SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order) I
A genus for one distinctive
fern-feeding species. Idiopterus
nephrelepidis Davis Plate 19g (Fig.43i) Apterae are black with pale legs, pale
antennae ringed with black, siphunculi black basally and pale distally, and a
black cauda; BL 1.2-1.6 mm. Alatae have broadly black-bordered forewing veins
and a large white spot on the pterostigma.
On many genera and species of ferns
(Acrostichum, Adiantum, Asplenium, Anemia, Blechnum, Blotiella, Ceropteris, Dicranopteris, Dryopteris, Gymnocarpium, Nephrolepis, Pityrogramma, Polypodium, Polystichum, Pteridium,
Pteris). Records from other plants
are probably all vagrants. Perhaps neotropical in origin (Holman 1974), now
almost cosmopolitan, but confined to glasshouses and caves in northern
temperate regions. Apparently entirely anholocyclic. See Steffan (1962) and
Heie (1994) for further information. 2n=13.
About 45 North American species and one from the
Caucasus. Illinoia are pale spindle-shaped aphids with long appendages,
most easily recognised by the long weakly to moderately clavate siphunculi,
similar to those of Amphorophora but ornamented on the distal,
constricted region with a few rows
of polygonal reticulation. There is no host alternation.
Evolution of host relationships within the genus is somewhat obscure; many
species in the two principal subgeneric groups are associated with Ericaceae,
but others feed on taxonomically diverse plants, including some on trees. The species from cultivated Rhododendron mostly bear 5 hairs on the first tarsal
segments. The subgenus Masonaphis contains about 8 species, 6
of them on Ericaceae in North America.
Illinoia s.str. contains
about 30 species, 9 of them (including the Russian one) associated with
Ericaceae, with 6 of the species known from eastern North America. Seven species of Illinoia s.str. and one Masonaphis
live on Compositae. The 3‑4
species of subgenus Oestlundia are
associated with Rubus and the 2
subspecies of Amphorinophora live
on Lonicera. MacGillivray (1958) revised Illinoia (= Ericobium) using the name Masonaphis
for the full genus. Blackman (2010) reviewed and keyed the 5
species introduced to Europe from North America. Illinoia
alni
(Mason) Apterae are spindle-shaped,
pale green with a dark green median stripe and dark spots at bases of
siphunculi; BL 2.5-3.1 mm. Immatures
are pale green. In small colonies on
undersides of leaves of Alnus rugosa
in north-eastern USA and eastern Canada (MacGillivray 1958). Oviparae were collected in late August
. 2n=10. Illinoia
andromedae (MacGillivray) Apterae are
shiny bright green with darker appendages, BL 1.9-2.3 mm. Specimens beaten
from mixed vegetation of Andromeda glaucophylla and Vaccinium myrtilloides fed
on leaves of A. glaucophylla,
which is thus the only confirmed host plant.
Only known from apterae collected in Quebec, Canada.
Illinoia
azaleae (Mason) (Fig.46a) Apterae are rather shiny deep green; BL
1.9-2.7 mm. Immatures
are pale greyish green. Alatae have
quite dark wing-veins. On young stems and leaves of Rhododendron spp. (especially azaleas); also sometimes on other
Ericaceae (Andromeda, Leucothoe, Pieris, Vaccinium) and
occasionally on some other plants (Comptonia,
Viola, Tulipa). In eastern North America, and introduced to Europe,
South Africa, Hawaii, Australia, New Zealand and Argentina (Ortego
et al. 2004). Probably mainly anholocyclic; oviparae sometimes occur, but males are
undescribed. In colder climates it is an indoor or glasshouse pest of potted
azaleas and can cause defoliation. A form described by Tissot & Pepper
(1944) from Kalmia latifolia in Pennsylvania
was regarded by MacGillivray (1958) as a subspecies, Illinoia azaleae ssp. kalmiaflora.
It differs in the ratio of R IV+V to HT II from I. azaleae on Rhododendron (1.6-1.7
in kalmiaflora, 1.1-1.45 in azaleae), and is probably a good
species. On the other hand, Amphorophora
rhododendronia Mason, regarded as a subspecies of azaleae by MacGillivray, should probably be treated as a synonym.
