiaria, Chloris, Dactylotenia, Digitaria, Eleusine, Panicum, Phalaris, Phragmites, Triticum), and also recorded from Cyperus exaltata, but without having
the phytotoxic effects of S. graminum.
In West Bengal, Sri Lanka, Thailand, Philippines, Irian Jaya (Indonesia) and
Australia (Eastop 1966, as a form of graminum).
No sexual morphs have been recorded. Mordvilko's (1921) description of S. graminum is possibly referable to
this species. Schizaphis jaroslavi (Mordvilko)
Apterae are pale green to yellow greeen with a darker green median
stripe and dark tips to siphunculi, slightly wax-powdered; BL 1.5-2.1 mm. Al.
have 3-6 secondary rhinaria on ANT III only. Recorded from various grasses (Calamagrostis, Phalaris, Phleum, Secale) in Europe (Scotland,
Netherlands, Denmark, Sweden, Finland, Poland, Ukraine) and Asia (Kazakhstan,
Mongolia, Korea, Japan). However, the name is perhaps being applied to more
than one species, as in Europe S.
jaroslavi occurs on Calamagrostis
epigeios in dry sandy habitats (Heie 1986), whereas Japanese populations
have a shorter antennal terminal process than European ones and were only
found on the wetland grass Phalaris
arundinacea (Miyazaki 1988). In Korea it is recorded from a dicot, Cocculus trilobus (Lee et al. 2002c). The European aphid has
apterous males, and is often ant-attended. Schizaphis longicaudata Hille Ris Lambers Apterae are greenish, with black-tipped
siphunculi; BL 1.8-2.2 mm. Alatae have secondary rhinaria distributed ANT III
6-10, IV 0-1. On grasses (Alopecurus,
Apera, Bromus, Luziola, Phalaris); Pettersson (1971c) studied
host preferences. Widely distributed in Europe. Monoecious holocyclic with
alate (sometimes brachypterous) males (Heie 1986). Schizaphis (Paraschizaphis) longisetosa
(Higuchi) Apterae are dark green to nearly black;
BL 1.9-2.3 mm. Alatae have secondary rhinaria distributed III 12-19, IV 2-7,
V 0-3. On Carex spp. in Japan
(Higuchi 1970). Ilharco (2002) provided a redescription, noting that the
original description included specimens of S. brachytarsus. Schizaphis mali Shaposhnikov Apterae
are yellowish green with concolorous siphunculi and cauda; BL c. 2.8 mm. In
spring colonies on Malus baccata in
Russia, curling the leaf into a pod-shaped pseudogall. Migration occurs in
June to Carex spp. Appearance of apterae
on secondary host is undescribed; BL c.1.8-1.9 mm. Alatae produced on Carex have secondary rhinaria
distributed III 12-14, IV 2-6, V 0. 2n=8. Alate males
were collected on Carex, and
oviparae on Malus, in early October
(Shaposhnikov 1979b). This species seems closely related to species alternating from Pyrus to Cyperaceae in Russia (S. pyri) and Japan (S. piricola). 2n=8. Schizaphis minuta (van der Goot)
Apterae are green, BL 1.2-1.8 mm. Alatae have secondary rhinaria
distributed III 5-7, IV 2-3, V 0-1. On Cyperus
spp. It is usually found on small, short-leaved species growing in sand,
for example along dry stream beds. In Africa, Iran, east and south-east Asia
(India to Philippines), Australia and also recorded from Scirpus lacustris in Italy (Patti 1983), and in suction trap
samples in Florida, USA (Halbert et al.
2002). In Korea three samples were collected from Mazus miquelii (Lee et al. 2002c). Sexual morphs are
apparently unknown. Schizaphis muhlenbergiae (Phillips & Davis) Apterae are pale green, BL c 1.3 mm. On Muhlenbergia sp., which appears to be
the normal host; also on Poa pratensis,
and colonies were established on wheat under
insectary conditions (original description). In USA (Indiana, Ohio, Colorado,
Utah). A record from Lithuania on Leymus
arenarius (Rakauskas et al.
2008) needs further confirmation. Monoecious holocyclic with apterous males and
oviparae in October. Schizaphis nigerrima (Hille Ris Lambers) ( = Schizaphis
laingi Eastop; see Eastop & Blackman 2005) Apterae are coal black, BL 1.7-2.3 mm.
Alatae have secondary rhinaria distributed ANT III 4-7, VI 0-1. On upper
sides of leaves of Festuca pratensis,
yellowing the tips, and also recorded from Alopecurus pratensis and Calamagrostis
spp. In Europe (UK,
Netherlands, Finland, Germany, Poland, Austria, Czech Republic and Hungary).
Monoecious holocyclic with oviparae and alate males in October (original
description). This species is very similar to S. cuprea in Japan. Schizaphis nigra (Baker) (
= Toxoptera viridirubra Gillette
& Palmer; see Eastop & Blackman 2005) Apterae are blackish green (original
description; Washington DC) or reddish brown to olive green with orange
around bases of siphunculi (original description of viridirubra, Colorado); BL 1.1-2.0 mm. Alatae have secondary
rhinaria distributed III 6-14, IV 3-7. V 0-3. On Carex spp. in USA. Monoecious holocyclic, with apterous,
brachypterous and alate males (original description) or alate males (viridirubra, original
description). Ilharco (2002)
redescribed apterous and alate viviparae (as Paraschizaphis nigra). Schizaphis (Euschizaphis) palustris (Theobald) (Fig.35c,d) Apterae are yellowish brown to olive brown
or dark green; BL 1.5-1.8 mm. Described from Triglochin palustre, and also recorded from T. maritima and Poa annua, but the more common hosts are Juncus spp. In marshy situations in
Europe (UK, Netherlands, Germany, Slovakia, Spain (BMNH collection, leg. D.
Hille Ris Lambers – “Franz Sp.145”), Ukraine (Mamontova 1963) and India (New
Delhi; BMNH collection, coll. R. Menon). Monoecious holocyclic with apterous
males (Hille Ris Lambers 1947b). Schizaphis (Paraschizaphis)
pashtshenkoae (Ilharco) Apterae are dark
green to almost black; BL c.2.0 mm. Alatae have secondary rhinaria
distributed (1 specimen) III 15-16, IV 7, V 4. On Carex sp. in east Siberia (Ilharco 2002). Schizaphis phlei Orlob
Apterae are yellow to pale yellowish, to pale green, with siphunculi
dark-tipped; BL 1.2-1.9 mm. Alatae have secondary rhinaria distributed ANT
III 4-6, IV 0-1. On various grasses (Aegilops, Alopecurus, Apera, Arrenatherum, Phleum)
in Sweden and Finland, with Arrhenatherum
elatius as a preferred host (Pettersson 1971c, as arrhenatheri), and found causing brown leaf chlorosis on Phleum pratense in New Brunswick,
Canada (original description). Monoecious holocyclic with oviparae and alate
males in September (original description, and Pettersson 1971b). It is a
member of the S. graminum group.
Pettersson (1970, 1971a) studied the sex pheromone of this species (as borealis). Pettersson (1971b) provided
morphometric data and illustrated four morphs (as arrhenatheri). Schizaphis pilipes (Ossiannilsson) (Fig.18g) Colour of apterae in life is unknown,
probably dark brownish; BL 1.6-2.4 mm. Alatae have secondary rhinaria
distributed III 17-20, IV 7-13, V 2 (Heie 1986). On Carex acuta and possibly
other Carex spp. in Europe.
Monoecious holocyclic; oviparae were found in Sweden in November
(Ossiannilsson 1959). Also recorded
from Finland, Denmark, Netherlands, Germany, Czechoslovakia and UK. Schizaphis piricola
(Matsumura) Apterae are
yellowish-brown or green, with dark antennae and black-tipped siphunculi; BL
c. 1.4 mm. Spring colonies live in rolled edges of leaves of Pyrus pyrifolia (Essig & Kuwana
1918), and it has also been recorded from other Pyrus spp. The Schizaphis
recorded from Sorbus amurensis (= S. pohuashanensis) in Korea by Paik (1972) was also probably this
species. In Japan, Korea and China, and recently introduced to Spain, where it
was found colonising P. calleryana,
and attended by the ant Tapinoma
nigerrimum (Pérez-Hidalgo et al.
2011a). Alatae are produced in May in Japan and migrate to Cyperus rotundus (Miyazaki 1988). They
are also found on other Cyperus and
Carex spp. Apterae on secondary hosts are yellowish green to
deep green, with black siphunculi and antennae black except at bases; BL
0.96-1.85 mm. Alatae have secondary rhinaria distributed III 15-22, IV 7-11,
V 0-5 (cf. rotundiventris). S. piricola is very similar to S. pyri and S. mali in Russia. 2n=8 (Chen & Zhang 1985b). Schizaphis priori Stroyan
Colour of apterae in life is unrecorded, probably pale green with
darker spinal stripe; BL 2.2-2.6 mm. On Leymus
arenarius in Iceland. Monoecious holocyclic, with apterous males
collected in late July (original description, as S. geijskesi ssp. priori).
Schizaphis pyri
Shaposhnikov Apterae are green, often
with darker green longitudinal stripe, and dark siphunculi and cauda; BL
1.4-2.2mm. In spring colonies on Pyrus
communis, feeding on upper sides of leaves and folding them
longitudinally in half, with the underside turned outward, to form a similar
pod-shaped pseudogall to that of S.
rotundiventris (Shaposhnikov 1952). Presumed to be heteroecious
holocyclic, alatae migrating in May to form colonies on Cyperaceae, although
this still needs experimental confirmation. Apterae on secondary hosts are pale to dusky
green with black antennae and siphunculi, and a dark cauda; BL 1.5-2.0 mm.
They live singly on the undersides of leaves of Carex (Leclant 1967, as fritzmuelleri)
and Cyperus. Alatae produced on Carex in France had secondary rhinaria
distributed III 20-29, IV 7-15, V 4-5. In continental Europe (France, Spain,
Portugal, Czech Republic, Moldova, Russia, Ukraine), Atlantic islands
(Azores, Canaries, Madeira), and also recorded from Israel, Iran, Kazakhstan,
west Siberia, east Siberia (Pashchenko 1988a), and Australia
(Victoria). Sexual
morphs have not been described; secondary host populations are more widely distributed
and are probably mostly anholocyclic. Schizaphis (Paraschizaphis) rosazevedoi
(Ilharco) Apterae are of two colour-forms,
orange-brown to rusty brown, or dark green to almost black; BL 1.3-2.4 mm.
Alatae have secondary rhinaria distributed III 6-16, IV 0-6, V 0-2. On Typha spp. and Strelitzia reginae. Recorded from
Portugal, Madeira, France, Sicily, Israel, Iran, Pakistan, Saudi
Arabia, Sudan, Burundi, South Africa and Martinique. (Some of these records
were as misidentified S. eastopi). Apparently anholocyclic (Ilharco
2002). 2n=8. Schizaphis rotundiventris (Signoret) Apterae on Pyrus are greyish or blackish, dusted with white wax, with black
siphunculi and cauda; BL c. 1.9 mm. Feeding by fundatrices on developing
leaves of Pyrus communis in early
spring prevents them from unfolding and causes the edges of the leaves to
stick together forming a slightly inflated pod-shaped pseudogall (Das 1918,
as S. punjabipyri). Pear-feeding
generations are recorded from northern India, Pakistan and Nepal. In Pakistan
they are common in the plains and lower hills, especially in humid,
monsoon-exposed areas (Naumann-Etienne & Remaudière 1995). Alatae migrate
in May to Cyperaceae. Colonies occur most commonly on Cyperus
spp., although it can form small colonies in young unfolding leaves of palms such as Elaeis guineensis, and there are also records from Poaceae (Pennisetum, Polytrias)and other monocots (Acorus,
Typha). Apterae on secondary hosts
are dark green to almost black; BL 1.2-1.7 mm. Alatae have secondary rhinaria
distributed III 6-18, IV 0-10, V 0-5. Presumed anholocyclic populations on Cyperus and other monocots are
recorded from southern Europe, the Middle East, Africa, Mauritius, east and
south-east Asia, Australia, New Zealand, Hawaii, and most recently
south-eastern USA (Halbert et al.,
2000) and Costa Rica (Sánchez-Monge et
al. 2010). In Pakistan
gynoparae and alate males return to Pyrus
in early December or earlier, oviparae laying eggs in late December-January
which hatch in February-March. [The synonymy with S. punjabipyri (Das) proposed by Shaposhnikov (1979a) is followed
here, but still requires experimental confirmation.] 2n=8. Schizaphis rufula (Walker)
Apterae are yellowish green with darker green spinal stripe, slightly
wax-powdered, with dark- tipped siphunculi; BL 1.4-3.1 mm. Alatae have
secondary rhinaria distributed ANT III 4-8, IV 0-2. Pettersson (1971b)
provided morphometric data and illustrated four morphs. On leaves of various
grasses, especially in sand-dune habitats (Aegilops, Agropyron, Agrostis, Alopecurus, Apera, Ammophila, Bromus, Elymus, Leymus). Infested leaves become
yellowish brown. Pettersson (1971c) studied host preferences, and Van Moorleghem & de la Peña (2016) studied the
effects of its feeding on various dune grasses. Records from Salsola, including the original description, are presumably
casual occurrences. In north-west
and northern Europe. Monoecious holocyclic with apterous males. Schizaphis (Paraschizaphis) scirpi (Passerini) Plate 9c Apterae are shiny dark bronze-brown to
reddish brown or blackish (see influential
points/Gallery); BL 1.5-2.8 mm. Alatae have secondary rhinaria
distributed III 7-16, IV 0-8, V 0(-1). In ant-attended colonies at leaf-bases
of Typha and Sparganium, also on Cyperaceae
(Carex, Eriophorum, Scirpus)
and sometimes on Araceae (Colocasia),
Juncaceae (Juncus) or Iridaceae. Throughout Europe, and
in south-west Asia, China, Korea and Japan. Monoecious holocyclic, with alate
males (cf. S. caricis). European
populations on Eriophorum spp. are
regarded as a subspecies, S. scirpi ssp. eriophori Müller. 2n=8. Schizaphis (Paraschizaphis) scirpicola Hille Ris Lambers Colour of apterae in life is unknown;
1.4-1.7 mm. Alatae have secondary rhinaria distributed III 9-10, IV 3, V 0 (1
specimen, redescribed by Ilharco 2002). On Scirpus atrocinctus in Canada (New Brunswick). It has
subsequently been recorded from Scirpus
spp. and Carex sp. in Ontario,
Nova Scotia, Quebec and Manitoba, and from North Carolina, USA. Schizaphis thunebergi Heie
Apterae are pale, colour in life unknown; BL c.2.1-2.3 mm. The single
known alata has 6 secondary rhinaria on ANT III only. On Calamagrostis purpurea in Finland. Schizaphis variegata Ossiannilsson
Apterae are broadly oval, mottled dark green and brown, BL c.1.8-2.0
mm. On Carex juncella in Sweden
(Ossiannilsson 1959). Alatae and biology are unknown. Schizaphis wahlgreni (Ossiannilsson)
(Fig.18h) Colour of apterae in life is unknown,
probably pale, with black antennae (except at base) and siphunculi, and a
pale cauda; BL c.2.1 mm. On Carex
vesicaria in Sweden, Poland, Czech Republic and Hungary (Heie 1986), and
also found on Eleocharis palustris
in Latvia (Rupais 1989a). Alatae have been recorded (Ossiannilsson 1964b),
but are undescribed. Schizaphis weingaertneriae Hille Ris Lambers Apterae are dark green powdered with
bluish grey wax; BL 1.2-1.7 mm. Alatae have secondary rhinaria distributed
ANT III 5-8, IV 0-2. On undersides of leaves of Corynephorus canescens, usually arranged in single file. In
Netherlands, Poland and Czechoslovakia. Oviparae and apterous males were
collected in early October (original description). It is not visited by ants. Schizaphis werderi Börner
Colour of apterae in life is unrecorded, BL 1.6-1.9 mm. On Poa alpina in the Alps.
Seven species of oval-bodied, hairy, wax-secreting
aphids, all feeding on Pinus
needles, four nearctic and three palaearctic. Phylogenetic relations with Cinara and Eulachnus were studied by Sorensen (1990), who concluded that Schizolachnus was more closely related
to Eulachnus. However, a recent
molecular phylogenetic study combining both nuclear and mitochondrial data
placed Schizolachnus within Cinara and proposed that it should be
treated as a subgenus of that genus (R. Chen et al. 2016). Hottes (1956a) keyed the North American species
then recognized. Heie (1995) and Nieto Nafría et al. (2002a) reviewed the two European species. Pauesia unilachni is a specialized
parasitoid. Schizolachnus curvispinosus
Hottes, Essig & Knowlton Colour
of aptera in life is not recorded, 'probably a dusky green', with black hind
tibiae; BL 2.1-2.8 mm. On needles of Pinus
ponderosa, and occasionally found on non-native pines, in western North America.
Oviparae and alate males occur in October (Hottes et al. 1954). The alata was described by Hottes (1956d). Schizolachnus flocculosus
(Williams) Apterae have a black head
and thorax with grey wax markings, abdomen light brown or green with narrow
cross-bands of wax (Hottes 1952a; as S.
wahlea) and black hind legs; BL 2.4-2.9mm. On needles of Pinus arizonica and P. ponderosa in US Rocky Mountain
region (Colorado, Utah) and Arizona; there are now also records from the
Great Smoky Mountains in eastern USA, on P.
rigida and P. virginiana (Favret
et al. 2010). Oviparae and alate
males in September-November (Hottes 1954b; as S. wahlea). Schizolachnus obscurus Börner Apterae are brownish with greyish white
covering of wax powder (see
influentialpoints.com/Gallery); BL 1.9-2.7 mm. On needles of Pinus nigra (also sometimes on mugo, pinaster and pinea) in
Europe, east to Turkey (Klimaszewski & Wojcieckowski 1976) and Georgia
(Barjadze et al. 2010a), and
introduced to North America, where it was originally identified as C. pineti (see
Skvarla et al. 2017) .
In Spain, fundatrices were found in April-May, alatae in May, and oviparae
and males from the end of October to December (Nieto Nafría et al. 2002a). Schizolachnus orientalis
(Takahashi) Apterae are
yellowish-brown in original description from Taiwan (Takahashi 1924a), but
recorded as blackish-brown in India (A.K. Ghosh 1982b), covered in fine,
dirty white wax meal; BL 1.9-2.4 mm. Feeding in rows along needles of Pinus spp. in India, Japan, Korea,
China and Taiwan. A record from Turkey (Şenol et al. 2014b) requires further confirmation. Oviparae and
alate males occur in October-November (Inouye, 1970). Schizolachnus parvus
(Wilson) Apterae are brownish tinged
with green, covered with bluish-white wax threads which also extend onto
needles; BL 1.6-1.8 mm. On needles of Pinus
spp. in eastern USA, and recently recorded from Costa Rica (Villalobos Muller
et al. 2010). Oviparae and alate
males occur in USA in late October (Wilson 1915). Schizolachnus pineti
(Fabricius) Plate
16e Apterae are dark greyish-green
covered in wax meal giving a light bluish-grey appearance (see
influentialpoints.com/Gallery); BL 1.2-2.5 mm. On numerous Pinus spp., especially on young trees,
forming dense colonies in rows along the previous year's needles. Oviparae
and alate males occur in October-December; in some years colonies persist
into early winter and possibly anholocyclic overwintering occurs in mild
winters (Carter & Maslen 1982). Common and widespread in Europe; also in
Kazakhstan, east Siberia (Pashchenko 1988a) and China (Zhang & Zhong
1985e) although some of these records may apply to S. obscurus. Holman (1990) studied morphological changes in
spring generations, Kidd et al.