2n=10. Illinoia
borealis (Mason)
Colour in life is unknown, probably dark green or brownish green with
dark siphunculi; BL 1.7-2.5 mm. On new growth of Ericaceae (Vaccinium, Gaultheria),
and also recorded from Myrica
aspleniifolia (= Comptonia
peregrina) (MacGillivray
1958). In north-eastern USA and Canada
(Quebec). Oviparae
and apterous males are recorded from Quebec early in August (aphidtrek.org).
Specimens in the BMNH collection from Clintonia identified as borealis seem to be pallida (q.v.). Illinoia
brevitarsis (Gillette
& Palmer) Apterae are pale
yellowish green to yellow; BL 2.3-3.2 mm. On leaves of Clematis ligusticifolia in western USA. Monoecious holocyclic,
alate males and oviparae occurring in late September-October (Palmer 1952).
[In the BMNH collection there is an undescribed Macrosiphum sp. from the same host in Oregon (leg. D. Hille Ris
Lambers, no. T85), distinguishable by the characters given in the key.] Illinoia
canadensis (MacGillivray)
Apterae are rather shiny, yellowish green mid-dorsally, darker green
laterally with dark spots at the bases of the dark-tipped siphunculi; BL c.
2.7 mm. On Myrica sp. in New Brunswick, Canada (MacGillivray 1958). Life cycle is unknown. Illinoia
ceanothi (Bartholomew) Apterae are green, BL 2.0-2.5 mm. On
leaves of Ceanothus spp. in western
USA (California, Oregon, Idaho, Washington; aphidtrek.org).
Alatae have 5-9 secondary rhinaria on basal part of ANT III (BMNH collection,
leg. R.C. Dickson and H.L.G. Stroyan). Monoecious holocyclic, with oviparae
and apterous males in late August to October (BMNH collection, leg. R.C.
Dickson). Illinoia
corylina (Davidson) Apterae are green with dark-tipped siphunculi:
BL 2.3-2.5 mm. Apterae have 2-4 secondary rhinaria on ANT III, alatae have
23-28. Described from Corylus rostrata (= cornuta var. californica), and
perhaps monoecious holocyclic on this host, as there are photographs by A. Jensen
of a fundatrix and an aptera on C.
cornuta at http://www.flickr.com/photos/sandnine/
(and see also aphidtrek.org).
However Davidson (cited in Mason 1925) stated that it was more common on
“ninebark” (Physocarpus capitatus).
In western USA (California, Oregon, Washington). [In the BMNH collection
(leg. D. Voegtlin) there are 2 apterae and 2 immatures that run to I. corylina in MacGillivray’s (1958)
key, but collected from Aquilegia
formosa. A. Jensen has collected an unidentified Illinoia from Aquilegia in
Washington and British Columbia (aphidtrek.org),
so perhaps there is another similar but undescribed species of Illinoia on this host.] Illinoia (Amphorinophora) crystleae (Smith &
Knowlton) Apterae are pale whitish
green, with pale appendages, dark only on tarsi and R IV+V; BL 3.7-5.0 mm. On
leaves of Lonicera involucrata in
western North America. Oviparae and alate males occur in August-September
(MacGillivray 1958, Palmer 1952). A form described by Essig on L. ledebourii in California (as Amphorophora bartholomewi) is regarded
as a subspecies, I. crystleae ssp. bartholomewi. 2n=16*. Illinoia (Oestlundia) davidsoni (Mason) Plate 25d Apterae are pale green, slightly waxy, with
brown-black siphunculi; BL 3.3-4.3 mm. On leaf petioles and undersides of
leaves of Rubus parviflorus in
western North America. Amphorophora arnicae Glendenning is
regarded as a synonym, with host alternation between Rubus and Arnica spp.
suspected (MacGillivray 1958), but this has still not been confirmed
experimentally. The collection data for the BMNH specimens suggests that
there may be two similar species on Arnica,
one migrating to Rubus and the
other not. 2n=12. Illinoia
dzhibladzeae Shaposhnikov Apterae are yellowish brown, shining
brown dorsally, siphunculi paler with black apices; BL 1.9-2.5 mm. On
undersides of leaves and on young shoots, flowers and fruits of Rhododendron flavum and Rhododendron sp. in Ukraine, Georgia
and Turkey (BMNH collection, leg. N. Tuatay). Illinoia
finni (MacGillivray) Apterae are undescribed, but some large
greenish white apt. collected on Vaccinium
sp. in Idaho (BMNH collection, leg. J.H. Martin) could be this species;
BL of these apterae is 3.3-3.4 mm. BL of alate paratypes is 2.2-2.5 mm (2
specimens), collected on “huckleberry” (Gaylussacia
or Vaccinium; we have included
it under both these genera) in Ohio, USA. This species is similar to I. pepperi (MacGillivray 1958). Illinoia
goldmaryae (Knowlton) (Fig.49a) Apterae are bright green (Essig 1942, as Amphorophora patchiae); BL 1.8-2.9 mm.
Forming dense colonies on young growth of species of Aster, Conyza, Erigeron and Solidago (MacGillivray 1958), and there are also records from
other genera of composite plants.
Widespread in USA and eastern Canada, also in Central America (Panama;
Quiros et al. 2009) and South
America (Peru, Argentina; Ortego & Nieto Nafría 2017). Also introduced to
England, where it was common in the London area on cultivated Michaelmas
daisies and golden rods in 1960-62, and has since been recorded from suction
traps (Blackman 2010). It has also
been reported from the Azores (Borges et
al. 2010), and a trapped alata is recorded from Tunisia (Boukhris-Bouhachem et al.
2007). Monoecious holocyclic in USA, with fundatrices on Solidago in April and males in September (aphidtrek.org);
the population introduced to England was apparently anholocyclic. Illinoia
gracilicornis (MacGillivray) Apterae (oviparae) are reddish with
“delicate glassy extremities”; BL 1.7-2.0 mm. On Cercocarpus montanus in Colorado, USA. Only oviparae are known. Illinoia
grindeliae (Williams) Apterae are greyish green, siphunculi
pale with dusky apices; BL 1.6-2.5 mm. Alatae may have 1-2 secondary rhinaria
on ANT IV and V, as well as 17-21 on III. On Grindelia squarrosa, and possibly specific to this host, in
western and central USA and Canada. MacGillivray (1958) differentiated some
samples from Colorado and Utah as a subspecies (palmerae), giving the host as Grindelia
sp., but paratypes that she donated to the BMNH collection are labelled
as collected from G. squarrosa, and
specimens subsequently collected do not provide evidence of any correlation
between morphology and geographical distribution or host plant. Illinoia lambersi
(MacGillivray) (Fig.46b) Apterae and alatae exist as
three distinct colour morphs, green, pink and yellow, which can often occur
in mixed populations (see influentialpoints.com/Gallery);
BL 2.0-3.7 mm. Alatae have 21-30 secondary rhinaria on ANT III. On
young leaves, shoots and flowers of Rhododendron
spp., including deciduous varieties. The principal
host is Rhododendron, but colonies
sometimes occur on Ilex aquilifolium
(records from British Columbia, Canada, Forbes & Chan 1989, and England,
BMNH collection, leg. V.F. Eastop). In
western North America, and introduced into Europe in about 1970 (Stroyan
1972b, Hille Ris Lambers 1973a), South America (Chile, Fuentes-Contreras et al. 1997), and Japan (Morimoto
& Kiritani 1995). Holocyclic on Rhododendron in California, USA, but
European populations are anholocyclic (Hille Ris Lambers 1973a). 2n=10. Illinoia liriodendri
(Monell) Apterae are spindle-shaped,
pale green lightly dusted with wax, with antennae and siphunculi black except
at bases, and legs pale green except for black tibial apices and tarsi (see influentialpoints.com/Gallery);
BL 1.7-2.5 mm. A red colour form also
occurs. Monoecious holocyclic on Liriodendron tulipifera in North
America, feeding on undersides of leaves. Introduced to Japan in 1991
(Sugimoto 1999), more recently to France (1998), Italy (2001), UK and Germany
(2004), and Korea (Kim et al.
2011). It is also now recorded from Luxembourg, Slovenia, Poland, Hungary
(Bozsik 2012), Serbia, Greece and Egypt. Oviparae and alate males occur in
October. Davis (1909) gave a full description,
Dreistadt (1987) studied honeydew production, Zuparko & Dahlsten (1993)
surveyed its parasitoids in California, and Dreistadt & Dahlsten (1988)
discussed the timing of control measures.