(1985) studied the association with Eulachnus
agilis and Thompson (1977) studied its effect on tree growth. 2n = 18
(Russia; Rukavishnikov 1972) or 2n = 10 (England; Blackman 1980); the first
chromosome number may possibly apply to S.
obscurus. Schizolachnus
piniradiatae (Davidson) Plate 16f Apterae are dark olive green to brown,
with grey flocculent wax coating; BL 2.0-2.5 mm. On needles of numerous Pinus spp., widely distributed in
North America. Oviparae and alate males occur in September-November. The male genitalia were
described and illustrated by Wieczorek (2012). There have
been studies in Canada of its life history and ecology (Gröbler 1962),
population dynamics on Pinus resinosa
(Sharma & Laviolette 1968a), oviposition behaviour (Sharma &
Laviolette 1968b), predators (Gagne & Martin 1968) and fungal diseases
(Gröbler et al., 1962; Tyrrell
& Macleod, 1975).
Eight
species are known, at least one of them having host alternation between Distylium galls and Litsea (in Java; Noordam 1991).
Closely related to Nipponaphis, but
the aleyrodiform exules have long, basally thick dorsal (prosomal) hairs. Schizoneuraphis
gallarum van der Goot
Galls on Distylium stellare
in Java are oval, balloon-shaped, 25-50 mm long, apparently arising from the
stem but formed from the leaf (fig. 130J);
pale green to peach-coloured when young, becoming dark grey-brown and woody,
but with a thin circular window in the side which becomes the emergence hole
for alatae in August-October. Emigrant alatae (BL 1.6-2.2 mm) have secondary
rhinaria distributed III 17-22, IV 8-11, V 4-8. Apterous exules feeding along
the veins on the undersides of leaves of Litsea
glutinosa are aleyrodiform, shiny dark brown to black, with upright pale
brown dorsal hairs and a white marginal band of wax; BL 0.7-0.9 mm. The host
alternation still needs to be confirmed experimentally. Hille Ris Lambers
& Takahashi (1959) claimed to have established it on the basis of the
similarity of embryos within emigrant alatae to first instars on Litsea, but Noordam (1991) noted some
differences. Schizoneuraphis
himalayensis (Ghosh & Raychaudhuri) Apterae are broadly oval, dark brown to
black; BL 0.9-1.1 mm (A.K. Ghosh & Raychaudhuri 1973b, as Nipponaphis). In colonies on leaves
and stems of Litsea spp., Lindera sp. and Persea bombycana (= Machilus
gamblei) in India (Assam, West Bengal, Manipur). Alatae (sexuparae?) were
produced in January. The life cycle is unknown. Nipponaphis machiliphaga of A.K. Ghosh et al. (1971d) seems to be this species. A record from Quercus dilatata (Holman 2009) needs
verification. Schizoneuraphis
litseicola Noordam
Apterae on the undersides of leaves of Litsea amara (= umbellata)
are aleyrodiform, shiny brown-black, similar in life to S. gallarum; BL 0.8-0.9 mm. Only known from Java (Noordam 1991).
The life cycle is unknown. The description is very similar to that of S. machiliphaga (Takahashi), which has
also now been recorded from Litsea. Schizoneuraphis
siamensis (Takahashi) described as shining purplish brown without wax and
feeding along mid-rib on the undersides of leaves an unidentified tree of
Lauraceae in Thailand (Takahashi 1941a, as Thoracaphis), could be this or a very similar species. Schizoneuraphis
longisetosa Noordam Galls on Distylium stellare in Java are not properly described, but
believed to be semiglobular, turning red when mature (possibly gall no. 487
of Docters van Leeuwen-Reijnvaan & Docters van Leeuwen 1926). Emigrant
alatae (BL 2.4-2.8 mm, secondary rhinaria distributed III 25-31, IV 12-16, V
8-13) were collected in late May (Noordam 1991). Secondary host populations
are unknown. Schizoneuraphis
machiliphaga (Takahashi) Apterae are almost circular, dorsally
flattened, reddish-brown, not strongly sclerotised; BL 0.8-1.0mm. On the
undersides of leaves of Machilus
sp. in Japan (Takahashi 1959b), and recorded from Litsea polyantha and Litsea
sp. in India (Assam, West Bengal, Meghalaya; A.K. Ghosh 1988, as Nipponaphis). The life cycle is
unknown. Schizoneuraphis
malayna (Takahashi) Apterae are aleyrodiform, almost
circular, dorsally flattened, shiny brownish-black, without evident wax; BL
c. 1.3mm. On undersides of leaves of Quercus
sp. in Cameron Highlands of Malaya (Takahashi 1950, as Thoracaphis malaynus). The life cycle is unknown. Schizoneuraphis
querciphaga (Ghosh and Raychaudhuri) Apterae are
yellowish to dark brown; BL 1.3-1.5 mm. On apical stems of Quercus sp., attended by ants, in
Sikkim, India (A.K. Ghosh and Raychaudhuri 1973b), and also on Castanopsis in Nepal (Miyazaki 1977).
The life cycle is unknown. Very similar to S. malayna.
One
little-known North American species closely related to Eriosoma but lacking siphunculi in morphs developing on Crataegus, the presumed secondary host.
The primary host is unknown, but presumed to be Ulmaceae (Hille Ris Lambers
1973c). Schizoneurata tissoti
Hille Ris Lambers Only immatures and
alate sexuparae are described, and its appearance in life is unknown. Apterae
and immatures are presumably wax-covered. Sexuparae have BL 1.6-2.2 mm and
secondary rhinaria distributed III 15-20, IV 2-5, V 1-3, VI 0-1. On Crataegus uniflora and Crataegus
sp. in Florida, USA (Hille Ris Lambers 1973c).
One
species in the Himalayan region, with empodial hairs reduced (in alatae) or
absent (in apterous exules); related to Eriosoma
but without siphunculi in the morphs on the secondary host plant, and
distinct from the nearctic genus Schizoneurata
because it has embryos with spinulose hind tarsi, as in palaearctic Eriosoma (Schizoneura) (Hille Ris Lambers 1973b). A cladistic study by Sano
& Akimoto (2011) placed it close to Aphidounguis.
[A second species described in this genus, Schizoneurella gei Bozhko, was synonymised with Eriosoma japonicum by Eastop &
Blackman (2005).] Schizoneurella indica
Hille Ris Lambers Galls are of the
open leaf-roll type, on leaves of Ulmus
villosa in India (Kashmir, Jammu) and Pakistan, and
possibly China (see below). Bhagat (1982b) reported that it formed clavate,
asymmetrical pouch galls on U.
wallichiana, but this was probably based on a misidentification (S.
Chakrabarti, cited in Sano & Akimoto 2011). Alatae (BL 1.8-2.3 mm) (fig.118B)
are produced in May-June. Host alternation occurs to Malus pumila, where colonies live, presumably in woolly wax, on
the roots. Qiao et al. (2005d)
described apterous and alate exules collected in September-October from Malus in China, and reported that
their root feeding may cause considerable economic damage. [However, to judge
from alatae and a single aptera representative of this material that have
been deposited in the BMNH collection, the identity of these aphids is
questionable. The aptera with 4-segmented antennae seems to be Aphidounguis mali; it has wax gland
plates of the form typical of that species. Its tarsi all have only one claw
fully developed, and embryos within the abdomen have tarsi with the long
pointed single claws typical of apterous exules of A. mali. The alatae (collected on sticky bands around the trunks
of the apple trees) are sexuparae of a species of Eriosomatinae, not Aphidounguis, and may be those of S. indica, although the differences
from the spring migrants from Ulmus galls
are not those usually found between these two morphs in one species, so
further confirmation seems necessary.] Bhagat (1982b) reported a parasitoid and
hyperparasitoid of the gall-feeding generations (but note that the identity
of the aphid host requires confirmation).
Two
east Asian species related to Melaphis,
and likewise having host alternation between Rhus and mosses, but alatae emerging from the galls on Rhus have an elongate, curved
pterostigma. In the light of recent molecular studies by H.C. Zhang &
Qiao (2007) and Yang et al. (2010),
as well as numerical taxonomic analysis by Yang et al. (2008), we have reinstated the genus Meitanaphis Tsai & Tang for two of the three species (elongallis and microgallis) treated under Schlechtendalia
in Blackman & Eastop (1994). Schlechtendalia chinensis
(Bell) Plate
6f Chinese Gall-nut Aphid Galls
('Chinese gall-nuts') are greenish-yellow, densely covered with short, fine
hairs, ovoid, becoming woody and irregular in shape with horn-like protrusions,
varying from 1 to 6 cm in maximum length (fig. 133).
On Rhus chinensis (= javanica, semialata) in China, Japan,
Korea, Taiwan and Malaysia. Heteroecious holocyclic, with an unusual one-year
cycle; after several generations in the gall, emigrant alatae (BL c. 1.5mm)
are produced in October and fly to found overwintering colonies on mosses (Mnium, Plagiomnium). The appearance of the overwintering immature stages
on mosses is undescribed. Alatae sexuparae return to Rhus in spring to produce sexuales. Sexual females live in bark
crevices for three weeks after mating and then each produces a single
fundatrix larva ovoviviparously – a phenomenon not known in any other aphid
(Tang 1976; Takada 1991a). The galls have been collected for many years; they
have been used in tanning leather, in the preparation of dyes and for
medicinal purposes (including a treatment for haemorrhoids; Gardener 1981).
Takada (1991a) and P. Liu et al. (2014) studied
gall development, and Shao et al. (2013) made a detailed study of
the generations developing in the gall. Qiu et al. (1999) examined the influence
of environmental factors on development of sexual females. Wang et al.(2008) and Ren et al. (2008) examined
genetic diversity in and between populations. 2n = c. 36. Schlechtendalia peitan (Tsai
& Tang) was previously treated as a synonym of S. chinensis, but evidently the galls of this species do not
develop horn-like protrusions (see Fig. 4d in C. Zhang et al. 2008), and there are differences in the relative lengths
of the antennal segments of the emigrant alatae (Yang et al. 2009). On
Rhus chinensis and R. punjabensis var. sinica in China.
Two species on woody Euphorbiaceae/Phyllanthaceae
belonging to an ancient group (also including Eonaphis and Paulianaphis)
of Gondwanaian distribution. Remaudière (1988, 1990) reviewed the genus,
synonymising most of the described species. DNA barcoding also failed to
distinguish the two currently recognised species (Q. Liu et al. 2013); further work using nuclear markers is needed. A.K.
Ghosh & Agarwala (1993) reviewed the genus in India. Schoutedenia emblica
Patel & Kulkarni Apterae are
yellow-green to green; BL 1.4-1.6 mm. In ant-attended colonies, only on Phyllanthus emblica (= Emblica officinalis) in India,
Pakistan, Nepal, Thailand and China (D. Zhang et al. 2006). It was recorded from Spiraea sp. in western Himalaya (Bhattacharya & Dey 2001, as
ssp. andhraka), but the host is
unlikely and there are errors in their morphometric data. Monoecious
holocyclic; in southern India, apterous (brachypterous?) males and oviparae
are produced in August (David & Hille Ris Lambers 1956; as ssp. andrhaka). Schoutedenia
ralumensis Rubsaamen
Plate 10e, f Apterae are lemon-yellow or green; BL
1.4-1.8 mm. Alatae have thickly black-bordered forewing veins. On young
shoots, leaves and stems of woody Euphorbiaceae/Phyllanthaceae (Breynia, Bridelia, Fluggea, Glochidion, Phyllanthus), attended by ants. Records from other plants are
probably all of vagrants. Widespread in south-east Asia; India; Africa; and
down the east coast of Australia. Monoecious and partially holocyclic;
oviparae and alate males occur concurrently with parthenogenetic generations
throughout summer and autumn. Presumably all eggs laid enter diapause, as
fundatrices occur only in spring (Hales & Carver 1976). The oviparae are
unusual in having scent glands on the hind femora. The male genitalia were described and illustrated
by Wieczorek (2012). Relations with predators and
parasitoids in Australia were studied by Hales & Carver (1976) and genetic (allozyme) variation was studied by Tomiuk
et al. (1991). 2n
= 16 (female), 14 (male) or !5* (sex unknown; Papua New Guinea, leg. J.H.
Martin); the highly anomalous spermatogenesis of this aphid was studied by
Hales (1989).
A genus for one Indian species on Corylopsis with a spinulose head but
differing from Myzus in having
weakly-developed antennal tubercles and post-siphuncular sclerites (R.C. Basu
et al. 1976). Scleromyzus
corylopsis (Basu, Ghosh & Raychaudhuri) Colour in life unrecorded, BL of apterae
1.6-2.5 mm. On unidentified Corylopsis
sp(p). in India (Shillong, Assam; R.C. Basu et al. 1973, as Myzus).
Other morphs and life cycle are unknown.
About
16 palaearctic species similar to Hyadaphis
but with very short siphunculi. They are typically heteroecious between Lonicera and Umbelliferae/Apiaceae, or
live without host alternation on one or other of these host‑plant groups.
A few species occur on other plants, including two on Impatiens. The genus is not well-known biologically and the life
cycles of many species are not completely known. Populations on Lonicera are particularly difficult to
assign to species. Accounts are available for north-west Europe (Heie 1992,
Blackman 2010), central Europe (Heinze 1960), European Russia (Shaposhnikov
1964), Siberia (Ivanoskaya 1977; Pashchenko 1988a), and Japan (Miyazaki
1971). Semiaphis aizenbergi (Narzikulov)
Apterae are pale green, wax-covered; BL c. 1.7 mm. On Lonicera spp. in Central Asia.
Oviparae and alate males occur in November (Mukhamediev & Akhmedov 1982). Semiaphis anthrisci (Kaltenbach)
Apterae are dirty green, wax-powdered; BL 1.8-2.2 mm. Recorded from
several genera of Umbelliferae/Apiaceae (Aegopodium,
Anthriscus, Peucedanum, Torilis),
slightly rolling the leaves. In north-west, northern, central and eastern
Europe, Kazakhstan (Kadyrbekov 2012a) and west Siberia (Heie 1992).
Differences from several other species (dauci,
pastinacae, pimpinellae, sphondylii) are minimal, and therefore host records
must be treated warily; possibly the true host range is more restricted.
Heikinheimo (1997) gave additional data to distinguish anthrisci from dauci and
pimpinellae on the basis of hair
lengths. Monoecious holocyclic; oviparae were collected in September on Peucedanum palustre. Stekolshchikov et al. (2008b) provided detailed
descriptions of males and oviparae. Semiaphis cervariae (Börner) (Fig.41c) Apterae are pale yellow-ochre to bright
greenish yellow, with fine wax powder; BL 1.9-2.3 mm. On Peucedanum spp., causing yellow and red spotting, and desiccating
terminal parts of leaflets. In Europe (Austria, Gemany, Netherlands,
Switzerland). Semiaphis coniumi Bozhko
Apterae are greenish with brown legs and dark green siphunculi and
cauda; BL c.1.9 mm. On Conium maculatum
in Ukraine. Semiaphis coryspermi Mamontova ex Shaposhnikov Apterae are greenish white, with waxy
bloom; BL c.1.5 mm. On Coryspermum sp.,
in leaves rolled into tubes. In Ukraine. Mamontova (1979) provided a more
detailed description. Semiaphis dauci (Fabricius) Plate 13h Apterae are pale blue-green with dark brown
head and waxy bloom; BL 1.3-2.1 mm. On wild and cultivated Daucus carota, feeding on upper sides
of rolled young leaves and leaflets in spring, and later in umbels. It is
also recorded from Aegopodium and
several other genera of Umbelliferae/Apiaceae. In Europe, North Africa (Tunisia; Boukhris-Bouhachem et al. 2007), the Middle
East and Central Asia (south-east
Kazakhstan; Kadyrbekov 2005c). Records
from USA require confirmation (Foottit et
al. 2006), although a carrot-feeding species of Semiaphis does occur there (Skvarla et al. 2017). Monoecious holocyclic (with alate males) on Daucus, or anholocyclic. Similar
aphids on Seseli spp. are currently
regarded as a subspecies, S. dauci ssp. seselii Börner. Semiaphis heraclei (Takahashi)
(Fig.10k,
Fig.38d) Apterae are pale grey-green to
yellow-green, wax-dusted, with antennae, legs, siphunculi and cauda rather
pale; BL c.2.1 mm. On stems or curled leaves of various Umbelliferae/Apiaceae
(Angelica, Apium, Bupleurum, Carum, Changium, Cnidium, Coriandrum, Cryptotaenia, Daucus,
Glehnia, Heracleum, Ligusticum, Oenanthe, Osmorhiza, Seseli,
Torilis) in east and south-east Asia, India and Pakistan, Algeria
(Laamari et al. 2013), and Hawaii.
Heteroecious holocyclic with Lonicera spp.
as primary hosts in Japan, China and east Siberia. Hyadaphis albus (Monzen), described from Angelica acutiloba in Japan, is possibly this species. 2n=8. [Leycesteria formosana is possibly an
alternative primary host in India (Pal & Khuda Bukhsh 1983), but as these
authors recorded 2n=10 they possibly had a different species.] Semiaphis horvathi Szelegiewicz
(Fig.41d) Apterae are pale brown, lightly
wax-powdered; BL 1.6-2.0 mm. Described from Peucedanum officinale, colonising shoot tips and deforming
leaves. In Hungary, also in Czech Republic (BMNH collection, leg. H.L.G.
Stroyan) and recorded from P. morisonii
and several Ferula species in Kazakhstan (Kadyrbekov 2009b,
2014d), and on P. alsaticum in
Russia (Altai; Kadyrbekov
2014g). Semiaphis iliensis Kadyrbekov
Apterae are yellow-green; BL 1.2-1.7 mm. On Lonicera iliensis in Central Asia. Semiaphis longissima Narzikulov
Apterae are green, wax-dusted; BL 1.8-2.2 mm. On Lonicera microphylla in Central Asia. Oviparae and alate males
occur in October-November (Mukhamediev & Akhmedov 1982). Semiaphis moiwaensis Takahashi
Apterae are white with pale appendages; BL c. 2mm. On stem and roots
of Impatiens noli-tangere in Japan,
and Korea (BMNH collection, leg. W.H. Paik). This is likely to be a synonym
of S. nolitangere. Semiaphis nolitangere Aizenberg
(Fig.33e) Apterae are pale yellowish green; BL
1.8-2.3 mm. On undersides of young apical leaves of Impatiens noli-tangere, which become slightly rolled, and later
on flower stalks (Heie 1995). In
Finland and northern Russia, including west Siberia. Heteroecious holocyclic
with oviparae and alate males in September in Finland, where transfers were
made to the primary host Lonicera
tatarica, causing yellowish spots on leaves (Heikinheimo, cited by Heie
1995). Semiaphis pastinacae Börner
Apterae are probably grey-green to yellow-green, with dark antennae,
legs, siphunculi and cauda; BL c.1.3 mm. In downwardly rolled leaflets of Pastinaca
sativa in central Europe, (Austria, Czech Republic, Switzerland). Host
alternation probably occurs with Lonicera spp. as primary hosts, but
this needs confirmation (Heinze 1960). Semiaphis pimpinellae (Kaltenbach)
Apterae are dull green mottled with darker green, coated with greyish
wax powder; BL 1.4-1.8 mm. On Pimpinella
spp., esp. saxifraga, causing
leaves to curl upwards, and later in season on umbels. Ossiannilsson (1959) recorded it from Aegopodium podagraria. In Europe (not Iberian peninsula), and
Iran. It is very similar to other species in the S. anthrisci group. Semiaphis rabotkinae Pashchenko
Apterae are green, wax-dusted; BL c. 2.2 mm. On Lonicera edulis in east Siberia. Semiaphis sphondylii (Koch) (Fig.10j) Apterae are greenish white, wax-powdered;
BL 2.0-2.5 mm. In rolled leaves on Lonicera
nigra in spring, heteroecious holocyclic, migrating to various
Umbelliferae/Apiaceae (Angelica, Heracleum, Peucedanum), although some host records may be due to confusion
with other similar species such as anthrisci
and pimpinellae. In north-west,
northern and central Europe (Finland, Latvia, Netherlands, Germany, Austria),
and also reported to occur in Kazakhstan on Lonicera tatarica, Bupleurum
and Libanotis (Kadyrbekov
2009a,b).