2n=10. Illinoia
macgillivrayae (Hille Ris
Lambers) Apterae are pale yellowish
green; BL 2.6-3.2 mm. Alatae have c.7-9 secondary rhinaria on ANT III (1
specimen, BMNH collection). In small colonies on young leaves of Raphiolepis indica, and subsequently
found on other Rosaceae (Chaenomeles
japonica, Pyrus kawakamii, Holodiscus
discolor and Spiraea sp.), as
well as two samples from Corylus
cornuta (BMNH collection, leg. C.-k. Chan, and aphidtrek.org).
In western USA (California, Oregon) and Canada (British Columbia). It is very
similar to I. spiraeae, but larger
and with a relatively longer R IV+V. Illinoia
masoni (Knowlton) Apterae are bluish green, with pale
appendages; BL 1.8-2.8 mm. On leaves of Helianthus
spp. in western USA (Palmer 1952). In laboratory tests, Rogers &
Thompson (1978) found various degrees of resistance to this species in North
American perennial species of Helianthus. Illinoia
(Oestlundia) maxima (Mason) Apterae are white
with black siphunculi (unpublished
observations, R.L. Blackman); BL 2.9-3.9 mm. Alatae have a dark pterostigma
and a dark spot at tip of forewing. On leaves of Rubus parviflorus in western North America (British Columbia to
California). Monoecious holocyclic with an abbreviated life cycle, oviparae
and alate males being produced in the 3rd or 4th
generation from late May (California) to July (Frazer & Forbes 1968).
Population dynamics of this aphid (Gilbert 1980) and its interactions with a
parasitoid (Gilbert & Gutierrez 1973) have been studied intensively.
2n=12. Illinoia (Masonaphis) menziesiae Robinson Apterae are grey-green to salmon pink (see http://www.flickr.com/photos/sandnine/page3/); BL 2.0-2.5 mm. Alatae are undescribed. On undersides of leaves of Menziesia ferruginea, where it causes a yellowing and downward curling of apical portions of leaves. In western USA (Idaho, Montana, Washington, A. Jensen, aphidtrek.org) and Canada (Alberta, original description, and British Columbia, BMNH collection, leg. C.-k. Chan.) Monoecious holocyclic, with oviparae and apterous males collected in early September-October. (The siphunculi are unusual for this genus in having little or no distal swelling. The specimens described by Robinson are probably fundatrices.) 2n=10*. Illinoia
morrisoni (Swain)
Apterae are spindle-shaped, long-legged, rather dark apple green, with
antennae dark beyond the base of ANT III, and legs and siphunculi tipped with
black (see influential
points/Gallery); BL 1.5-2.3 mm. On
terminal leaves and shoots of numerous conifers, usually Cupressaceae (Actinostrobus, Callitris, Calocedrus, Chamaecyparis, Cupressocyparis, Cupressus,
Juniperus, Thuja, Widdringtonia);
also on Araucariaceae (Araucaria)
and Taxodiaceae (Metasequoia, Sequoia, Taxodium). There is only
one record from Pinaceae (Cedrus
deodora; BMNH collection, leg. H.G. Walker). In western North America, Panama (Quirós et al. 2009), Costa Rica and Venezuela
(Villalobos Muller et al. 2010),
and introduced into England, Wales and France (Rabasse et al. 2005b). Alatae have also been trapped in northern Italy,
but apparently it has not spread more widely.
Life cycle is unknown; no sexual morphs are described and, in
California at least, it seems to be anholocyclic. 2n=10. Illinoia
pallida (Mason) Colour of apterae in life is unrecorded;
BL 1.7-2.6 mm. Alatae are undescribed. On Clintonia
borealis in north-eastern USA and Canada (Maine, New Brunswick). The
aptera was redescribed by MacGillivray (1958). Illinoia (Masonaphis) paqueti (MacGillivray) (Fig.46c) Apterae are whitish, yellowish or reddish,
wax-dusted, with black-tipped siphunculi; BL 3.1-4.0 mm. On felty undersides
of old senescing leaves of Rhododendron
(=Ledum) groenlandicum in Canada (Quebec, Labrador) and Alaska (BMNH
collection). Monoecious holocyclic, with oviparae and alate males in August
(MacGillivray 1958). Illinoia patriciae
(Robinson) Apterae are spindle-shaped,
yellowish green to green; BL 1.7-2.3 mm.
On new needles of Tsuga
heterophylla in British Columbia, Canada (Robinson 1969) and Oregon, USA
(aphidtrek.org). Differences from the one other conifer-feeding
Illinoia species, I. morrisoni (which has not been
recorded from Tsuga) are given by
Robinson. The life cycle is not known.
Illinoia
pepperi (MacGillivray) (Fig.56c,d) Apterae are green or red, with rather dark
antennae, legs and siphunculi; BL 1.7-2.8 mm. On Vaccinium spp. in north-eastern North America. Monoecious
holocyclic with oviparae and alate males in October (MacGillivray 1958).
Elsner & Kriegel (1989) studied distribution and seasonal phenology in
Michigan. 2n=28. Illinoia
phacelia (Essig) Apterae are green with dark-tipped
appendages; BL c. 2.5-2.7 mm. On undersides of leaves of Phacelia sp. in western USA (California, Oregon). The alata was
redescribed by MacGillivray (1958). Illinoia (Masonaphis) pinawae (Robinson) (Fig.46e) Apterae are pale yellowish green; BL
1.9-2.5 mm. Alatae are undescribed. On Rhododendron
(=Ledum) groenlandicum in Canada (Manitoba, Ontario). Illinoia (Masonaphis) rhododendri (Wilson) (Fig.46b,d) Apterae are green or pink, sometimes with
dorsal cuticle sclerotic and rather smoky; BL 2.4-2.8 mm. On Rhododendron sp(p) in north-western
USA. The life cycle is uncertain but may be abnormal, with fundatrices in
late July, and possible early production of sexuales (MacGillivray 1958). Illinoia (Masonaphis) rhokalaza (Tissot
& Pepper) Apterae are light yellowish
brown, sometimes with greenish tinge, with dark brown to antennae, legs and
siphunculi; BL 2.2-2.9 mm. On Rhododendron
spp., Leucothoe fontanesiana
and Kalmia latifolia in eastern USA
(MacGillivray 1958). Illinoia
richardsi (MacGillivray) Apterae are
white with pale appendages, with only apices of antennal segments, apices of
tibiae and tarsi darker to blackish; BL 2.6-3.2 mm. Alatae are also very pale but have brownish
head, thorax, antennae and legs, and rather dark wing veins. On undersides of
leaves and in inflorescences of Anaphalis
margaritacea and Gnaphalium bicolor (= Pseudognaphalium bioletti);
often found in the inflorescences with Uroleucon russellae.
Widely distributed in USA and Canada. Hille Ris Lambers (1966) described a
form in California with thicker and less swollen siphunculi as a subspecies, I. richardsi ssp. pacifica.
Sexual morphs and life cycle unknown.
2n=10.
Illinoia (Oestlundia) rubicola
(Oestlund) Apterae are greenish
yellow, often with a broad dark green spinal stripe; BL 3.1-4.6 mm. Alatae
have a dark pterostigma and a large black spot at the tip of each forewing.
On stems of Rubus idaeus var. strigosus, and occasionally on other closely-related Rubus spp., widely distributed in North America. Monoecious
holocyclic, with oviparae and alate males in October. 2n=12. Illinoia
simpsoni (MacGillivray) Colour of apterae in life is unknown; BL
1.8-2.1 mm. Alatae are undescribed. On Linnaea
borealis in Maine, USA. Illinoia
spiraeae (MacGillivray)
Colour of apterae in life is
unknown; BL 2.7-2.9 mm. Alatae have not been described. On Spiraea spp. in north-western USA (Oregon, Idaho) and British Columbia, Canada,
and there are also samples in the BMNH collection from certain other woody
Rosaceae, including Photinia davidiana,
and also from plant genera in other families (Deutzia, Rhus, Salix, Yucca) indicating that this species has polyphagous tendencies.