A
genus for one Aphis-like species in
south-west Europe completely lacking siphunculi. It has dark legs and antennal
sensoriation characteristic of a group of Aphis
(incl. jacobeae in Europe and lugentis in North America) that feed on Senecio, which suggests that, despite lacking siphunculi, its
separate generic status may not be warranted. Seneciobium balachowskyi Remaudière
Apterae are greenish to black, with a waxy bloom, and appendages all
dark, including cauda; BL 1.8-2.2 mm. On stems and flowers of Senecio adonidifolius in the eastern
Pyrenees. Monoecious holocyclic with apterous males.
Four species on Nothofagus in Papua New Guinea, Australia and New Zealand, and
one on Melaleuca in Western
Australia, where Nothofagus occurs
only in fossil form. Sensoriaphis is
closely related to Taiwanaphis, and
regarded by some authors as a subgenus or even a synonym of that genus, but
we follow Remaudière & Remaudière (1997) in giving it separate generic
status, as it forms a distinct group with biological as well as morphological
differences. Carver (1978) provided a key to species, and Quednau (2010)
reviewed the genus (as a subgenus of Taiwanaphis)
and provided illustrations of all available morphs. Sensoriaphis furcifera
Carver & White (fig.
62A) Alatae have a
pale green abdomen, head and pronotum with two longitudinal dark stripes, and
mainly black antennae and legs; BL 1.8-2.8 mm. On tender young shoots and
suckers of Nothofagus moorei in New
South Wales, Australia. Carver & Hales (1974) described all morphs and
gave life cycle details; monoecious holocyclic, with oviparae and alate males
occurring concurrently with alate viviparae through spring, summer and
autumn. Eggs are flat, colourless and wax-covered, and are laid on leaves. Sensoriaphis melaleucica (Quednau) Colour of apterae in life unknown; BL
1.4-2.2 mm. On Melaleuca spp. (lanceolata, teretifolia) in Western Australia (Quednau 2010). The life cycle
is unknown. Sensoriaphis
niuginii Carver (fig.
62B) Both apterae
and alatae occur, colour in life not recorded; BL of aptera 1.5-2.2 mm, of
alata 2.0-2.4 mm. On undersides of young leaves of Nothofagus carrii in Papua New Guinea (Carver 1978). The life
cycle is unknown. Sensoriaphis
nothofagi Cottier (fig.
62D) Apterae are
yellowish-green to brownish; BL 1.1-1.5 mm. Well camouflaged, feeding on
young stems of Nothofagus spp., generally
on trees close to forest edges. Indigenous to New Zealand, where it is rather
common and widespread (Teulon et al.
2013). Alatae become mature on the undersides of leaves. Monoecious
holocyclic, with oviparae and alate males in September to February (BMNH
collection, leg. V.F. Eastop). 2n = 10. Sensoriaphis
tasmaniae Carver & Martyn (fig. 62C) Apterae have a dirty yellow
head and prothorax, rest of body olive green and rather shiny; BL 1.5-2.0 mm.
On stems of young shoots of Nothofagus
cunninghami in Tasmania (Carver & Martyn 1962). The alatae morph was
described by Carver & Martyn (1965). The life cycle is unknown.
One
Asian oak-feeding species, related to Hoplochaetaphis
but with spiculose dorsal hairs and clusters of spinal papillae on the
thorax. Quednau (1999) provided illustrations of all available morphs. Serratocallis takahashii Quednau
& Chakrabarti Apterae are brown;
BL 1.6-2.3 mm. Alatae have a short broad pterostigma with a black spot at its
base. On twigs, leaf petioles and undersides of leaves of Quercus spp. (baloot, dealbata, oblongata,
semicarpifolia) in Afghanistan, Pakistan (Naumann-Etienne &
Remaudière (1995) and India (Himachal Pradesh, Uttar Pradesh; A.K. Ghosh
& Quednau, 1990). Sexual morphs are unknown.
One
North American species on Acer,
related to Drepanaphis and
considered as a subgenus by some authors; the differences were discussed by
Smith & Dillery (1968), and the case for regarding it as a separate genus
was convincingly argued by Wieczorek et
al. (2013c). Shenahweum minutum
(Davis) (fig. 22D)
All
viviparae are alate, with brown or black thorax and very pale greenish-yellow
to whitish abdomen; BL 1.0-1.25 mm. Immatures are pale greenish-yellow. On
the undersides of leaves of Acer
saccharum in eastern North America. Oviparae and alate males collected in
Maine and North Carolina in November were described by Wieczorek et al. (2013c).
One east
Asian species with greatly reduced tarsi, host-alternating between Viburnum and ferns. There is a rather
strange parallel with a fern-feeding North American species in another
monotypic genus, Mastopoda pteridis,
with similarly vestigeal tarsi and apparently with similar biology, but not
closely related. Shinjia orientalis (Mordvilko) Plate 22c (Fig.43a) Apterae are shining yellow with black
antennae and dark tips to siphunculi; BL 1.3-2.0 mm. Alatae have paired brown
patches partially fused across midline. On young growth of Viburnum spp. in spring. Heteroecious holocyclic, migrating to ferns (Athyrium, Eriosorus, Polypodium, Pteridium, Pteris) in Japan (Sorin 1962b, as pteridifoliae). Recorded from east India, Nepal, China, Korea,
east Siberia, Japan, Philippines and Australia. It is probably anholocyclic
on ferns in Australia (Eastop 1966). 2n=12*.
Seven Asiatic species associated with Celtis and Pteroceltis. The Indian species were reviewed by A.K. Ghosh &
Quednau (1990) and the Chinese species by W. Zhang et al. (1994) and Qiao et
al. (2005b). Quednau (2003) reviewed the genus and provided a key to
species and illustrations of all available morphs. Shivaphis
catalpinari Quednau & Remaudiere Apterae are green, with long wax filaments
(W. Zhang et al. 1994, as S. similicelti); BL of aptera 1.4-2.1 mm,
of alata 1.5-1.9 mm. On Celtis
australis in Turkey, and on Celtis
koraiensis and C. sinensis in
China (Quednau 2003, Qiao et al.
2005b). Oviparae and alate males occur in Turkey in October (Quednau &
Remaudiere 1985). Shivaphis
celti Das Plate l0c, d Apterae are pale to dusky greenish,
sometimes pinkish, covered in bluish-white flocculent wax; BL 1.9-2.6 mm. Alatae
also wax-covered, with forewing veins thickly bordered with fuscous distally
and large black patches on pterostigma and distal part of Cu1b. On
undersides of leaves or on shoot tips of Celtis
spp. in Turkey, India, Pakistan, Nepal, Sri Lanka, Korea, Japan, Loochoo,
China, Taiwan and Philippines. S. celti was introduced into
south-eastern USA in the late 1990s (Halbert & Choate 2003) and had
spread to California by 2002. Das (1918) gave a full account of its biology
and natural enemies in Pakistan; oviparae and alate males occurred in
December, eggs were laid in January and hatched in March. In Japan, Shibata
(1955; as S. celticolens) obtained
sexuales in September-November. In Taiwan, S. celti is mainly
anholocyclic, but produces some sexuales in December-January. 2n = 10. Shivaphis (Sinishivaphis)
celticola
(Nevsky) All viviparae are alate, pale
green to greenish-yellow covered with bluish wax; BL 1.8-2.2 mm. Forewings
are marked with fuscous as in S. celti,
except that there is a band joining the first fork of the media and Cu1a
(A.K. Ghosh & Quednau 1990). On the undersides of leaves of Celtis caucasica (= australis), recorded from
Tadzhikistan, Uzbekistan, Kazakhstan, Pakistan and India (Kashmir). Oviparae
and alate males occur in October-November; Narzikulov (1961a) gave a detailed
account of the life cycle in Tadzhikistan, with descriptions of all morphs. Shivaphis
(Sinishivaphis) hangzhouensis (Zhang & Zhong) Only alate viviparae are known; yellow,
covered with wax, with very thick bands of fuscous along forewing veins, BL
1.9-2.3 mm. Described from Osmanthus
asiaticus in China (Zhang & Zhong, 1982c), but subsequently found on
the undersides of leaves of more likely hosts, Celtis julianae and C.
sinensis (Quednau 2003). Quednau illustrated an ovipara collected on C. sinensis in October. 2n = 10
(Blackman & Eastop 1994). Shivaphis (Sinishivaphis)
pteroceltis Jiang, An, Li &
Qiao Apterae are red-brown in life,
covered in thick powdery or filamentous white wax; BL 1.5-1.8 mm. In large
colonies on leaves of Pteroceltis
tatarinowii in China (Shandong and Anhui provinces); colonies with
powdery wax are attended by ants, but those with wax filaments are
unattended. Fundatrices hatch in March, and oviparae and alate males occur in
October. Jiang et al. (2014)
provided descriptions of all morphs. Shivaphis (Sinishivaphis)
szelegiewiczi Quednau Only alate viviparae are known; colour in
life not recorded, presumably covered in wax, BL c. 2.5 mm. Forewings are
marked with fuscous as in S. celti,
but with a spot below the pterostigma, next to base of media. On Celtis aurantiaca in Korea (Quednau,
1979), and on C. biondii in Hong
Kong. An immature ovipara collected in Hong Kong in December is illustrated
by Quednau (2003). Shivaphis
(Sinishivaphis) tilisucta Zhang Only alate viviparae are known; bright
green covered with thick wax powder, BL c. 1.9mm. Described from Tilia tuan in China (Zhang & Zhong
1990), but the host was possibly misidentified and it has subsequently been
collected from Celtis sinensis in
China, and from C. japonica and C. jessoensis in Korea. Quednau (2003)
illustrated an ovipara collected on C.
jessoensis in Korea in early November.
One species in the Mediterranean region on Quercus, the apterae of which have
remarkable variation in form of dorsal hairs. Siculaphis vittoriensis
Quednau & Barbagallo Apterae are
pale yellowish green with longitudinal olive green dorsal stripes leaving a
pale spinal line; BL 1.0-1.5 mm. Alatae have smoky wings with Cu1b
strongly bordered with fuscous. On Quercus
calliprinos and Q. ilex,
forming ant-attended colonies mainly along mid-ribs on basal parts of
undersides of leaves. In Italy; described from Sicily, also found in Puglia,
Tuscany and Sardinia, and also now recorded from Tunisia (Ben Halima-Kamel
2012). A holocycle occurs in Puglia; the ovipara and apterous male were
described from Q. ilex by Patti
& Barbagallo (1997). Populations in Sicily were suspected to be
anholocyclic (Quednau & Barbagallo 1991).
One
species on Ulmus in east Asia
perhaps related to Pseudochromaphis,
the apterae having extensive dark dorsal sclerotisation. Quednau (2003)
reviewed the genus and decided that it comprised one species with vary
variable pigmentation. Sinochaitophorus maoi
Takahashi (= mordvilkoi Quednau
& Shaposhnikov) Colour of aptera is not recorded, probably blackish; BL
c. 1.3 mm. Alatae are blackish-brown, mottled with red, with maculate wings
(Holman & Szelegiewicz 1974a). On young shoots and along mid-ribs on the
undersides of the leaves of Ulmus
japonica in east Siberia (Quednau & Shaposhnikov 1988, as mordvilkoi), and on U. pumila in Mongolia and China. It is
attended by ants. Quednau (2003) illustrated oviparae collected in China and
east Siberia in September-October.
Two
species described from Taiwan and one from Bhutan, distinguished in
particular by the very numerous protuberant secondary rhinaria on the
antennae of alatae (Hille Ris Lambers 1956d). One of the Taiwanese species is
only known from a trapped alata. Sinolachnus
elaeagnensis Chakrabarti & Das Apterae are undescribed, alatae have BL
3.7-4.0 mm. Collected (with immatures) from young stems of an Elaeagnus sp. in Bhutan (Chakrabarti
& Das 2014). Attended by black ants. Sexual morphs and life cycle are
unknown. Sinolachnus niitakayamensis
(Takahashi) Apterae are
yellowish-brown, blackish on dorsal abdomen; BL c. 2.5mm. On branches of Elaeagnus oldhami in Taiwan (Takahashi
1927a). Sexual morphs and life cycle are unknown.
Eight species with head ventrally spinulose and
with weakly clavate, calf-shaped siphunculi and a long cauda, living on
various trees and shrubs in east and south-east Asia. Accounts are available
for Japan (Miyazaki 1971), China (Tao 1963), north-east India (Raychaudhuri
1980) and Java (Noordam 1986). Sinomegoura
citricola (van der Goot)
Plate 24i Apterae are broadly
spindle-shaped, shiny dark brown to reddish-brown with bright red eyes,
siphunculi pale at bases and dark distally, and a long dark cauda; BL 1.4-2.7
mm. On undersides of leaves or young growth of numerous tropical shrubs and
some trees, in about 20 families, the most frequent records being from Lauraceae
and Rutaceae.
Distributed throughout east and south-east Asia (India, Sri Lanka, China, Thailand, Korea, Taiwan,
Malaysia, Indonesia, the Philippines); also in
Australia (Queensland), and recently found in southern California (Halbert
2009). No sexual morphs are known, and it is apparently mainly or entirely
anholocyclic. Various karyotypes have been reported, and the variation in
chromosome number warrants further investigation; 2n = 12 (Kulkarni 1984,
Dutta & Gautam 1993), 2n = 16 (Chen & Zhang 1985b) and 2n = 18 (Kar
& Khuda-Bukhsh 1986). Sinomegoura coffeae Noordam
Apterae are broadly spindle-shaped, shining yellow with black antennae
and tibiae, siphunculi browner towards
apices, and cauda yellow or slightly grey; BL 2.4-3.0 mm. On undersides of
leaves of Coffea sp. in Java and Timor (Indonesia). Coffea is not indigenous to
south-east Asia and there is presumably a native host (see Noordam 1986). Sinomegoura nepalensis
Das & Raychaudhuri Apterae are
yellowish, with siphunculi basally pale and distally dark as in citricola, but with a pale cauda; BL
2.8-2.9 mm. On young shoots of Melia
sp. in Nepal (B.C. Das & Raychaudhuri 1983). The life cycle is unknown. Sinomegoura
photiniae (Takahashi)
Apterae are green, often with a pinkish tinge (Japan, Takahashi 1960c)
or 'faded orange' (China, Takahashi 1936a), with black siphunculi and green
or yellowish cauda; BL 2.0-2.8 mm. On Photinia
spp. and Rhaphiolepis spp. (Miyazaki 1971), and also recorded
from Phoebe sp. (Tao 1999). In
China, Taiwan, Japan and India. Sexual morphs and life cycle are unknown. 2n
= 18 (Khuda-Bukhsh & Kar 1990). Sinomegoura
pyri
Ghosh & Raychaudhuri Apterae are
pale yellowish, with pale siphunculi and cauda; BL c. 2.4 mm. On undersides
of young leaves of Pyrus sp. in
Assam, India (A.K. Ghosh & Raychaudhuri 1968b). Ardisia sp. is also recorded as a host plant (Raychaudhuri et al. 1980b). Other morphs and life
cycle are unknown. Sinomegoura rhododendri (Takahashi)
Apterae are green, with dark-tipped siphunculi and black cauda; BL
c.1.4 mm. On Rhododendron oldhami in
Taiwan. Records from other hosts in
Philippines and India are likely to be referable to other species. Sinomegoura symplocois (van der Goot)
Apterae are pale reddish brown with eyes, antennae greyish white with
apices of segments black, legs pale brown with apices of tibiae and tarsi
black, siphunculi pale brownish yellow
with dark apices, cauda black; BL c.2.9-3.0 mm. On Eurya japonica and Symplocos
spp. (cochinchinensis, sessilifolia). In Java (van
der Goot 1917, Noordam 1986) and Malaya (BMNH collection, leg. R. Takahashi). The
life cycle is unknown. [The record from Acronychia laurifolia in north-west India (Shillong; A.K. Ghosh
& Raychaudhuri 1970) seems to be another, apparently undescribed,
species, to judge from two alatae (leg. A.K. Ghosh) with the same date, host
and locality data in the BMNH collection.]
Three
or four species in east Asia in which the dorsum of the aleyrodiform aptera
is densely covered with spine-like processes. Sinonipponaphis formosana
(Takahashi) Apterae are
blackish-brown, with a small yellowish-brown patch on 'margin near caudal
apex' (Takahashi 1927b); BL c. 1.7mm. Alatae are black with brownish abdomen,
with secondary rhinaria distributed ANT III 32-36, IV 18-20, V 10-13. Apterae
are fixed firmly to branches of Persea (=
Machilus) sp. (Takahashi 1929) in
Taiwan, China (Szechuan, Tao 1966) and Thailand. In Taiwan it is also
recorded from Phoebe sp. (Tao
1999). The life cycle is unknown. [The aphid described under the name Sinonipponaphis monzeni from Machilus sp. in Meghalaya, India (A.K.
Ghosh 1988) may be this or a related species.] Sinonipponaphis
hispida Noordam
Apterae dull black or brownish-black with reddish-brown cauda, without
wax; BL 0.8-1.0mm. Living scattered on undersides of leaves of Lithocarpus indutus in Java (Noordam
1991). Alatae and life cycle are unknown. Sinonipponaphis holboelliae (A.K. Ghosh & Raychaudhuri) Apterae are brownish, covered with mealy
wax; BL 1.7-2.1 mm. Heavily infesting stems of Holboellia latifolia in West Bengal. Other morphs are unknown.
Eleven
species of spiny-haired oval grass-feeding aphids with stump-shaped
siphunculi, 5-segmented antennae and a sclerotic tergum. The four species of Sipha s.str. have a knobbed cauda; two
are nearctic with a smooth tergum and two palaearctic with a spinulose tergum.
The eight species of subgenus Rungsia have
a broadly rounded cauda and a smooth tergum. Regional accounts are available
for UK (Stroyan 1977), north-west Europe (Heie 1982), the Iberian peninsula
(Nieto-Nafría & Mier Durante 1998), European Russia (Shaposhnikov 1964),
North America (Halbert et al. 2013b) and USA Rocky Mountain region (Palmer 1952).
Wieczorek (2010) fully reviewed the genus with distribution maps and
illustrations, host lists and keys to all available morphs. Sipha agropyronensis (Gillette) Apterae are long-bodied, rusty yellow
with brown pleural stripes; BL 1.6-2.2 mm. On upper surfaces of leaves of Elymus glaucus (as Agropyron glaucum), attended by ants
(Gillette 1911, as Chaitophorus).
Also collected on Elymus smithii (as
Agropyron occidentale; BMNH
collection, leg. M.A. Palmer) and E.
hispidus (Wieczorek 2010). USA; Colorado, with records also from
California and Montana (see Halbert et
al. 2013b),
and Gillette mentions collections “throughout the Northwest”. Monoecious
holocyclic, with oviparae in October (Palmer 1952). Sipha (Rungsia) arenarii Mordvilko Apterae are greenish yellow to brown with
pale longitudinal stripes; BL 2.0-2.4 mm. The usual host is Leymus arenarius, but it is also recorded from Elymus
hispidus, E. repens and Avena sterilis ssp. ludoviciana. In Europe, and eastward to west Siberia, Kazakhstan and Inner
Mongolia. A record from Turkey (on Triticum
aestivum; Akyürek et al. 2011) requires
confirmation. Monoecious
holocyclic, with oviparae and apterous males in October. The sexual morphs
were redescribed by Wieczorek (2010). Sipha (Rungsia) burakowskii Holman &
Szelegiewicz Apterae are greyish
green with brownish spots, very slightly pruinose; BL 1.8-2.3 mm. In small
colonies on upper sides of leaves of Leymus
chinensis (= Aneurolepidium pseudoagropyron) in Mongolia, and also recorded from Elymus repens in Kazakhstan (Kadyrbekov 2002e). Wieczorek (2010)
provided a redescription of the aptera. Alatae and sexual morphs are unknown.