2n=10. Illinoia
spiraecola (Patch) Apterae are various hues of red, green
and yellow; BL 2.5-3.2 mm. On Spiraea spp.,
widely distributed in North America. Specimens identified as this species
have also been collected from undersides of leaves of a tall Thalictrum sp. It was redescribed by
MacGillivray (1958). Illinoia
subviride (MacDougall) Apterae are delicate green, similar in
colour to hairs of the host plant. On Aster
alpinus in Washington, USA. 2n=10. Illinoia
thalictri (MacGillivray) Apterae are very pale sea-green to
yellowish green, lightly wax-dusted; BL 2.0-2.3 mm. On leaves and petioles of
an unidentified Thalictrum sp. in
Colorado, USA, and recorded from T.
fendleri in New Mexico (Miller et
al. 2016); a photograph of an Illinoia
sp. on Thalictrum (http://www.flickr.com/photos/sandnine/)
is probably this species. Monoecious holocyclic with oviparae and alate males
in September (MacGillivray 1958). Illinoia
wahnaga (Hottes) Apterae are pale whitish green to milky
white, with black eyes; BL 2.2-2.6 mm. Living singly on undersides of leaves
of plants in genera formerly placed in
Convallariaceae (Convallaria, Maianthemum) in North America
(Colorado, British Columbia, Manitoba). Oviparae and apterous males occur in
September-October in Manitoba (Robinson 1965). 2n=10. Illinoia
wilhelminae (Hille Ris Lambers) Apterae are spindle-shaped, colour in life
is not recorded; BL c. 3.1 mm. The
siphunculi show little or no trace of swelling. Described from a single aptera collected in
Quebec, Canada, on Alnus rugosa,
but the form of the last rostral segment suggests that this was not its
normal host plant (Hille Ris Lambers 1962).
It does not seem to have been recorded since.
A genus which possibly has host-alternation between Smilax and Impatiens in Asia, although this needs further
confirmation; the two species that have spread to Europe produce their sexual
morphs on Impatiens.
Five species living on Cuphea in
America (Mexico, Panama and Brazil) have also been placed in this genus.
Reviews are available for Europe (Heie, 1994), Japan (Miyazaki, 1971), Korea
(Lee & Havelka, 2001a) and America (Nieto Nafría et al. 2012).
Impatientinum
americanum Remaudière Apterae have a shining black dorsum,
antennae dark except for basal part of ANT III, and dark siphunculi; BL
1.5-2.6 mm. On Cuphea aequipetala and Cuphea sp. in Mexico. Alatae were described by Nieto
Nafría et al. (2012).
The record from Panama (Quiros et al.
2009) should be referred to I.
chiriquense.
Impatientinum
asiaticum Nevsky Plate 25f (Fig 33a,c) Apterae are bright green, pink or red,
with an extensive shiny brown-black dorsal shield, black siphunculi and
yellow to whitish cauda (see influential
points/Gallery); BL 2.1-3.1
mm. On undersides of leaves along main veins, and later in dense colonies on
flowerstalks, of Impatiens spp.,
especially I. parviflora. Probably
originally in Central Asia (Tien Shan), now widely distributed in Europe
(Holman 1971, Heie 1994) and Central Asia (Kadyrbekov 2017a). A subspecies, I. asiaticum ssp. dalhousiense, was described from Smilax parvifolia (= elegans) in north-west India (Verma 1969, as I. impatiensae dalhousiensis). I.
asiaticum s.str. is monoecious holocyclic on Impatiens in Europe (with alate males), but the north Indian
subspecies may be heteroecious holocyclic between Smilax and Impatiens.
2n=16. Impatientinum
balsamines (Kaltenbach) Apterae are
green with an extensive shiny black dorsal shield, black siphunculi and a
pale cauda; BL 2.0-2.7 mm. Singly or in small colonies on undersides of
leaves of Impatiens spp.
but especially associated with I.
noli-tangere. Probably east Asian in origin, now widely distributed in Europe and
Asia. Monoecious holocyclic with alate males. 2n=16.
Impatientinum
chiriquense Nieto Nafría
& Mier Durante Apterae probably
have a shiny black dorsum in life, and dark antennae and siphunculi; BL
1.7-2.0 mm. Apterae and alatae were collected in May on Cuphea infundibulum (= appendiculata) in Panama (Nieto Nafría et al. 2012). Other morphs are
undescribed. Impatientinum
holmani Peña-Martínez & Nieto
Nafría Apterae probably have a shiny
black or dark brown dorsum in life with dark siphunculi, but antennae mainly
pale; BL 1.7-2.3 mm. Monoecious holocyclic on Cuphea sp. in Mexico, with alate males collected in October
(Nieto Nafría et al. 2012). Impatientinum
impatiens (Shinji) (Fig.33b,d) Apterae are yellowish green to bluish
green or orange-red with an extensive shiny black dorsal shield, black siphucunli
and a pale cauda; BL 2.4-3.5 mm. On Impatiens
spp. (especially I. textori) in
Asia (north India, east Siberia, Japan, Korea, Taiwan, Laos). Fundatrices
were described from Smilax china in
Japan (Miyazaki 1971), and it has also been collected from this host in Laos
(Yoshitomi et al. 2014). There is
presumably a host alternation between Smilax
(primary host) and Impatiens,
but this needs to be confirmed experimentally. Macrosiphum smilaceti Takahashi is probably a synonym. 2n=16. Impatientinum
oaxacense Nieto Nafría & Pérez Hidalgo Apterae probably have a shining black
dorsum in life, with dark siphunculi, the antennae and legs mainly pale; BL
c.2.3 mm. Only known from the holotype aptera, collected in August on Cuphea sp. in Mexico (Nieto Nafría et al. 2012).