Sipha (Rungsia) elegans del Guercio (=
aegilopis Bozhko; synonymy by Wieczorek 2010) Apterae are yellowish brown to brown with
a paler spinal stripe; BL 1.4-2.1 mm. Alatae have black dorsal abdominal
markings. In small colonies on upper sides of leaves of various grasses and
cereals (Aegilops, Agropyron, Agrostis, Ammophila, Arrhenatherum, Bromus, Elymus, Festuca, Hordeum, Phleum, Puccinellia, Setaria, Triticum)
often causing leaves to roll upwards and develop yellow patches. Havličkova et al. (1996) recorded its
differential colonisation of Aegilops
and wild Triticum species in Czech
Republic. In
Europe, across Asia to China and east Siberia, and in North America (see
Wieczorek 2010 for full details of distribution). Monoecious holocyclic with slender brown-black
apterous males, in both Europe and North America. See also Blackman &
Eastop (2000). 2n=6. Sipha flava (Forbes) Yellow Sugarcane Aphid Plate 5d Apterae are straw to bright yellow (or
green at low temperatures) with dusky transverse intersegmental markings; BL
1.3-2.0 mm. Alatae have variably-developed dark dorsal abdominal markings. On
leaf blades of grasses in numerous genera including Andropogon,
Avena, Cenchrus, Cymbopogon, Cynodon, Digitaria, Holcus, Hordeum, Miscanthus, Panicum, Paspalum,
Pennisetum, Saccharum, Setaria, Sorghum and Triticum, often forming large colonies, not attended by ants. Bradshaw et al. (2010) found large populations
on young stands of the bioenergy crop Miscanthus
×giganteus in USA. Life table parameters of S. flava on this and another bioenergy crop were measured by
Pallipparambil et al. (2014). It is also recorded from
Cyperaceae (Carex, Cyperus). In USA (including Hawaii),
Central and South America, Madeira, the Azores, and recently found in Morocco
(Abdelmajid 2008), Spain (Hernández-Castellano & Pérez-Hidalgo 2014) and
South Africa (Way et al. 2014).
Monoecious holocyclic with apterous
males in areas with cold winters, anholocyclic in warmer climates. See also
Blackman & Eastop (2000). Sipha glyceriae (Kaltenbach) Apterae are dull
pale green to mid-green, sometimes reddish, often with a paler spinal stripe;
BL 1.5-2.4 mm. On many species of Graminae, especially in wetlands; genera
include Agrostis, Aira, Alopecurus, Glyceria, Elymus, Hydrochloa, Oryza, Paspalum, Phalaris, Poa, Puccinellia. Also collected from
Ceratophyllaceae (Ceratophyllum), Cyperaceae (Carex, Cyperus, Eleocharis, Schoenus, Scirpus),
Juncaceae (Juncus), Typhaceae (Typha)
and Alismataceae (Sagittaria). In
Europe, Middle East, eastward to west Siberia and Central Asia, and in USA
and Canada (see Wieczorek 2010 for full details of distribution). Monoecious
holocyclic, with dark apterous males. 2n=12 (Gut 1976) or 10. Sipha littoralis (Walker)
Apterae are green, dark green or bluish green with brownish markings,
head and thorax more brownish; BL 1.4-2.2 mm. In leaf sheaths of grasses in
coastal situations (e.g. Spartina spp.,
Puccinellia maritima, Festuca rubra),
where they are able to survive tidal submergence. In north-west and northern
Europe. Monoecious holocyclic with brown oviparae and apterous males in
October (Heie 1982), or as early as August in northern Russia (Wieczorek
2010). Sipha (Rungsia) maydis Passerini Plate 5e Apterae are shining dark brown to almost black
on dorsal surface; BL 1.0-2.1 mm. Alatae have a solid black dorsal abdominal
patch. On numerous species of Poaceae, in more than 30 genera. It
feeds on upper sides of leaf blades near bases, sometimes on stems or
inflorescences, often attended by ants. In Europe, Mediterranean region,
Middle East, Central Asia, India, Pakistan, South Africa, and now in
Argentina (Corrales et al. 2007)
and USA (Sorensen 2007, Halbert et al.
2013b) (for full details of
distribution see Wieczorek 2010). Monoecious holocyclic on Poaceae with
apterous males recorded from Turkey (Tuatay & Remaudière (1965) and
Poland (Wieczorek 2010), but probably anholocyclic in most countries. The
male genitalia were described and illustrated by Wieczorek (2011). In drier
climates outside north-west Europe it can be a pest of cereal crops (Blackman
& Eastop 2000). 2n=12. Sipha (Rungsia) praecocis (Bozhko) Apterae are yellowish, mottled with
green; BL c.1.6 mm. In small colonies on upper sides of leaf blades of Carex preacox in Ukraine, and also
recorded from Kazakhstan. Alatae and
sexual morphs are unknown. Sipha (Rungsia) taurica (Mamontova) Apterae are lemon yellow; BL 1.8-2.2 mm.
On undersides of leaves of Leymus
racemosus, turning them yellow. In Ukraine (Crimea), Russia (Volgograd;
Kadyrbekov 2014g) and
also recorded from Kazakhstan. Alatae and sexual morphs are unknown. Sipha (Rungsia) uvarovi Mordvilko Apterae are green to dark green; BL
c.1.5-1.6 mm. On wheat (Triticum) and various grasses (Agropyron, Festuca). In Ukraine, southern Russia, west Siberia, and
Kazakhstan. Alatae and sexual morphs are unknown.
One
species which is a specialized feeder on Cupressaceae in North and Central
America (Remaudière & Munoz-Viveros 1991). Siphonatrophia cupressi
(Swain) Apterae are pale green
(concolorous with leaves of host), with body strongly domed dorsally and
flattened against leaf ventrally; BL 1.0-1.7 mm. Alatae have the forewing
media once- or twice-branched. On Cupressaceae (Juniperus, Cupressus, Widdringtonia) in western USA,
Mexico and Central America (Honduras,
Costa Rica), and recently introduced to southern Europe (France, Rabasse et al. 2005a; Italy, Bella &
Squarcini 2009; Spain, Lumbierres et al.
2015) and Japan (Sano & Yoshitomi 2012).
In the south of France and Spain A significant amount of parasitization by Lysiphlebus testaceipes has been
observed in southern France and Spain (Rabasse et al. 2005a;
Lumbierres et al. 2015).
Monoecious holocyclic; oviparae and both alate and apterous males occur in
October-November (Palmer 1952).
About
66 species resembling Macrosiphum in
the reticulation of their siphunculi, but with shorter hairs, less well‑developed
antennal tubercles, and often with a more sclerotic dorsum. The
genus is hard to define; the karyotype (2n = 16 or 2n == 18) suggests that
the strong similarity to Macrosiphum
(which have 2n = 10) may be due to morphological convergence. Probably they have a
closer relationship with Metopolophium,
which also have 2n=16 or 18 and have a similar biology. Sitobion have a broad range of host plant
relationships, with
species about equally distributed between monocots (particularly Poaceae) and
dicots. Several species host-alternate
between dicots (Rubus, Rosa, Akebia) and Poaceae, and some of the other dicot‑feeding
species may prove to be the primary host forms of species on grasses. Sitobion are
distributed throughout the northern hemisphere, and have also undergone an
adaptive radiation in Africa south of the Sahara. The four European species are all associated with
Poaceae. The 24 African species include 9 grass‑feeders and 13 from
dicots. The 30 Asiatic species include only 6 from Poaceae, but 10 from orchids
and other monocots and 13 from dicots.
Twelve American and one European species previously in Sitobion were placed in Macrosiphum after a cladistic analysis
by Jensen (1997). Five grass-feeding
species with characters intermediate between Sitobion and Metopolophium are
placed in subgenus Metobion; a key
to these species was provided by Stekolshchikov (2011). Accounts are
available for Europe (Holman 1961c, added to by Stroyan 1969b), UK and
Ireland (Blackman 2010), Fennoscandia and Denmark (Heie 1994), east Africa
(Eastop 1958), west Africa (Eastop 1961a), Africa south of the Sahara
(Remaudière & Autrique 1985), Japan (Miyazaki 1971, in Macrosiphum), China (Tao 1963), India
(David 1976), north-east India (Raychaudhuri, 1980, in Macrosiphum) and east Siberia (Pashchenko 1988a). Many of the
African species are parasitised by Aphidius
camerunensis (see Remaudière & Autrique 1985). Sunnucks et al. (1996) and Hales et al. (2010) discussed the introduced species in
Australia. Sitobion africanum (Hille Ris Lambers) Plate 26d Apterae are variably coloured yellowish
green to green, reddish, yellowish, or greenish brown, reddish or brownish
forms being usually densely wax‑covered; BL 1.6‑3.1 mm. There is
also a very variably developed dorsal abdominal pattern of dark pigmentation;
there may be an extensive, broken pattern of sclerotization of the whole
dorsal surface or, at the other extreme, the cuticle of the paler green forms
may be almost completely pale. Alatae have a pattern of transverse, narrow,
often interrupted dark bars on the dorsal abdomen. On Poaceae (Aristida, Bothriochloa, Brachyaria, Bromus, Calamagrostis, Cenchrus, Cynodon, Dichanthium, Digitaria, Eragrostis, Exotheca, Hyparrhenia, Panicum, Pennisetum, Setaria, Sorghum, Sporobolus, Stenotaphrum, Tricholaena, Triticum, Zea)
in Africa, Indian Ocean islands, and also recorded from Yemen. A record from
Chile was based on a misidentification (E. Fuentes-Contreras, pers. comm.).
In Africa S. africanum is recorded frequently on various other
monocots (Asparagus, Cyperus, Gladiolus, Watsonia)
and also on dicots (Aeschynomene, Aster, Clematis, Desmodium, Droogmansia, Euphorbia, Hakea, Hypericum, Lactuca, Lefeburea, Pavetta, Tephrosia, Thesium, Ximenia), including
some trees (e.g. Dombeya, Ficus, Phyllanthus). Euphorbiaceae-feeders
identified as S. africanum have
lateral abdominal tubercles; possibly more than one species is being
identified as africanum, and it may eventually transpire
that this name is being applied to a complex of species. Müller & Schöll
(1958) gave an account of the variation and host plants of S. africanum in South Africa. The
life cycle is unclear; it is probably mainly anholocyclic, but Müller & Schöll (1958) described
oviparae from Ficus. Sitobion akebiae (Shinji) Apterae are variably coloured green,
yellowish green or orange red, with dorsal abdomen pale or variably
pigmented. Heteroecious holocyclic between Akebia and secondary hosts mainly in Poaceae according to Moritsu
(1944, as granarium) and Takahashi
(1964), but Miyazaki (1971) found fundatrices on Stellaria and Platanus,
so overwintering eggs are apparently laid on various plants. Summer host
plants include Elymus, Festuca, Oryza, Poa, and Triticum; also Gladiolus, Plantago, Ranunculus and Trifolium. In Japan and Korea. Kanehira et al. (1988) studied population
dynamics of aphids identified as S.
akebiae in wheat fields in Japan. The more widely distributed S. miscanthi is very difficult to separate
from S. akebiae and may perhaps be
the same species. For further information see Yano et al. (1983). Sitobion alopecuri (Takahashi)
(= Sitobion salicornii Richards;
synonymy by Jensen, 1997) Apterae are
yellowish brown to mid apple-green with pale to dusky siphunculi, darker at
apices; BL 1.8-3.2 mm. On grasses, particularly Alopecurus, but also in
sandy habitats on Ammophila, Elymus, Hordeum and Poa. Asia
(Taiwan, north India), Europe (Netherlands, UK, Poland) and North America
(California, Hille Ris Lambers 1966a, as ssp. sylvesteri; Oregon, A. Jensen, aphidtrek.org;
British Columbia, Richards 1963, as salicornii
– vagrants?; Manitoba, BMNH collection, leg. A.G. Robinson). Monoecious
holocyclic on Alopecurus, with
oviparae and alate males produced
in October (Stroyan 1991, and BMNH collection). It is characteristic of the
European and North American populations to have marginal abdominal tubercles
on all or most of abdominal tergites 2-5, whereas these are not developed in
the single oriental specimen seen (from India). S. pseudoalupecuri Chakrabarti, described from an unidentified
grass in Himachal Pradesh, India, is
very similar but has a shorter, blunter cauda. 2n=18 (sample from British
Columbia). Sitobion anselliae (Hall)
Apterae are grass green to yellow green with black siphunculi; BL
1.4-2.2 mm. Alatae have no dark dorsal abdominal markings. On wild and cultivated
orchids (Ansellia, Cattleya, Epidendrum, Eulophia). Common on Eulophia streptopetala in Burundi
(Remaudière & Autrique 1985). In Africa south of the Sahara (Angola,
Burundi, Kenya, Rwanda, South Africa and Zimbabwe). Colonies are ant-attended
(Pheidole sp.; van Harten &
Ilharco 1976). Sitobion asirum Aldryhim & Ilharco
Apterae are pale green; BL 1.7-2.0 mm. In small colonies on upper
sides of leaves of Phragmanthera
regularis (a parasite on Acacia) in Saudi Arabia. Closely related to
an African species on Loranthaceae, S.
loranthi. Sitobion aulacorthoides David, Narayanan & Rajasingh Apterae are green, with dark brown apices
to femora and tibiae, and siphunculi also dark brown on distal half; BL
1.8-2.4 mm. Described from undersides of leaves of an unidentified plant (a
“weed”) in NW India (Himachal Pradesh). There are subsequent records of this
species from various plants in north-west and north-east India, mainly
Lamiaceae (Ajuga, Elsholtzia, Perilla, Plectranthus),
but also from species of Commelina,
Rubus and Valeriana, which are
presumably not normal host plants. Biology is unknown. 2n=18. Sitobion autriquei Remaudière Apterae are shining brown-black, with
black SIPH, and cauda pale except for dusky tip; BL 2.0-2.7 mm. On various Compositae
(Conyza, Crassocephalum, Helichrysum),
and also collected from species of Plectranthus (Lamiaceae) and Laportea (Urticaceae). In Africa; as
yet known only from Burundi (Remaudière & Autrique 1985). The life cycle
is unknown. Sitobion avenae (Fabricius) Grain Aphid Plate 26e Apterae are yellowish green or dirty
reddish brown, sometimes rather shiny, with black antennae and siphunculi (see
influentialpoints.com/Gallery); BL 1.3‑3.3 mm. On numerous species of
Poaceae, including all the cereals and pasture grasses of temperate climates,
and many other monocots. Certain dicots may also occasionally be colonised,
e.g. Capsella bursa-pastoris and Spergula arvensis. On cereals the aphids prefer to feed on the
upper leaves, and on the ears once these have emerged. Throughout Europe, the
Mediterranean, the Middle East, Central Asia, India, Nepal, Pakistan, Africa;
North, Central and South America (oriental records refer to S. miscanthi). Monoecious holocyclic,
with alate males, on many species of Poaceae (Müller 1977a). Anholocyclic
overwintering is common in regions with mild winters. Rautapää (1970) studied
preferences among a range of host plants. DNA studies in southern
England (Sunnucks et al. 1997) and
France (Haack et al. 2000) have
revealed host specialisation including races preferentially colonising wheat
and Dactylis glomerata, the latter
form apparently hybridising naturally with S. fragariae. Economic
implications of limited genetic exchange between populations on different
hosts were discussed by Vialatte et al.
(2005). The genetics of introduced populations in Chile was studied by
Figueroa et al. (2005). [S.
avenae, S. miscanthi and S. akebiae form a complex of closely
related species. Choe et al. (2006) compared variation in morphology and in an
mtDNA sequence of S. avenae from
England and North America with the oriental taxa S. miscanthi and S. akebiae,
and concluded that the latter two should be treated as synonyms of S. avenae. As a result in some recent
papers cereal-feeding populations of this complex in east Asia have been
identified as S. avenae (e.g. Xu et al. 2011; Xin et al. 2014; B. Zhang et
al. 2016). However we agree with Hales et al. (2010) that there is insufficient evidence for such a
step, and a more thorough study of oriental Sitobion is needed. It is possible that there is a complex
situation in China involving both introduced S. avenae and indigenous S.
miscanthi, which could explain the high levels of genetic diversity
revealed in a recent study using microsatellite markers (Xin et al. 2014), and may also need to be
taken into account in studies of secondary symbionts (Alkhedir et al. 2015). The complete
mitochondrial DNA of a sample from China identified as S. avenae has been published (B. Zhang et al. 2016).] See also Blackman & Eastop (2000, 2007).
2n=18. Sitobion
bambusicola (L.K. Ghosh) Apterae are yellowish; BL c. 3.7mm. In a
colony under young, tender leaves of an unidentified Bambusa sp. in Himachal Pradesh, India (L.K. Ghosh 1986). Other
morphs and life cycle are unknown. Sitobion bamendae (Eastop)
Apterae are green with dark dorsal abdominal cross-bands, pinkish
green or olive-green to almost black; BL 1.7-2.5 mm. On grasses (Pennisetum, Setaria) in Africa (Burundi, Cameroon). Sitobion (Metobion) beiquei (Hille Ris
Lambers) Apterae are pale green with
bright green spinal and marginal longitudinal stripes; BL 3.9-4.2 mm.
Originally beaten from mixed vegetation, subsequently collected from Poaceae
(Agrostis, Calamagrostis) (BMNH collection,
leg. G.W. Simpson and M.E. MacGillivray). In
north-eastern USA (Maine) and Canada (Quebec). There are also recent records
from western USA (Idaho, Oregon, Utah) including oviparae collected from a
lakeside sedge in October (A. Jensen, aphidtrek.org). Sitobion
berchemiae (Takahashi) Apterae are blue-green, BL c. 1.8
mm. On Berchemia lineata and B.
floribunda in China. S. berkemiae (Shinji)
is possibly a synonym. Sitobion
berkemiae (Shinji) Apterae are deep green with mainly black
antennae and legs, black siphunculi and a pale yellow cauda; BL 1.3-1.8 mm
(Miyazaki 1980a). On Berchemia racemosa in Japan. The life cycle is unknown. Sitobion (Metobion) brevirostre (Heikinheimo) Apterae are blackish green with dark
antennae and pale siphunculi; BL c. 4.1 mm. The host plant is unknown, but
assumed here to be a species of Poaceae. Only known from Finland. Sitobion breyniae Noordam
Apterae are dull green with pale brown head, black siphunculi and brownish yellow cauda; BL
1.8-2.3 mm. On Breynia microphylla in
Java (Noordam 2004). Sitobion burundiense Remaudière Apterae are shining dark green with black
siphunculi and rather dark antennae, legs and cauda; BL 2.1-2.9 mm. It
differs from other African Sitobion
in having 4 hairs (not 3) on first tarsal segments. On Kotschya
africana and Cassia sp. in
Burundi (Remaudière
& Autrique 1985). It apparently only occurs on Leguminosae/Fabaceae. The
life cycle is unknown. Sitobion (Metobion) calvulum (Ossiannilsson) Apterae are dull bluish green, immatures
yellowish green; 2.1-2.5 mm. Originally described from Poa arctica and there have
been subsequent collections from this plant, but colonies have also been
found apparently feeding on Salix
polaris and above-ground stems of its root parasite Pedicularis hirsuta. Only
known from Spitzbergen. Monoecious
holocyclic; the life cycle is abbreviated with oviparae and apterous males in
late July-early August (Hodkinson et al.
2004, as Acyrthosiphon). It is
parasitised by an endemic aphidiine species (Chaubet et al. 2013). Sitobion caricis (Glendenning)
Only alatae are described (possibly of more than one species), on Carex sp. in northern British
Columbia, Canada. Sitobion cissi (Theobald) Apterae are variable in colour, from
pinkish brown to almost black, with some whitish wax intersegmentally; BL
2.0-2.5 mm. The extent of dark dorsal pigmentation varies from total absence
in samples from South Africa, to broad dark bars or paired patches on all
tergites in a collection at over 2000m altitude in Kenya (BMNH collection,
leg. J.H. Martin). Alatae are brown to black with similarly variable dorsal
pigmentation. On Cissus and Rhoicissus in Kenya, Zimbabwe and
South Africa. Monoecious holocyclic, with oviparae and alate males collected
in June in Pretoria (Eastop 1958). Sitobion colei (Eastop) Apterae are probably
blackish in life; BL 1.7-2.3 mm. On several genera of Lamiaceae (Aeollanthus, Ocimum, Plectranthus, Salvia, Solenostemon, Tetradenia) in
southern Africa (Remaudière & Autrique 1985). Sitobion congolense (Doncaster & Hille Ris Lambers) (Fig.57b) Apterae are mid- to dark green; BL 2.2-2.5
mm. On young growth of Compositae (Vernonia
spp., Elephantopus scaber),
with records also from the guttiferous tree Harungana madagascarensis, which seems able to act as a reserve
host for both this aphid and S.
nigeriense (Remaudière & Autrique 1985). In Africa (Angola, Burundi,
Cameroon, Congo, Ghana, Ivory Coast, Kenya, Nigeria, Zaire). Attended by ants
(Duviard 1969), and parasitized by Aphidius
camerunensis (Remaudière & Autrique 1985). The life cycle
is unknown. Sitobion dismilaceti (Zhang)
Apterae are yellowish; BL c.2.7 mm. On Smilax china in Yunnan, China. This is possibly a synonym of S. sikkimense (q.v.). Sitobion eulophiae Remaudière
Apterae are black in life (but macerated specimens are pale with black
siphunculi and femoral apices and a dark cauda; BL 2.1-2.6 mm. Alatae are
without dark dorsal abdominal markings. On the orchid species Eulophia horsfallii (= rosea) in Burundi. (Not attacking E.
streptopetala, the host of S.
anselliae at the same locations). It was also found on Liparis neglecta (Remaudière &
Autrique 1985). Sitobion fragariae (Walker) Plate 26f Apterae on Rubus and other Rosaceae in spring are yellowish green with small
brown intersegmental markings, with siphunculi either entirely dark or pale
with darker apices (see
influentialpoints.com/Gallery); BL 2.0-3.0 mm. Heteroecious holocyclic,
migrating to Poaceae, where apterae are dirty greenish yellow with black
siphunculi, and BL 1.6-2.9 mm. Although the principal primary hosts are Rubus, especially R. fruticosus, the sexual phase may also sometimes occur on Fragaria, Rosa and Geum. In
Europe, Asia, and introduced to South Africa, North and South America, and
Hawaii. Type specimens of Uroleucon
qinghaiense described from vagrants on Artemisia sp, in China (Zhang et
al. 1999) are S. fragariae or a
close relative. An aphid introduced into Australia and New Zealand and
studied as Sitobion sp. “near fragariae” (e.g. Sunnucks et al. 1996) has a relatively long
cauda and very few microsatellite alleles in common with S. fragariae from southern England, but morphometric studies
indicate that these are probably founder effects (Hales et al. 2010). 2n=18. Sitobion (Metobion) graminearum (Mordvilko) Apterae are light
green or pink, with dark antennae ; BL 3.1-4.0 mm. On Poaceae (Arrhenatherum, Calamagrostis, Deschampsia,
Festuca, Phalaris, Poa) in Europe (Sweden, Finland, north-west Russia,
west Siberia, Czech Republic, and possibly Ukraine). Stekolshchikov (2011)
redescribed apterae, alatae and sexual morphs and designated a lectotype.
Monoecious holocyclic, with oviparae and alate males in September. Sitobion graminis Takahashi Apterae are dark reddish brown covered
with grey wax dust, often of a bluish appearance (red in alcohol), with black
antennae, legs, siphunculi and cauda; BL 1.8-2.9 mm. On many genera and
species of Poaceae, usually feeding on the flower stems. Also recorded from Cyperus and Juncus. In Africa,
Réunion, India, Malaya, Java, New Guinea and the Philippines. Apparently
anholocyclic. 2n=18. Sitobion gravelii (van der Goot) Apterae are pale yellow or pale brownish,
with distal segments of antennae dark brown, siphunculi dark brown and cauda
pale; BL 3.6-4 mm. Described from an unidentified host, but there are later
records from Artemisia and
unidentified Compositae (Raychaudhuri, 1980, as Macrosiphum spinotibium). However these seem unlikely hosts, and
it is more probably a grass feeder. In northern India. Oviparae were
collected from an unknown host in late
November (A.K. Ghosh et al. 1971d,
as M. spinotibium). 2n=12
(Khuda-Bukhsh & Basu 1987, as M.
spinotibium on Artemisia vulgaris). Sitobion
halli (Eastop)
Apterae are uniformly pale green, with black-tipped siphunculi and
pale cauda; BL 1.8-2.9 mm. Immatures are somewhat wax-powdered (Eastop 1959).
On young growth of trees in various families (Apocynaceae, Rubiaceae,
Moraceae, Euphorbiaceae, Melianthaceae, Myrtaceae) in central and southern
Africa (Burundi, Tanzania, Zimbabwe, South Africa; Remaudière & Autrique 1985).
The life cycle is unknown. Sitobion
hillerislambersi van Harten Apterae are bright grass-green with a
black oval dorsal abdominal patch and black siphunculi; BL 1.7-2.2 mm. On
undersides of leaves of Cyperus
esculentus in Angola. Sitobion himalayensis L.K. Ghosh
Apterae are greenish with dark siphunculi; BL 3.7-4.3 mm. On inflorescences
of Oryzopsis lateralis in Himachal
Pradesh, India. Sitobion hirsutirostris (Eastop)
Apterae are pale green with a slightly darker spinal stripe,
siphunculi black and cauda pale; BL 2.5-3.1 mm. On Anisopappus africanus, and less commonly on Helichrysum setosum (Remaudière & Autrique 1985). In Africa (Burundi, Cameroun). Sitobion ibarae (Matsumura) Apterae yellowish
green or yellowish brown with pale brownish head, mainly dark antennae and
legs, long blackish siphunculi and a long pale cauda; BL 2.1-3.1 mm. On wild
and cultivated Rosa spp. in east
and south-east Asia, and introduced to Hawaii (BMNH collection, leg. E.
Hardy). The life cycle is unknown, perhaps anholocyclic at least in warmer
countries. Dwarf, brownish apterae may occur in Japan in midsummer (Takahashi
1964). Sitobion indicum A.N. Basu
(Fig.25c) Apterae are probably yellowish green, with
an irregular dark dorsal abdominal patch, black siphunculi and pale brown
cauda; BL c. 2.0-2.1 mm. On orchids, mostly Cymbidium spp., in hilly regions of north India (West Bengal,
Himachal Pradesh, Uttar Pradesh. Possibly it is entirely anholocyclic. A
record from Rosa indica (Kurl 1980)
is likely to be based on a misidentification. 2n=18. Sitobion javanicum Noordam
Apterae are shiny brownish red, with brown to blackish siphunculi and
brown cauda; BL 2.0-2.3 mm. On Themeda
arguens, living between spikelets, in Java (Noordam 2004). Sitobion kamtshaticum (Mordvilko)
Apterae are green with black antennae and siphunculi; BL 1.5-2.9 mm.
Alatae have rhinaria on ANT IV as well as III. On young growth of Rosa multiflora and Rubus sp. in spring. Heteroecious holocyclic, migrating to grasses (Digitaria adscendens and Microstegium vimineum; Miyazaki 1971,
as rubiphila). In Japan, Korea and east Siberia. Sitobion
krahi (Eastop)
Apterae are green, with siphunculi pale only on basal third and a pale
cauda; BL 2.5-3.0mm. Alatae have siphunculi wholly black and dark marginal
abdominal sclerites, small antesiphuncular sclerites and large
postsiphuncular sclerites. Collected on growing points of saplings of a
forest tree, probably Rauvolfia
vomitoria but possibly Tabernaemontana
chippii, in Ghana (Eastop 1959). Specimens trapped or on unidentified
hosts have been collected in Nigeria, Sierra Leone and Tanzania (BMNH
collection), and Remaudière & Autrique (1985) recorded krahi from Mangifera indica in Burundi. The life cycle is unknown. Sitobion lambersi David
Apterae are pale green with brown siphunculi and a pale cauda; BL
c.1.8 mm. On stems and in flower heads of tropical grasses (Bothriochloa, Chloris, Cynodon, Dicanthium, Digitaria, Ischaemum, Panicum, Paspalum, Themeda), and
occasionally on Cyperaceae. Widely
distributed in the Old World tropics; Africa, southern India, Sri Lanka,
Java, and also recorded from Fiji. Two alatae have also been trapped in
Brazil (C.L. Costa, pers. comm.).The name is possibly being used for a
complex of species in the Old World tropics, which share the distinctive
character of a cauda with only 2 long hairs. Sitobion leelamaniae (David) Banded Cereal Aphid (= chanikiwiti
Eastop 1959, = howlandae Eastop 1959; synonymy by Eastop
& Blackman 2005) Apterae
are pale yellowish green to green or bright yellow, antennae ringed with
brown, legs pale, siphunculi pale brown, darker towards apices; BL 1.6-2.3
mm. On undersides of leaves of many species of grasses and cereals (Brachiaria, Cynodon, Eleusine, Pennisetum, Sorghum, Zea).
Described from south India, and also in Sri Lanka, Réunion (as S. chanikiwiti), and widely
distributed in southern Africa, where most records are as S. chanikiwiti and S. howlandae. Possibly heteroecious
holocyclic with Hagenia abyssinica as
primary host in Burundi (Remaudière & Autrique 1985, as S. chanikiwiti). Sitobion
leonidasi Remaudière
Apterae are variably pigmented but usually with a shiny black abdomen
and both siphunculi and cauda black; BL 1.8-2.5 mm. Alatae have dark wing
veins and variably developed dark abdominal markings. On Bersama ugandensis, living particularly in leaf galls formed by a
psyllid. Only known from Burundi (Remaudière & Autrique 1985). Sitobion loranthi van Harten
Apterae are green with siphunculi and apices of antennal segments
black; BL 1.7-2.7 mm. On upper and undersides of leaves of Loranthus sp. in Angola. There are similar aphids in the BMNH
collection from Loranthus dregei in South Africa, but with a longer and more
pointed R IV+V. Sitobion luteum (Buckton) Yellow Orchid Aphid (Fig.25b) Apterae
are bright greenish yellow to pale yellowish green, with an oval black dorsal
abdominal patch, antennae black except for base of ANT III, black siphunculi
and a pale cauda; BL 1.3-2.1 mm. Alatae have only dark intersegmental
markings. On many species of orchids (Dendrobium,
Epidendrum, etc.) with an almost
world-wide distribution, in temperate regions usually under glass. In Europe,
Madagascar, Mauritius, India, Singapore, Java, New Guinea, Australia, Fiji,
Tahiti, USA (New Mexico, New York), central and South America. Apparently
entirely anholocyclic; no sexual morphs have been recorded. 2n = 12. Sitobion matatum (Eastop)
Apterae are shining black dorsally; BL 2.0-2.5 mm. On Tricholaena rosea (=Rhynchelytron repens) in Kenya. Sitobion microspinulosum David, Rajasingh &
Narayanan Apterae are olive green,
BL c.2.8-3.0 mm. Described from Arthraxon
lancifolius in West Bengal, feeding on stalk and inflorescence. Only
known from types (holotype aptera and paratype immature alata), which are in
BMNH collection. Sitobion milii Remaudière Apterae are grey
with black segmental cross-bands; BL 1.7-2.5 mm. On Euphorbia milii in Burundi. Closely related to S. phyllanthi; see Remaudière &
Autrique (1985) for a comparison of the two species. Sitobion miscanthi (Takahashi)
Apterae are variably coloured green, reddish brown to dark brown with
shiny black siphunculi, pale cauda, and dorsal cuticle variably tanned from
pale with only very small intersegmental sclerites to very dark (in the latter case the pigmented area may
be entire or segmentally divided); BL 1.7‑3.0 mm. On many genera and
species of Poaceae. Also on Cyperus,
and sometimes found in large numbers on dicots, particularly semi‑aquatic
species, e.g. Polygonum hydropiper
(Eastop, 1966). In Indian
subcontinent, east and south-east Asia, Australia, New Zealand, and Pacific
islands, including Hawaii. Records
from Turkey (Görür et al. 2011b, 2012)
require additional confirmation. Anholocyclic almost everywhere, but an ovipara
(on Polygonum chinense) and males
collected in India may be this species (see David 1976), and oviparae and
males have been reared from the introduced populations in Australia (Hales et al. 2010). S. akebiae in Japan and Korea is closely related and on secondary
hosts not practically distinguishable from S. miscanthi; it may be a synonym, in which case one would expect
the holocycle to involve overwintering as eggs on various dicots, as in
Japan. The taxonomy of the miscanthi/akebiae
group requires clarification by a combination of molecular and biological
studies (see also under S. avenae). Sunnucks et al. (1996) studied DNA and karyotype variation in the
introduced populations in Australia, and concluded that probably two
successful colonizations had occurred.
Genetic evidence suggests that hybridisation may have occurred between
introduced populations of S. miscanthi
and Sitobion ‘sp. near fragariae’ (see above) in New Zealand
(Hales et al., 1998), although the
sexual morphs of neither of these species have so far been found in the field
there. Recent studies on S. miscanthi
populations in China have revealed two strains of Wolbachia (Z.Wang et al.
2009) and a new Rickettsia-like
secondary symbiont (T. Li et al.
2011, 2016). See also Blackman & Eastop (2000). 2n = 18 in India (Kurl
& Chauhan 1988), but varies from 17 to 21 in Australian populations. Sitobion mucatha (Eastop)
Apterae are of two colour forms, both appearing translucent and
delicate, one very pale yellow with dark siphunculi and the other pale
yellow-green with dusky siphunculi; BL 2.0-2.5 mm. Living singly or in small
colonies on undersides of leaves of Vernonia
lasiopus in Kenya. Sitobion neusi (Eastop) Apterae are shining
black; BL 1.7-2.3 mm. On various tropical grasses (Digitaria, Eragrostis, Panicum, Setaria, Tricholaena),
and also found on Canna and Ensete (BMNH collection). In Kenya,
Ethiopia, Burundi (at 2000m) and Madagascar. Sitobion nigeriense (Eastop) (Fig.57c) Apterae are green with dark siphunculi; BL
1.5-2.2 mm. On young growth of Vernonia
spp. (BMNH collection), and also
recorded from Guttiferae (Harungana
madagascarensis and Psorospermum
febrifugum), which possibly act as reserve hosts (cf. S. congolense), although Remaudière
& Autrique (1985) recorded it twice colonizing H. madagascariensis in Burundi. Parasitism by a ?Trioxys sp. was observed. In Africa (Burundi, Congo,
Nigeria, Kenya, Ethiopia). The life cycle is unknown. Sitobion nigrinectarium (Theobald)
Apterae are green with black siphunculi and a long pale cauda; BL
1.8-2.9 mm. On various Leguminosae/Fabaceae (Cajanus cajan, Glycine
javanica (= Neonotonia wightii),
Crotalaria, Dolichos, Pisum, Rhynchosia) in Africa, and also on
Indian Ocean islands (Madagascar, Mauritius, Réunion, Anjouan). See also
Blackman & Eastop (2000). 2n=18. Sitobion
ochnearum (Eastop)
Apterae are salmon-pink (but this colour was possibly due to fungal
infection); BL 1.7-2.3 mm (Eastop 1959). On Ochna spp. in eastern and southern Africa (Angola, Kenya, South
Africa, Zambia, Zimbabwe). The life cycle is unknown. 2n=18. Sitobion orchidacearum (Franssen & Tiggelovend) Apterae are shiny
black with dark red around the siphunculi; BL 1.4-2.7 mm. On Orchidaceae (Coelogyne, Dendrobium) in Java (Noordam 2004). Sitobion paludum Müller Apterae
are shining green or brownish, with black head and antennae, legs with femora
black except at bases, black siphunculi and pale cauda; BL 1.7-3.0 mm. On Vaccinium uliginosum in northern
Europe. Heteroecious holocyclic, migrating to Rhynchospora alba (Cyperaceae), where apterae are darker than on
the primary host. Müller (original description) gave a full account of its
biology. Sitobion papillatum Remaudière
Apterae are matt dark brown, with black siphunculi sometimes paler at
base; BL 1.7-2.5 mm. In inflorescences of Pennisetum
purpureum in Burundi (Remaudière & Autrique 1985). A shorter-haired form
on a bamboo, Arundinaria (= Yushania) alpina, was distinguished as a subspecies, S. papillatum ssp. subnudum Remaudière (although if
the differences remain consistent after further collections this form should
probably be given separate species status). The life cycle
is unknown. Sitobion pauliani Remaudière
Apterae are pale green or pink, usually with a shiny dark olive green
or brown dorsal patch; BL 1.5-2.1 mm. In inflorescences of grasses,
especially Pennisetum, Setaria and Eleusine. In Africa south of the Sahara, Madagascar, Réunion
(BMNH collection, leg. G. Remaudière), Andaman & Niccobar Isles, India
(Orissa), Sri Lanka, Indonesia, West Malaysia (BMNH collection, leg. J.H.
Martin), Philippines (BMNH collection, leg. V.J. Calilung), Central America
(Venezuela, Honduras, Cuba, Jamaica, Costa Rica; BMNH collection, leg. J.H.
Martin) and South America (Brazil, S.P.; S. Matsuoka, pers. comm.). Sitobion phyllanthi
(Takahashi) Apterae are dirty olive green with dark brown head and dorsal
markings (Müller 1959; as S. adgnatum);
BL 1.5-2.1mm. Prepared specimens of apterae have variably developed narrow dorsal
abdominal cross-bands between dark marginal sclerites. Described from Phyllanthus sp. in Mauritius
(Takahashi 1937c). An alata probably of this species was trapped in Sri Lanka (BMNH
collection). Remaudière (1988) proposed that S. adgnatum Müller, widely distributed
on various Euphorbiaceae (Euphorbia, Breynia, Phyllanthus) in Africa (Millar 1994), is this
species, and the BMNH collection contains specimens collected by A. van
Harten on Phyllanthus maderaspatensis
in Angola. It is also reported to occur in Hawaii. Müller (1959) gave a account of this species (as S. adgnatum). The life cycle is unknown. Sitobion plectranthi M.R. Ghosh,
A.K. Ghosh & Raychaudhuri
Apterae are pale-coloured; BL 1.7-2.1 mm. On stems and flowerstalks of
Plectranthus coetsa, and also
recorded from Perilla sp. In
north-east India. Sitobion pseudoluteum A.K. Ghosh
Colour of apterae in life is unknown; BL 1.7-2.3 mm. On Masdevallia
sp. (? - recorded as Madenvallia)
and Cymbidium spp. in West Bengal and
Sikkim, India. The consistency of the unusual first tarsal chaetotaxy (4,4,4)
needs to be confirmed with additional material. It appears closely related to
S. orchidacearum in Indonesia, and
could be a synonym. The specimens with a dark cauda and long hairs referred
to by David (1976) under the name S.
indicum may be this species. 2n=18 (Kar et al., 1990). Sitobion raoi (Kulkarni) Apterae are pale
brown (in life?); BL 2.1-2.5 mm. On Iseilema
anthephoroides in Maharashtra, India. Sitobion rosaeiformis (Das)
Apterae are bright yellowish green, green or pinkish brown with
antennae and legs mainly pale, and siphunculi entirely or mainly dark with
pale bases; BL 2.1-3.3 mm. On both cultivated and wild Rosa in India, Pakistan, Bhutan (Chakrabarti & Das 2014) and
Vietnam (Szelegiewicz 1968). Also recorded
from Rubus lasiocarpus and Spiraea corymbosa. Probably heteroecious holocyclic with migration to grasses, as
it has been collected on Digitaria
adscendens (= ciliaris), although
host alternation has not been confirmed experimentally by host transfers, and
populations can persist on roses into the cold season. Oviparae and alate
males were found on Rosa in
November-March (David 1976). 2n=18. Sitobion rosivorum Zhang
Colour of apterae in life is unrecorded; BL c. 4.2 mm. On Rosa chinensis and Rosa sp. in China. 2n=18 (Chen &
Zhang 1985a). Sitobion schoelli (Müller)
Apterae are green, with head pale brown, antennae and legs with dark
apices to segments, siphunculi black and cauda dusky; BL 1.4-2.2 mm. On Kniphofia sp. in South Africa, and
Angola (BMNH collection, leg. A. van Harten). Sitobion (Metobion) scoticum (Stroyan) Apterae are dull apple-green with dark
antennae and pale brownish siphunculi; BL 2.3-3.0 mm. On Poa trivialis, living on uppersides of leaves and hidden in
inflorescences, in marshy habitat. Only known from Scotland. Monoecious
holocyclic, with alate males. Sitobion sikkimense (A.K. Ghosh & Raychaudhuri) Apterae are undescribed, probably pale
green with black siphunculi, and BL 2.5-3 mm. On undersides of leaves of Smilax sp. in northern India, and
subsequently recorded from S. ferox (David
1976, as M. smilacifoliae ssp. ferocis). Monoecious holocyclic with
oviparae and alate males in late October (Raychaudhuri et al. 1980). S.
dismilaceti, described from Smilax in China, may be a synonym. Sitobion smilacicola (Takahashi)
Apterae are pale yellow with black siphunculi and yellow cauda; BL c.
3 mm. Alatae have 6-8 secondary rhinaria on ANT III. On young leaves of Smilax stenopetala in Taiwan, and
recorded also from S. china in
Korea (Paik 1965). Both apterae and alatae are described as having femora and
tibiae with numerous small circular sensoria or pores. Records from China and
India may apply to other species. Sitobion smilacifoliae (Takahashi)
Apterae are bright green to orange-red with very variably developed
shining brownish black dorsal sclerotisation, femora with distal half black,
siphunculi black and cauda yellow; BL 2-3 mm. Alatae have 5-8 secondary
rhinaria on ANT III. On undersides of leaves of Smilax spp. in east Asia (Taiwan, China, Korea, Japan). Specimens
from Smilax spp. in Australia, and
possibly those previously recorded from Hong Kong and Thailand, seem to be a
different species, the identity of which needs further study (Hales et al. 2010). Sitobion
takahashii (Eastop)
The colour of apterae in life is not recorded in the original
description (Takahashi 1938a, as Macrosiphum
phyllanthi), probably greenish, with black antennae and siphunculi; BL
1.7-2.3 mm. On Phyllanthus spp. in
China, Thailand and India, and perhaps also on Mimosa pudica in India (M.R. Ghosh et al. 1977a). [Specimens from Phyllanthus (maderaspatensis,
urinaria) from southern India, Java
and Singapore in the BMNH collection have a longer antennal PT and shorter R
IV+V than takahashii s. str., and
seem to be the form referred to as a subspecies by David (1958); possibly the
name mimosae Ghosh, Basu and
Raychaudhuri is applicable to this form, but further work is necessary to
confirm that there are indeed two very similar species, both living on Mimosa and Phyllanthus.] 2n = 18
(Khuda-Bukhsh & Kar 1989). Sitobion thalictri Remaudière
Apterae are dark green with black antennae and siphunculi, and dark
head, legs and cauda; BL 2.0-3.1 mm. On Thalictrum
rhynchocarpum in Burundi. Sitobion triumfettae Remaudière
Apterae are very pale yellowish green with a darker green spinal
stripe and sometimes dusky dorsal cross-bands; BL 1.7-2.3 mm. On Triumfetta
rhomboidea in Burundi. Sitobion
wikstroemiae (Mamet) Colour of apterae in life is unrecorded;
BL 1.8-2.4 mm. On Wikstroemia
viridiflora in Mauritius (original description) and Sri Lanka (BMNH collection, leg J.
Edirisinghe). Specimens collected in Kenya on Gnidia sp. (BMNH collection, leg. J.H. Martin) and trapped in
Zimbabwe (BMNH collection, leg. C.E. Taylor) also seem to be this species.
2n=16. Sitobion yakini (Eastop) Apterae are green,
with distal parts of antennae dusky to black, and siphunculi dark distally,
sometimes paler at base; BL 1.5-2.2 mm,.
On various tropical grasses (Andropogon,
Eleusine, Eragrostis, Pennisetum,
Poa, Tricholaena, Triticum)
in Africa (Burundi, Kenya, South Africa). Sitobion
yasumatsui (Moritsu) Apterae are shiny yellowish brown with a
variably-developed dark brown-black dorsal patch (see Moritsu 1983); BL c.2.1
mm. Alatae have more than 40 rhinaria on ANT III. On Spodiogon (= Eccoloipus) cotulifera in Japan, and Andropogon ascinodis in Thailand (BMNH
collection, leg. H. Banziger).
Four
species closely related to Aploneura,
producing sac-like or coral-like galls on Pistacia
spp. in south-west and central Asia. Three species produce emigrant alatae and
are apparently host-alternating, with a two-year life cycle as in most other
host-alternating Fordini, but their secondary host generations are still
unknown. The fourth lives exclusively on Pistacia.
Davatchi (1958) revised the genus. Slavum
esfandiarii Davatchi and Remaudiere Galls on twigs of Pistacia mutica in Iran are coral-like, red when mature, with
branches slightly larger than those of S.
wertheimae and S. mordvilkoi
(Davatchi & Remaudiere 1957). Galls were also recently found on Pistacia sp. in Turkey (Şenol et
al.
2014). Heteroecious holocyclic; alatae (BL 2.0-2.2 mm) emerge in November and
fly to an unknown secondary host. Slavum
lentiscoides Mordvilko Galls (fig. 126N)
are bag-like outgrowths of the leaves of Pistacia
spp., similar in position to those of Geoica,
but with pointed apices. In Crimea, Iran, Kazakhstan (Tien-Shan),
Tadzhikistan, Turkmenistan and Pakistan . Heteroecious; alatae (BL 1.6-2.2
mm, fig. 81B)
emerge in October and are reported to migrate to roots of various grasses
(Kadyrbekov 2002d, 2014d), but the secondary host generations are
undescribed. It is possible that S.
lentiscoides is the primary host form of Aploneura ampelina, which lives on roots of vines in southern
Europe, Africa and south-west and central Asia, and is also reported from grasses, but
experimental work is needed to confirm this. Slavum
mordvilkoi Kreutzberg
Galls on twigs of Pistacia vera
are coral-like, similar to those of S.
wertheimae. Recorded only from wild P.
vera, in Turkmenistan, Afghanistan and eastern Iran (Kreutzberg, 1953).
Apparently heteroecious, but only the primary host part of the life cycle is known. Slavum
wertheimae Hille Ris Lambers Galls are branching, coral- or
cauliflower-like (fig. 126O),
reddish when mature, 5-15 cm in diameter, on twigs and branches of Pistacia atlantica and P. mutica in Iran, Israel and Syria.
Monoecious holocyclic, presumably with a one-year cycle; alatae (BL 1.7-2.2
mm) emerging from galls in September are sexuparae and presumably give birth
to sexuales on the trunk, although this has not been recorded. Wool &
Manheim (1983) studied morphological variation, Aloni et al. (1989) studied the differentiation of plant tissue in the
galls, Álvarez et
al. (2016) studied their ultrastructure in comparison with other Fordini,
and
Martinez (2008) studied the impact of the galls on subsequent growth of the
host trees.
Four
palaearctic crucifer-feeding species related to Brevicoryne, with broad dark volcano-shaped siphunculi,
a short cauda, and apterae usually with rhinaria on ANT III. Smiela fusca Mordvilko Plate 12g
Apterae are grey-brown,
wax-powdered; BL 1.4-1.8 mm. On Berteroa
spp., deforming leaves and flower
buds (Müller 1975a), and now also recorded from Armoracia rusticana (Basky 2016). In eastern Europe (Germany,
Poland, Hungary, Bulgaria, Czech Republic, Ukraine). Oviparae occur in
October (BMNH collection, leg J. Holman). Smiela mongolica Holman & Szelegiewicz Apterae are green, densely covered in wax
powder, with mainly dark appendages; BL 0.9-1.2 mm. On Alyssum (= Ptilotrichum)
canescens, colonising terminal
parts of shoots, in Mongolia. Other morphs are undescribed. Smiela schneideri (Hille Ris
Lambers) Apterae are blackish,
probably wax-covered in life; BL 1.2-1.5 mm. Alatae have dark transverse bars
on the dorsal abdomen. On Isatis glauca,
colonising the stem apices and inflorescences, in Syria. A population found on Alyssum
tortuosum in Crimea, with fewer secondary rhinaria on the antennae (of
apterae) and a shorter antennal terminal process, was described as a
subspecies, S. schneideri ssp. alyssii Mamontova-Solukha. Sexual morphs and life cycles are unknown. Smiela syreniae Bozhko Apterae are greenish brown-black, covered
with dense white wax powder; BL c.1.6-1.7 mm. On Erysimum spp. and Syrenia spp.
In south-west Russia, Ukraine, Slovakia and Central Asia.
One
species related to Iziphya but with
a very different pattern of dorsal sclerotisation, all tergites bearing
numerous wax glands. Sminthuraphis ulrichi Quednau (Fig.18j) Apterae are plump-bodied, covered in bluish
white wax: BL c.1.5-1.6 mm. Living on ground-level stems of Carex ligerica in Germany, and also
recorded from Carex spp. in France,
Poland, Hungary and Czech Republic (BMNH collection), and east to Kazakhstan
(Kadyrbekov 2011b). Alatae have similar dorsal sclerotisation and wax gland
distribution to apterae, bordered wing veins with dark spots at the ends, and
secondary rhinaria distributed III 9-11, IV 1-3 (BMNH collection). Living all
year on Carex, with sexuales
collected in October in France (Leclant 1966).
One
species characterized by the elongate second antennal segment and the thick
sclerotic rims of the primary sensoria on the last two segments. Smynthurodes
betae Westwood Galls on Pistacia spp. (atlantica,
mutica and, rarely, vera) are
yellow-green or red, spindle-shaped, about 20mm long, formed by rolling of
the edge of the leaflet near its base (fig. 126P).
These are secondary galls, produced by the progeny of the fundatrix, which
lives in a small red mid-rib gall (Burstein & Wool 1991). The
ultrastructure of the galls was studied by Álvarez et al. (2016). Host-alternating,
with a two-year cycle; alatae (BL 1.3-1.6 mm, fig. 81O)
emerge in September-November and migrate to roots of numerous, mostly
dicotyledonous, plants. Secondary
hosts are particularly
Compositae/Asteraceae (Artemisia, Arctium), Leguminosae/Fabaceae (Phaseolus, Vicia, Trifolium), and
Solanaceae (Solanum tuberosum, S. nigrum, Lycopersicon
esculentum); also sometimes on Beta,
Brassica, Capsella, Gossypium, Heliotropum, Rumex, etc. Only rarely is it found on monocots (Poaceae, Cyperaceae).
Apterae on roots of secondary hosts are dirty yellowish white, wax‑dusted,
with light brown head, prothorax, ANT and legs; BL 1.6‑2.7 mm (Plate 2f). The
holocycle is recorded throughout the range of the primary hosts; Algeria,
Morocco, Israel, Syria, Iran, southern Crimea, Transcaucasus (Mordvilko,
1935, as Trifidaphis phaseoli;
Davatchi, 1958) and Pakistan (Naumann-Etienne & Remaudière 1995).
Anholocyclic populations occur commonly on secondary hosts in other parts of
the world (see also Blackman & Eastop 2000, p.347). Wool & Burstein
(1991a, b) studied phenology and ecology of gall generations and interactions
with a parasitoid in Israel. 2n=8.
One
species perhaps related to Sappaphis,
but with 4 instead of 3 hairs on 1st segments of fore and mid-tarsi. Sorbaphis
chaetosiphon Shaposhnikov Plate 10g Spring colonies curl leaves of Sorbus spp. downward and turn them
reddish. Host-alternating; emigrant alatae (BL 2.1-3.0 mm, with shining black
abdomen due to an extensive sclerotic patch), migrate in late May-June to Ligularia (Stekolshchikov &
Shaposhnikov 1998), where they live in ant shelters on stems or at bases of
leaves. Apterae on secondary hosts are grey-brown to blackish, with immature
stages dirty yellow; BL 1.7-2.6 mm. Stekolshchikov & Shaposhnikov (1998)
provided a redescription including fundatrix, emigant alata and apterous
exule. In Japan, Korea, China, west and east Siberia and the southern Urals.
2n = 38.
Two
species in Europe related to North American Landisaphis but with 2 hairs (no sense peg) on HT I, and without
the development of the spinal sclerites on abdominal tergites 6-8 into rugose
conical processes. Spatulophorus alyssi Holman
Apterae are greyish green with slight pruinosity, especially
ventrally; BL 1.1-1.3 mm. Found in
September feeding (with sexual morphs) on the upper sides of the youngest
terminal leaves of young shoots of Alyssum
saxatile in central Bohemia, Czech Republic. The inflorescences are
possibly colonised earlier in the season. Monoecious holocyclic, with
apterousmales. Alatae are unknown. Spatulophorus incanae Müller
Apterae are pale green to yellowish green or grey-green, BL 1.5-1.9
mm. Living inconspicuously on flower stalks and in inflorescences of Berteroa incana, and after flowering
on undersides of leaves. Also recorded from Capsella bursa-pastoris. In
eastern Europe (Germany, Hungary, Ukraine). Monoecious holocyclic on Berteroa, with apterous males. Müller
(original description) gave a very full account of this species.
One
or more palaearctic species on Galium and
related Rubiaceae, differing from Hydaphias
in the shape of the siphunculi. A species in Tasmania, two oviparae of
which were collected on Coprosma (BMNH
collection, leg. M Williams) may also belong to this genus. The unnamed
species from Sorbus aucuparia
described as a Staegeriella by
Bozhko (1976, p.30) probably does
not belong in this genus, and could perhaps be the primary host form of Pseudacaudella rubida. Staegeriella necopinata (Börner) Apterae are dark greyish green to
lead-coloured, powdered ventrally with grey wax; BL 1.3-2.2 mm. On Galium spp., and probably also Asperula spp., colonising the stems
and flowerheads, causing stunting and twisting of new growth. Throughout
Europe, and also recorded from Tunisia (Boukhris-Bouhachem et al. 2007). Oviparae and
alate males were collected in October (Hille Ris Lambers 1947c). A second
species, S. asperulae Bozhko, was described from Asperula cynanchica in Ukraine, and this name was also applied to
populations on A. aristata in
Switzerland (Jörg & Lampel 1988). However, the characters used for
separation of apterae of S. asperulae from those of S. necopinata (ratio of body width to
length and shape of siphunculi) are inconsistent and unreliable, and another
collection of Staegeriella from A. aristata in Switzerland (BMNH
collection, leg. D. Hille Ris Lambers) differs from both S. necopinata on Galium and
from S. asperulae according to Jörg
& Lampel (1988), so it appears that a single, variable taxon may be
involved.
About
12 species similar to Macrosiphoniella, associated with Plumbaginaceae in
salt-marsh and coastal habitats. The taxonomy is very problematic, as several
of the nominal species are poorly known and not clearly differentiated in the
literature. The better-known species exhibit much geographic variation,
accentuated by the fact that populations are distributed linearly along
coasts or isolated in salt marshes separated by deserts. Kadyrbekov (2003b) reviewed and keyed the
western palaearctic species. Staticobium caspicum Bozhko Apterae are shining dark brown; BL c. 2.0
mm. On Limonium sp. in Russia
(north Caucasus). Decribed as a subspecies of S. latifoliae, but probably just a geographical variant of that
species. Staticobium caucasicum Bozhko
Apterae are shining dark brown; BL c.2.2 mm. On upper and undersides
of leaves of a Limonium sp. in
Ukraine. Stekolshchikov (2005) recorded it (as limonii ssp. caucasicum)
from L. gmelinii in southern Russia
(Astrakhan), and Kadyrbekov (2011c) from the same host in west Kazakhstan. Staticobium gmelini Bozhko Apterae are brown, with pale cauda; BL c.
1.9-2.0 mm. On flowerstalks and root collars of Limonium and Goniolimon spp.
in Ukraine, west Siberia and Kazakhstan. S. zolotarenkoi (Ivanoskaya) is a
synonym according to Kadyrbekov (2003b), and its host was probably Limonium gmelinii rather than Helichrysum arenarium. S.
strongilosiphon Bozhko, and perhaps S. insularum, are probably also synonyms. Staticobium insularum Bozhko
Apterae are dark brown with greenish tinge; BL c. 2.0-2.1 mm. On Goniolimon tataricum in Ukraine
(Bozhko 1961 – but on Limonium sp.
according to Bozhko 1976), and Kadyrbekov (2011c, 2014g) recorded it on Limonium gmelinii in Kazakhstan and
Russia. Closely related to S. gmelini,
and possibly a synonym. Staticobium latifoliae Bozhko Apterae are dark brown to blackish; BL 1.6-2.6
mm. On undersides of leaves and flowerstalks of Limonium spp. in Pakistan, Kazakhstan, Tajikistan (Kadyrbekov
2013c), Iran (Rezwani 1990), Lebanon, Russia (north Caucasus, west Siberia),
Ukraine, Bulgaria, Romania, Hungary, Greece and Italy (Barbagallo &
Stroyan 1982). S. tauricum Bozhko
is a synonym according to Kadyrbekov (2003b). S. smailovae Kadyrbekov has morphology intermediate between latifoliae and staticis on the basis of characters given in Kadyrbekov’s (2003b)
description and key. Staticobium limonii (Contarini) Apterae are dark olive-green to dark
brown; BL 1.5-2.2 mm. On Limonium
spp. in southern Europe and the Mediterranean region. Accounts from
north-west Europe are referable to S.
staticis (q.v.). The distinction from S. latifoliae seems unclear, and the separate species status of
these two taxa requires further confirmation. 2n=12*. Staticobium longisetosum Kadyrbekov
Colour of apterae in life is unknown; BL c.2.0 mm (one specimen). On Limonium gmelinii in south-east Kazakhstan
(Kadyrbekov 2003b). Staticobium loochooense (Takahashi)
Colour of apterae in life is unrecorded, probably dark; BL c. 1.4-1.5
mm. On Limonium wrightii in Loochoo
(Japan), and on L. tetragonum in
Korea (Paik 1972). A sample collected from Aster tripolium were presumably vagrants. Redescribed by Sorin
(1967). Staticobium otolepidis Nevsky
Apterae are brown, with antennae dusky except yellow III-IV,
siphunculi black with pale bases, and cauda pale yellow; BL 1.6-2.3 mm. In
large colonies on flowerstalks of Limonium
otolepis and L. perfoliatum in
Central Asia and north-west China. S.
nevskyi Hille Ris Lambers is probably the fundatrix of this species, and
hence a synonym (Kadyrbekov 2003b). Staticobium smailovae Kadyrbekov
Apterae are brownish; BL 1.8-2.2 mm. On stems of Limonium gmelinii in Kazakhstan (Kadyrbekov 2003b) and Russia
(Kadyrbekov 2014g). Staticobium staticis (Theobald) Plate 27f Apterae are shiny, deep green, brown, or
dirty greenish or reddish, with antennae paler with dark apices, siphunculi
black with pale bases, and cauda green, pale red or yellow (see influentialpoints.com/Gallery); BL 2.3-2.7 mm. On leaves and on upper parts of
stems under inflorescences of Limonium spp.
especially L. vulgare (= Statice limonium), often flooded by
high tides and covered with mud (Foster 1984). In coastal areas of north-west
and north Europe. A similar undescribed species but with a shorter antennal
terminal process occurs in the Mediterranean region. Oviparae and apterous
males occur in September in Germany (Müller 1975b, as S. limonii). The sexuales described by Jacob (1948a, as S. limonii) from L. humile in North Wales were
possibly of another closely-related species. Staticobium suffruticosum Kadyrbekov Apterae are dark greenish or light
brownish; BL 1.9-2.3 mm. On stems of Limonium
suffruticosum in Kazakhstan (Kadyrbekov 2003b, 2011c). Very close to S. otolepidis.
Seven
species on nearctic Quercus. Alate
females apparently do not occur and males may be apterous or alate within a species.
Hille Ris Lambers (1966a) keyed the species then known and discussed the
confused nomenclature. Quednau (2010) fully reviewed the genus and provided
illustrations of all available morphs. Stegophylla
brevirostris Quednau Apterae pale greenish white, producing
abundant wax (Halbert 2013); BL 1.2-1.7 mm. On Quercus spp., seeming to prefer Q. virginiana, in
eastern USA. An ovipara and an alate male were collected in late September on
Q. alba in Maryland (Quednau 2010).
Populations on Q. virginiana in
Florida seem to be anholocyclic. Ammar et
al. (2013) studied the ultrastructure of the wax glands and the wax
secretions of this species. Stegophylla
davisi Quednau
(new name for Phyllaphis? querci Fitch of Davis 1911a) Apterae pale greenish, BL not recorded. On Quercus spp. of the white oak group (alba, macrocarpa, stellata)
in eastern North America, forming small colonies on upper and undersides of
leaves (Davis 1911a). Pale brownish oviparae and both apterous and alate
males occur in September-October, the apterous male having an alatiform,
sclerotised thorax (Quednau 2010). Stegophylla
essigi Hille Ris Lambers Apterae are broadly oval, pale grey-green
to olive, covered in white wax wool; BL 1.2-1.8 mm. Feeding by fundatrices in
spring causes edges of young leaves to curl upward and become thickened and
red. On native Quercus spp. in
California, USA, with oviparae and both apterous and alate males produced in
November, but populations may continue parthenogenetically on evergreen oaks
such as Q. agrifolia, especially in
leaves spun together by caterpillars (Hille Ris Lambers, 1966a). It is also
recorded from Oregon (A. Jensen, aphidtrek.org).
2n = 12. Stegophylla
mugnozae Remaudiere & Quednau Apterae are broadly pear-shaped, creamy
white, covered with white or bluish-white wax wool; BL 1.4-1.8 mm. On leaves
of Quercus crassipes and Q. castanea in Mexico (Remaudiere
& Quednau 1985), and subsequently recorded from Panama (Quirós et al. 2009). Other morphs and life
cycle are unknown. Stegophylla
quercicola (Monell)
(= querci of Blackman &
Eastop 1994) Apterae are oval, pale
greenish- or brownish-yellow, covered in dense white wax wool; BL 1.0-1.5 mm.
In small, scattered colonies on upper and undersides of leaves of Quercus spp., usually of the white oak
group (e.g. alba, stellata) in eastern North
America. Oviparae and alate males
occur in September-October (Hottes & Frison 1931; as quercicola Baker). Stegophylla
quercifoliae (Gillette) Apterae are yellowish to
yellowish-green with brown head, wholly covered in white wax wool; BL 1.1-1.5
mm. On leaves of Quercus spp.,
especially Q. gambellii, large
colonies causing leaf curl. Oviparae and alate males occur in October. In
western USA (California, Colorado, Utah, Idaho). Stegophylla
quercina Quednau (= quercicola of Blackman & Eastop
1994) Apterae are pale yellowish-green
with brown head, covered with bluish-white wax wool; BL 1.6-1.9 mm. On leaves
of Quercus spp., usually of the red
oak group, in dense flocculent masses which may cause curling and yellowing
(Quednau, 1966). Oviparae and apterous males occur in October (the males are
true apterae without a sclerotised thorax). In eastern North America. Quednau
(2010) distinguished two subspecies, with S.
quercina s.str. known from Quebec and New York (BMNH collection, leg.
R.L. Blackman), and S. quercina acutirostris Quednau,
known from Wisconsin and North Carolina. The latter subspecies makes pseudogalls
on the infested Quercus spp.
(Quednau 2010). 2n > 30 (for quercina
s. str.).
One
species in Mongolia possibly related to Spatulophorus,
but with more numerous body hairs arising from tubercles, short conical
siphunculi and rather hairy R IV+V and cauda. Stellariopsis songini Szelegiewicz
Apterae are pale green, lightly dusted with wax; BL 1.5-2.1 mm. Alatae
have secondary rhinaria distributed III 29-34, IV 5-6, V 0-1 (1 specimen). On
stems and undersides of leaves of Stellaria
dichotoma in Mongolia.
About
30 palaearctic species of very large, stem- or root-feeding aphids,
remarkable for their very long rostrum which enables them to probe down
through crevices in thick bark of oak and other trees. The morphology of
these remarkable mouthparts was studied by Brożek et al. (2015). Males are small, apterous, lack mouthparts, and
often exhibit morphological aberrations. The evolutionary significance of
male dwarfism in Stomaphis was
discussed by Depa et al. (2015).
This could be a stage in the evolutionary loss of males leading to permanent
parthenogenesis, but it seems more likely to be due to the sheltered,
confined habitat in which males reach maturity in close proximity to females,
so that their only function is to mate before they die. The male genitalia of
four species were compared by Wieczorek (2012). In some species (subgenus Parastomaphis) the males adopt an
unusual, ventro-ventral mating position, with associated morphological
modifications in both sexes (Depa et al.
2014). Colonies
are invariably attended by ants (usually Lasius
spp.). Protaphidius wissmannii is
a specialized parasitoid. Accounts and keys are available for Europe (Heinze
1962, Szelegiewicz 1962, Czylok & Blackman 1991), Iberian peninsula
(Nieto Nafría et al. 2002), Ukraine
(Mamontova 1972), Korea (Lee et al.
1994, Seo 1994), China (Qiao & Zhang 1999d) and Japan (Sorin 2012a). The Acer-feeding
species were reviewed and keyed by Binazzi & Blackman (2004). Depa et al. (2015) provided illustrations
and keys for oviparae and males of European species. Depa & Mróz (2013)
studied mitochondrial DNA and morphometric variation in Stomaphis populations on Acer
and Salix in Europe and discussed
their taxonomic status. Stomaphis
abieticola Sorin Colour in life was not recorded; BL of
aptera c.6.2 mm. On Abies firma in
Japan (Shikoku). Oviparae and apterous males were collected in late October
(Sorin 2012a). Stomaphis
aceris Takahashi
Colour in life was not recorded; BL of aptera 4.2-4.7 mm. On Acer sp. in Japan (Takahashi, 1960b).
Oviparae and apterous males were collected in November (Sorin 1995). Stomaphis
acquerinoi Binazzi
Apterae are elongate oval, greyish white powdered with wax, immatures
shiny yellowish white; BL 4.3-6.4 mm. On Acer
pseudoplatanus in Italy (Tuscany; Binazzi 2001), in colonies on smooth
exposed trunk of young trees, or in bark crevices of older trees. No sexuales
were found, and populations are believed to overwinter as alate viviparae
(Binazzi & Pennacchio 2002). Ant attendant behaviour and parasitism by Protaphidius wissmannii were studied
by Binazzi & Pennacchio (2002, 2005). Depa & Mróz (2013) suggested
that this might be a geographical variant of S. graffii. Stomaphis
alni
Sorin Apterae are dark brown; BL c.
6mm. On Alnus hirsuta in Japan.
Sexual morphs occur in September (Sorin 1965). Stomaphis
aphananthae Sorin Apterae are blackish-brown; BL c. 4.7 mm.
On Aphananthe aspera in Japan.
Sexual morphs occur in November (Sorin 1979a). Stomaphis
asiphon Szelegiewicz
Colour in life is unknown; BL of aptera 4.8-5.9 mm. Described from
roots of an undetermined tree species in Korea (Szelegiewicz 1975).
Pashchenko (1988b) recorded it living in cracks in bark at bases of trunks of
Juglans mandshurica in eastern
Siberia. Sexual morphs of S. asiphon s.str.
are undescribed. However, Sorin (2012a) described fundatrices, apterae
(fundatrigeniae), oviparae and apterous males of Japanese populations on Juglans mandshurica ssp. sieboldiana (= J. ailanthifolia) as a subspecies, S. asiphon ssp. sakuratanii.
The apterae had fully developed siphuncular cones, although these were absent
in fundatrices; they will run to S.
matsumotoi in our key to aphids on Juglans,
and the distinction from that species seems to require further confirmation.
The aphids were living on the surface of the trunk at or just below ground
level, attended by ants, and sexual morphs were produced in November. Stomaphis
betulidahuricae Zhang & Qiao Apterae are powdered with greyish-white
wax; BL c. 5.6 mm. In bark crevices of
Betula dahurica in Hebei, China
(Qiao & Zhang 1999d). Alatae and sexual morphs are undescribed. Stomaphis
bratislavensis Czylok & Blackman Apterae are brownish, coated with powdery
wax; BL 4.3-4.8 mm. On bark of Quercus
petraea in Slovakia, and subsequently recorded from Bulgaria and Turkey (Şenol et
al.
2014). Sexual morphs occur in October (Czylok and Blackman, 1991). The
ovipara was redescribed by Depa et al.
(2015). 2n = 8 (Blackman, 1990). Stomaphis
carpini Sorin
Colour in life is unrecorded; BL of aptera c. 5.5 mm. On Carpinus tschonoskii in Japan (Sorin,
1965). Sexual morphs are undescribed. Stomaphis
cupressi Pintera
Apterae are yellowish-brown, powdered with grey wax; BL 4.9-6.2 mm.
Described from specimens collected on trunk of Cupressus benthami in Kenya (Pintera, 1965), and since found on C. macrocarpa in France (BMNH
collection, leg. G. Remaudiere), on Cupressus
spp. in Italy (BMNH collection, leg. D. Roberti), on Juniperus spp. in Spain (Nieto Nafría et al. 2002), and on C.
sempervirens in Turkey (Akyürek et
al. 2010). Mamontova (1982) described a population on C. sempervirens in Abkhazia (Caucasus)
as a subspecies, S. cupressi sp. caucasica. Sexual morphs are unknown.
2n = 14. Stomaphis
fagi
Takahashi Colour in life was not
recorded; BL of aptera c. 5.3mm. On Fagus
sp. in Japan (Takahashi, 1960b). An oviparae and an apterous male were
described by Sorin (1995), collected in September . Stomaphis
(Parastomaphis) graffii Cholodkovsky Apterae are yellowish,
covered in white wax powder; BL 4.2-6.5 mm. In bark crevices on trunks of Acer spp., usually A. campestre, A. platanoides or A.
pseudoplatanus, often at or below soil level, attended by Lasius spp. (brunneus, balcanicus).
In Europe (UK, France, Italy, Poland, former Czechoslovakia, Rumania,
Ukraine), and east to Georgia. Sexual morphs have been collected from
mid-August to November; they were redescribed by Depa et al. (2015). Depa
(2012) studied populations in Poland. A population in Ukraine was described
by Mamontova-Solukha (1963, 1972) as a subspecies, S. graffii ssp. acerina,
but this was placed as synonym of S.
graffii graffii by Depa & Mróz (2013), who compared mitochondrial DNA
and morphometric data (see also under S.
longirostris). Stomaphis
hirukawai Sorin
Appearance in life unrecorded: BL c. 5.4 mm. On Chamaecyparis obtusa in Japan (Sorin 1995). Oviparae collected in
September were included in the original description. Matsuura & Yashiro
(2006) studied the behaviour of ants tending the eggs of this species. Stomaphis
japonica Takahashi Plate 15f Apterae are shiny brown, sometimes
wax-dusted; BL c. 5 mm. On trunks of Quercus
spp. in Japan (Takahashi 1960b; as S.
quercus japonica), China (Qiao & Zhang 1999d) and Korea (Seo 1994). Sexual
morphs occur in Japan in October-November. Takada (2008) found that sexual
reproduction in the region of Kyoto only took place on Q. serrata. He observed a partial migration to other Quercus spp. (acutissima, variabilis)
for the summer months, with alatae leaving these hosts in October-November.
Kamiju & Takada (1983) studied a parasitoid and hyperparasitoid of this
species. 2n = 10 (Blackman 1990). Stomaphis
(Parastomaphis) juglandis Petrović Apterae are elongate oval, shiny dark
brown, sometimes wax-dusted; BL 5.7-6.7 mm. On trunk of Juglans regia in Serbia (Petrović 1998). The original
description included alate viviparae collected in July and oviparae and
apterous males collected in October; the sexual morphs were redescribed by
Depa et al. (2015). Stomaphis
knechteli Binazzi & Blackman Colour in life unknown; BL of aptera
4.5-5.6 mm. On trunk of Acer campestre in
Rumania (Binazzi & Blackman 2004). An alata was collected in August. Depa
& Mróz (2013) suggested that this might be a geographical variant of S. graffii. Stomaphis
liquidambaris Takahashi Apterae are blackish; BL c. 5.2mm. On
stems of Liquidambar formosana in
Taiwan (Takahashi, 1925) and China (Ningxia; Qiao & Zhang 1999d). Sexual
morphs are unknown. Stomaphis
(Parastomaphis) longirostris (Fabricius) Apterae are
whitish powdered with greyish-white wax; BL 6.0-6.5 mm. On trunks of Populus spp. (nigra, tremula) and Salix spp. (alba, viminalis). There
is also a record of a population on Acer
campestre in Tuscany (Binazzi & Blackman 2004; but see Depa &
Mróz 2013). Tashev's (1961) record from Quercus
sp. in Bulgaria may be an error; however, the species resembling S. graffii that he found on Populus nigra is possibly longirostris. In continental Europe
(France, Netherlands, Germany, Austria, Czech republic, Slovakia, Italy,
Spain), Turkey and western Siberia. Depa et
al. (2015) described the sexual morphs, collected in September- October.
Depa & Mróz (2013) compared mitochondrial DNA and morphological
parameters of S. longirostris with
those of the closely-related European Acer-feeding
species (graffii, acquerinoi, knechteli), and concluded that the key characters used for
distinguishing species in this group were unreliable. Stomaphis
malloti Sorin
Apterae are dark brown; BL c. 5.2mm. On Mallotus japonicus in Japan. Sexual morphs occur in
October-November (Sorin 1979a, Takada 2009). Stomaphis
matsumotoi Sorin
Apterae are dark brown; BL c. 6.2 mm. On (trunk of?) Juglans mandshurica var. sachalinensis (= S. ailanthifolia) in Japan. Sexual morphs occur in October (Sorin
1995). Stomaphis
mordvilkoi Hille Ris Lambers
Apterae are dark brown; BL 5.3- 7.3mm. On trunk of Juglans regia; described from West Bengal,
India (Hille Ris Lambers 1933b) and subsequently rediscovered in northern
Italy (Colombo 1982). It is also now recorded from Hungary (Szathmáry et al. 2005). Sexual morphs are
unknown. Stomaphis
pini
Takahashi Colour in life unknown,
probably blackish; BL c. 4.5 mm. On basal part of trunk or roots of Pinus densiflora in Japan (Takahashi,
1960b; Inouye, 1970). There is also a record from P. halepensis in Algeria (Laamari et al. 2013; it needs to be confirmed that this is the same
species). Oviparae and apterous males collected in early December were
described by Sorin (1995). Apterae and an ovipara collected in late August on
Abies firma in Japan were described
by Sorin (2012a) as a subspecies, S. pini ssp. takaoensis. Stomaphis pistacicola Zhang
& Qiao Apterae are greyish brown;
BL c. 4.3 mm. On Pistacia sinensis
in China (Qiao & Zhang 1999d).
Other morphs and life cycle are unknown. Stomaphis
pterocaryae Sorin
Colour in life was not recorded; BL of aptera c.5.3 mm. On Pterocarya rhoifolia in Japan
(Honshu). Oviparae and apterous males
were collected in October (Sorin 2012a). Stomaphis
quercisucta Qiao & Zhang Apterae are elliptical, greyish white,
dusted with white powder; BL c. 5.2 mm. On Quercus sp. in Hebei, China (Qiao & Zhang 1999d). Apart from
the appearance in life the description does not indicate any clear
distinction from the western palaearctic species S. quercus. Stomaphis
quercus (Linnaeus)
Plate 15e Apterae are elliptical to elongate oval,
shining dark brown (see influential
points/Gallery); BL 5.3-7.0mm. In bark crevices on trunk of Quercus spp.,
also on Betula pendula and
occasionally Alnus glutinosa.
Throughout Europe, and west Siberia. Sexual morphs occur in
September-November; they were redescribed by Depa et al. (2015). Goidanich (1958) made a detailed study of its life
cycle and relation with the attendant ant Lasius
fuliginosus, and Lorenz & Scheurer (1998) made a comparable study of
populations on Betula pendula. In Sardinia it is associated with L. brunneus on Q. suber,
causing damage to cork bark (Loi et al.
2012). Depa
et al. (2012) confirmed using
mitochondrial DNA that the populations on Quercus
and Betula are conspecific, and
that S. betulae Mamontova,
described from Betula in the
Ukraine, is a synonym. 2n = 10. Stomaphis
radicicola Hille Ris Lambers Apterae are broadly oval, powdered with
grey dust; BL 4.5-6.0mm. On roots of Betula
sp., deep underground, attended by Lasius
umbratus,in the Netherlands (Hille Ris Lambers 1947b). Depa et al. (2013) have now described the
ovipara, collected in October on Alnus
glutinosa in Slovakia, and the male was described by Depa et al. (2015). This specimen was in a
deep bark crevice above ground level, attended by Myrmica rubra. Stomaphis
rhusivernicifluae Zhang Appearance in life is unknown; BL of
aptera c. 5.4mm. On Toxicodendron
vernicifluum (= Rhus verniciflua)
in Yunnan, China (Zhang & Zhong 1985e). Other morphs and biology are
unknown. Stomaphis
sinisalicis Zhang Apterae are without wax powder; BL c. 6.0
mm. On Salix matsudana, collected at Beijing, China (Zhang & Zhong 1982b);
subsequently recorded from Populus
sp., and from other parts of China (Qiao & Zhang 1999d). Other morphs and
biology are unknown. Stomaphis
takahashii Sorin
Appearance in life is not recorded; BL of aptera 4.7- 5.0 mm. On Acer spp. and Betula latifolia in Japan (Hokkaido). Sexual morphs occur in
September (Sorin 1965). Stomaphis ulmicola
Inouye Apterae are shiny brown-black;
BL c. 6.3mm. On trunk of Ulmus japonica
in Hokkaido, Japan (Inouye, 1938; Takahashi, 1960b) and also recorded
from Hebei, China (Qiao & Zhang
1999d). Sorin (1995) described oviparae and an apterous male collected in
October. Stomaphis wojciechowskii Depa Apterae are elliptical, pale fuscous to
pale brownish, dull, slightly powdered with wax; BL 6.1-6.8 mm. On basal part
of trunk of Quercus robur, at ground level or up to c.1.8m
above ground, in ant shelters constructed within bark crevices and covered by
soil (Depa et al. 2012). The
attendant ants are usually Lasius
brunneus, never L. fuliginosus
(cf. S. quercus). As yet it is only
known from southern Poland, but it has probably been overlooked and is much
more widely distributed. Sexual morphs occur in September to early November;
they were redescribed by Depa et al.
(2015). Depa & Mróz (2012b)
described the fundatrix, collected from Q.
robur in early June.. Stomaphis
yanonis Takahashi
Apterae are grey-brown with two rows of dark brown dorsal patches; BL
c. 5.0mm. In crevices on bark of trunk of Celtis
sinensis and Zelkova serrata,
in Japan, Korea, China and Taiwan. Takahashi (1960b) described a population
on Aesculus turbinata as a
subspecies, S. yanonis ssp. aesculi. Sexual morphs occur in
October-November (Sorin, 1965, Takada 2009). 2n = 20 (Honda 1921) or 16?
(Blackman 1990).
A
genus for one distinctive aphid related to Saltusaphis but with a dome-like protrusion of the front of the
head. The name is a replacement for Stenaphis
Quednau 1954 which is preoccupied by Stenaphis
del Guercio 1913 (Quednau in Favret et
al. 2007, p.397). Strenaphis elongata (Baker)
Apterae are very long-bodied, yellow to orange-yellow with paired dark
orange flecks on dorsal abdomen; BL 2.5-3.1 mm. An alata trapped in Canada
(New Brunswick) was illustrated by Quednau (2010). On Carex spp. (stricta, strictior) in eastern North America.
Oviparae (original description) and apterous males (Hottes & Frison 1931)
occur in October. 2n=10.
One
species living on Trifolium in
north-west Europe, with characters between Acyrthosiphon and Aulacorthum.
The peculiar habitat, the presence of
antesiphuncular sclerites in the aptera, and the unusual antennal
sensoriation of the alatae which have 2-14 rhinaria usually restricted to the
basal half of ANT III, are distinctive features. Subacyrthosiphon cryptobium Hille Ris Lambers Plate 24a (Fig.53c) Apterae are pale olive-green with head
sometimes faintly reddish; BL 1.6-2.3 mm. Living hidden on older parts of
prostrate stems of Trifolium repens,
and dropping when disturbed so that they are rarely observed or collected.
The plant shows no reaction. Monoecious holocyclic, with apterous males. In
Europe, and introduced to USA.
One
nearctic species related to Iziphya and
Saltusaphis but with exclusively mushroom-shaped
dorsal hairs and a distinctive, rounded and very hairy abdominal tergite 8. Subiziphya clauseni Quednau Apterae are long-bodied, colour in life
unknown but probably dark, with antennae black except at bases; BL c.1.8 mm.
Described from Poa sp., but the
true host must surely be Cyperaceae, probably a Carex species (hence inclusion in the key to sedge aphids). Alatae have broad dark dorsal abdominal
cross-bars, bordered wing veins with spots at their apices, and 15-17
secondary rhinaria on ANT III (see also Quednau 2010).
One
species on Lamiaceae in India resembling Capitophorus, with short capitate dorsal hairs and
smooth clavate siphunculi. Subovatomyzus leucosceptri A.N. Basu Apterae are spindle-shaped, pale
translucent green with a darker green spinal stripe, and whitish appendages;
BL 1.5-1.9 mm. Alatae have blackish head, thorax and antennae, a dark central
abdominal patch, and numerous secondary rhinaria on ANT III-V. On undersides
of leaves of various labiate species in several genera (Callicarpa, Colebrookia, Elsholtzia, Leucosceptrum, Pogostemon),
and also recorded from Vernonia (Compositae;
as Capitophorus vernoniae A.K.Ghosh
& Raychaudhuri), but these were possibly vagrants. Described from West
Bengal, India, where it is most common in April to June and reproduces
parthenogenetically throughout the
year (original description). Also
recorded from Sikkim and Meghalaya (A.K. Ghosh 1973), and from Nepal (BMNH
collection).
About 12 Carex-feeding
long-bodied aphids with spatulate empodial hairs, and with dorsal hairs
mostly very short and mushroom-shaped, with an incised “cap” so that they are
stellate in dorsal view. Apterae lack
secondary rhinaria. Alatae have a dark central abdominal patch on tergites
3-5, incised intersegmentally. Richards (1971) reviewed the world fauna, and
there are reviews of the European species by Stroyan (UK, 1977), Heie
(north-west Europe, 1982), and Nieto Nafría & Mier Durante (Iberian
peninsula, 1998) and of the Japanese species by Sorin (2005). Quednau (2010) reviewed
the world fauna, provided keys and
illustrated all available morphs. Subsaltusaphis aquatilis (Ossiannilsson)
Apterae are long-bodied, pale yellowish, with dark intersegmental
muscle sclerites; BL 2.1-2.6 mm. On Carex
spp. (aquatilis, acutiformis, riparia) in northern and central Europe. Alatae have 10-17 secondary rhinaria on ANT
III (Heikinheimo 1984). It is almost indistinguishable from the early
seasonal form of S. picta. 2n=8
(cf. picta). Subsaltusaphis canadensis Richards
Apterae are long bodied, pale yellow with black antennae; BL c. 2 mm.
On Carex sp. in Ontario,
Canada. Alatae have 8-12 secondary rhinaria
on ANT III. Apart from its small size and lack of dorsal pigmentation this
species closely resembles S. lambersi.
It could also be a variant form of S.
virginica (Quednau 2010). Subsaltusaphis flava (Hille Ris Lambers) (Fig.20c) Apterae are dull yellow, with dark
intersegmental muscle sclerites (no
longitudinal dark stripes), and antennae black except at bases; BL 1.8-2.1
mm. Alatae have 8-9 secondary rhinaria on ANT III. On Carex nigra in Europe. Oviparae occur in October in Denmark (Heie
1982), and the male genitalia were described and illustrated by Wieczorek
(2011). 2n=8. Subsaltusaphis intermedia (Hille Ris
Lambers) Apterae are yellowish white,
with rather indistinct dark intersegmental muscle sclerites, and antennae
black except at bases; BL 1.9-2.2 mm. Alatae have 9-12 secondary rhinaria on
ANT III. On Carex hirta, living
between the leaf bases on “dry
waysides”. In Europe (Netherlands, Austria, Germany, Czech Republic). Subsaltusaphis lambersi (Quednau) Apterae are pale
yellowish, with narrow brownish intersegmental muscle plates, and a pair of
brownish, sometimes rather vague and incomplete, pleural longitudinal
stripes; BL 2.4-2.8 mm. On Carex acuta in Sweden, apparently
preferring shady habitats and
waterside situations (Ossiannilsson 1959). It is also recorded from Finland
(Albrecht 2010), Poland (Achremowicz 1972) and Spain (Nieto Nafría & Mier
Durante 1998). Oviparae were found in Sweden in early September. Alatae are
undescribed. Similar aphids described from Carex ?reinii in
Japan are treated by Quednau (2010) as a subspecies, S. lambersi
kamijiensis Sorin. This Japanese form has alatae with 12-18 secondary
rhinaria on ANT III, and its oviparae were found in late January (Sorin
2005). 2n=6 for ssp. kamijiensis (Blackman 1980, erroneously listed as S. saracola). Subsaltusaphis
ornata (Theobald) (Fig.20e) Apterae are yellow to ochreous yellow with
two very distinct pleural longitudinal dark lines, clearly defining a pale
spinal band; BL c. 2.7-2.8 mm. Alatae have c.13 secondary rhinaria on ANT
III. On various Carex spp., with a
record also from Scirpus sylvaticus.
In Europe, Iran (BMNH collection), and now also recorded from Kazakhstan
(Kadyrbekov 2014e). 2n=8. Subsaltusaphis pallida (Hille Ris Lambers) (Fig.20b) Apterae are pale yellow, with dark
intersegmental muscle sclerites; BL c.2.2-2.7 mm. Alatae are unknown. On Carex
spp. (gracilis, stellulata) on river banks, and there is also a
record from Juncus (Shaposhnikov 1964). In Europe, Central Asia
((Kazakhstan) and east Siberia (Quednau & Shaposhnikov 1988). Subsaltusaphis paniceae (Quednau)
Apterae are pale yellow, with dark intersegmental muscle sclerites and
variably developed longitudinal dark markings similar to those of S.
picta (q.v.), and with antennae banded on segments III-V, VI being wholly
dark; BL 1.3-2.3 mm. On Carex spp.
in wet meadows in Europe (UK, Sweden, Poland, Germany, Italy, Spain). Very similar to the North American S. virginica. Subsaltusaphis picta (Hille Ris Lambers) Plate 4h Apterae are whitish yellow, with dark
transverse intersegmental muscle sclerites, and somewhat less dark and
variably developed longitudinal dark markings, the most heavily marked
specimens having dark marginal and pleural stripes and a shorter spinal stripe; antennae black
beyond basal half of segment III (see influentialpoints.com/Gallery);
BL 2.4-3.0 mm. Alatae have 11-16 secondary rhinaria on ANT III. On Carex spp. growing at the edge of
water, and sometimes also on Scirpus in
the same situation. Widely
distributed in Europe; also recorded from Kazakhstan, and east Siberia
(Quednau & Shaposhnikov, 1988). Oviparae and apterous males occur in
September-October in Sweden and Scotland (Heie 1982 and BMNH collection, leg.
H.L.G. Stroyan). 2n=10. Subsaltusaphis (Primoriaphis) primoriensis Quednau &
Shaposhnikov Apterae are
undescribed, alatae are pale yellow with blackish grey markings, and 4
rhinaria on ANT III; BL c 1.6 mm. On Carex
laevirostris in east Siberia. Subsaltusaphis rossneri (Börner) (Fig.20d) Apterae are ochreous yellow, usually with
dusky pleural longitudinal stripes on head and thorax, and paired segmental
patches plus a faint spinal stripe on abdomen; BL c.2.3-2.5 mm. Alatae have
10-13 secondary rhinaria on ANT III.
On Carex spp. (elata, nigra, rostrata, ?vesicaria) in Europe, and one alata is
recorded from Pakistan (Naumann-Etienne & Remaudière 1995). Subsaltusaphis (Primoriaphis) taoi Hsu ex Tao (= pulchra Quednau & Shaposhnikov; synonymy by Quednau 2010) Apterae are lemon yellow, slightly shiny,
with four longitudinal black stripes; BL c. 2.3mm. Alatae have 6-9 secondary
rhinaria on ANT III. On Carex sp.
In Taiwan, and on C. rubra in east
Siberia (Quednau & Shaposhnikov 1988). Sorin (2005) redescribed apterae,
alatae and oviparae from Japan, collected on C. dispalata. Subsaltusaphis virginica (Baker)
Apterae are pale yellow, with or without faint dark longitudinal
pleural and marginal stripes, and sometimes a very faint spinal stripe. The
antennae are pale basally and dark distally, with the base of segment IV
often paler than the distal part of
III but clearly darker than I and II. BL 1.9-2.9 mm. Alatae have 11-15
secondary rhinaria on ANT III. On Carex
spp. in North America (records mainly from eastern US and Canada, but
also from Utah) and Japan (as S. saracola
Higuchi). Oviparae were found in Illinois in early November, on both Carex and Scirpus eriophorum (= cyperinus);
BMNH collection, leg. T.H. Frison and H.H. Ross. [Normally this species has a
pair of long posterior marginal hairs on abdominal tergite 7, and the
posterior edge of this tergite projects laterally, as is typical for Subsaltusaphis. However, a collection
of specimens from North Carolina (BMNH collection., leg. D. Hille Ris
Lambers, no. 234) has no long marginal hairs on abdominal tergite 7, only
mushroom-shaped ones, and the posterior margin does not project
laterally. It is possible that two
species are involved, but no other distinguishing features between the two
forms were apparent.]
Two
bamboo-feeding species in India, resembling Takecallis in the anterior projection of the clypeus, but
differing in that the ANT PT/BASE ratio is only 0.24-0.55 and the tibiae are evidently
glandular. A.K. Ghosh & Quednau (1990) reviewed the genus and Quednau
(2003) illustrated all available morphs. Subtakecallis
brevisetosa Raychaudhuri & Pal All viviparae are alate, appearance in
life not recorded, probably wax-covered; BL 1.9-2.3 mm. On unidentified
bamboo in West Bengal, India (Raychaudhuri & Pal 1974). Sexual morphs are
unknown. Subtakecallis
pilosa (David, Rajasingh & Narayanan) All viviparae
are alate, covered in white powdery wax, with brown antennae, legs and dorsal
abdominal sclerites; BL 2.3-2.6 mm. On Bambusa
sp(p). in north-east India (Meghalaya, West Bengal). Oviparae and males were
described by Agarwala & Mahapatra (1990).
One
oriental species on Ulmaceae with flattened apterae having corrugated dorsal
cuticle and protruding spiracles. Sumatraphis celti
Takahashi Apterae are dull green (in
Sulawesi; J.H. Martin, pers. comm.) or yellow to deep brown or brownish green
(in India; A.K. Ghosh et al. 1971e,
A.K. Ghosh & Agarwala 1993); BL 1.5-1.7 mm. On new growth of Celtis tetrandra or ?Trema orientalis; the apterae are
flattened into angles of new shoots (J.H. Martin, pers. comm.). In India it
is also recorded from Mallotus sp.,
Quercus sp. and Sida sp., but these may be “casual”
visits. In Indonesia (Sulawesi, Sumatra), India
(Manipur, Meghalaya, Sikkim, Tamil Nadu, West Bengal), Nepal and China
(Yunnan). The alate vivipara was described by A.K. Ghosh & Agarwala
(1993), who also noted that S. celti
is active in India during the winter months. Sexual morphs are unknown. Sumatraphis tubercaudatus Qiao &
Zhang, described from Celtis sp. in
China (Qiao & Zhang 2000a), is probably a synonym.
One species in the Middle East apparently related to Aphis,
but with long thick dorsal hairs. A second species described in this genus
from China, S.
bambuciepula Zhang & Zhang is probably a Melanaphis, perhaps M. pahanensis. Swirskiaphis polychaeta Hille Ris Lambers
Plate 8e Colour of apterae in life is unknown; BL
1.9-2.6 mm. Alatae, described by Remaudière et al. (2006), have c.15-16 secondary rhinaria on ANT III. On Umbelliferae/Apiaceae
(Dorema, Ferula, Prangos) in the
Middle East (Iran, Lebanon, Turkey). The life cycle
is unknown.
About eight species of medium to large, rather
shiny dark brown aphids associated with Betula
and Alnus, forming colonies on the
branches and twigs, invariably attended by ants. All the apterae (even the
oviparae) are 'alatiform', with a similar pattern of sclerotization and
pigmentation to that of the fully-winged adults. Monoecious holocyclic; males
are apterous, and oviparae have the posterior abdominal segments extended
into an ovipositor-like structure. Qiao & Zhang (2002b) reviewed the
genus for China; they erected a subgenus, Antisymydobius,
which becomes a junior synonym of Yezocallis
Matsumura 1917, this having the same type species (S. kabae). Sorin (2014) reviewed and keyed the Japanese species. Trioxys betulae is a specialized
parasitoid. Symydobius alniarius
(Matsumura) Reddish or yellowish-brown,
with dark brown cross-bands and often with white powder on posterior
segments; BL 3.1-3.4 mm. On stems and twigs of Alnus spp. in Japan (Higuchi 1972), Korea and eastern Siberia
(Primorskiya Kray). The ovipara, collected in Japan in September, was
described by Sorin (2014), who also described a subspecies (nipponica), collected not only on Alnus spp. but also on Betula ermanii. 2n = 20. Symydobius americanus
Baker Shiny dark brown to black; BL
2.4-2.7 mm. On stems and twigs of Betula
papyrifera in northern USA and Canada. Sexual morphs occur in
September-October (Baker 1918). Symydobius intermedius Gillette
and Palmer (fig. 32B)
Olive
green to dark grey-brown; BL 2.2-2.6 mm.
On stems and twigs of Betula
occidentalis in western USA (Colorado, Utah, Idaho) and also recorded
from British Columbia. Sexual morphs occur in September-October (Gillette
& Palmer 1931). 2n = 16. Symydobius kabae
(Matsumura) (fig. 29C, fig. 32C)
Dark brown; BL 2.5-3.0mm. On bark of last
year's branches and twigs of Betula maximowicziana
and B. platyphylla var. japonica in Japan (Higuchi 1972); also
recorded from B. gmelini in
Mongolia (as alniarius - see Holman
and Szelegiewicz 1974), from B.
platyphylla var. latifolia and Alnus (hirsuta var. sibirica, japonica) in Korea and from B. dahurica in eastern Siberia
(Primorskiya Kray). Sorin (2014) described ovipara and apterous/alatiform
males, collected in Japan in late September-October. Paik (1972) recorded
oviparae in Korea in November. S.
paucisensorius Zhang & Zhang, described from Betula sp. in China (G. Zhang et
al. 1993a), is probably a synonym, and S. careofasciatus, described from apterae found on grasses (Qiao
& Zhang 2002b), are probably vagrant individuals of S kabae. 2n = 26? (Shinji
1931; but see Blackman 1986). Symydobius minutus
Quednau and Shaposhnikov
Yellowish-brown, with dark transverse bars and white wax powder
posteriorly; BL 2.2-2.4 mm. On shoots of Alnus
japonica in eastern Siberia (Primorskiya Kray) and Korea (Quednau &
Shaposhnikov 1988). Sexual morphs are unknown. Symydobius nanae Holman Brown to dark brown, with dark appendages
except for two pale rings on distal halves of antennae; BL not recorded,
probably 2-3 mm. In small aggregations on young branches of Betula nana; only known from apterous
viviparae, oviparae and males collected in September in Czech Republic
(Holman 1996). Symydobius oblongus
(von Heyden) (fig. 32D)
Shiny
dark brown (see
influentialpoints.com/Gallery); BL 2.0-3.5 mm. On twigs,
young stems and branches of Betula
spp. throughout Europe and across Asia. Sexual morphs occur in
October-November (Heie 1982). Holman (1990) studied developmental morphology
of fundatrices and their progeny, and Fowler & Macgarvin (1985) studied
the effects of ant attendance on population growth. See Heie (1982) and
Stroyan (1977) for general accounts. Two species described by Qiao &
Zhang (2002b) from Betula platyphylla in
China, S. fumus and S. brevicapillus, are close to S. oblongus and may be variant
populations of that species. 2n(female) = 15, 2n (male) = 14 (Blackman 1988). Symydobius quednaui Sorin Dark brown with blackish-brown dorsal
abdominal cross-bands, and dark brown antennae and legs; BL 2.5-3.4 mm. On Alnus
spp. in montane regions of Japan (Sorin 2014).
One species in east Asia apparently related to Aphis, but
apterae and alatae both have 4-segmented antennae, and apterae have a smooth
dorsal sclerotic shield. Szelegiewicziella chamaerhodi Holman
Apterae are green with greyish wax exudation; BL 1.2-1.45. Alatae have
just 1-3 secondary rhinaria on ANT III.
Described from specimens collected on flower stems of Chamaerhodis
erecta in August in Mongolia, and also recorded from eastern Kazakhstan
(Kadyrbekov 2009a). Similar aphids were collected on Artemisia vulgaris in June in China (Liaoning province; Zhang
& Zhong 1990). The dates of
collection do not support a host alternation from Chamaerhodes to Artemisia, so this possibility needs to be
tested experimentally. |