Impatientinum paranaense de Carvalho, Cardoso & Lazzari Apterae are green with abdomen olive green, antennae, legs and siphunculi dark
brown, and cauda green; BL 1.5-2.0 mm. Alatae have 15-24 secondary rhinaria
distributed over the length of ANT III. On flower stems of Cuphea
calophylla in the state of Paraná,
Brazil (de Carvalho et al. 2004). The life cycle is unknown. Impatientinum
(Neoimpatientinum) smilaceti Agarwala,
Mondal & Raychaudhuri Colour of apterae
in life is not recorded, presumably green or reddish with broad dark marginal
longitudinal bands, black siphunculi and a pale cauda; BL 2.8-3.0 mm. On Smilax macrophylla in Sikkim, India.
[This is a secondary homonym of Macrosiphum
smilaceti Takahashi, if that species is I. impatiens.]
A genus erected for one
gall-forming Middle-Eastern species originally described in Geoica, distinguished by the very large
single rhinaria on each of ANT III-VI in the emigrant alatae (Barjadze et al. 2018c). Inbaria swirskii (Remaudière & Inbar) Galls
on Pistacia atlantica are of
irregular shape, bulbous, c. 25 mm in diameter, arising from leaf stem
between bases of leaflets. Apterous fundatrigeniae and emigrant alatae were
collected from the galls in October (Remaudière et al. 2004). Secondary hosts and other life cycle stages are
unknown. Only known from Jordan.
Four Indian species on
Ericaceae, closely related to Ericolophium,
and possibly not warranting a separate
genus. A.K. Ghosh (1991) provided a key to species. Alatae have dark
dorsal abdominal cross bands or a quadrate patch. Indiaphis
crassicornis A.N. Basu Plate 23g (Fig.46h) Apterae are pale glassy white, dusted with
wax; BL 1.7-2.0 mm. Alatae have secondary rhinaria distributed III 21-40, IV
0-8. On Rhododendron spp. in India (Punjab, West Bengal). Alate males
were collected in Punjab in November (BMNH collection, leg. Commonwealth
Institute of Biological Control). Indiaphis
indica (L.K. Ghosh, Verma & Raychaudhuri) Colour of apterae in life is unrecorded,
probably pale with dark-tipped siphunculi; BL 1.4-2.4 mm. Alatae have
secondary rhinaria distributed III 44-56, IV 1-7, V 0-2. On an
unidentified Rhododendron sp.
in India (Himachal Pradesh, Uttar Pradesh).
Indiaphis rostrata
A.K. Ghosh &
Raychaudhuri Apterae are pale
yellow; BL 1.8-2.0 mm. On undersides of leaves of an unidentified Rhododendron sp. in India (Sikkim).
Only known from apterae; these were collected in March and may be
fundatrices, which would explain the 5-segmented antennae with a short
terminal process. (However they are probably not fundatrices of I. crassicornis, because of the long R
IV+V.) Indiaphis
setosa (Hille Ris Lambers & A.N.
Basu) Apterae are green to dark
green, with brown legs and dark brown siphunculi; BL 1.6-2.2 mm. Alatae have
secondary rhinaria distributed III 8-12, IV 0. On Agapetes (= Pentapterygium) serpens, often in rather large
colonies on tender parts of shoots and sometimes on undersides of leaves. In
West Bengal, India.
One
elm-feeding species related to Chromocallis,
but with shorter antennal hairs, unbordered forewing veins and a heavily
sclerotised apterous morph (see Quednau 2003) Indiochaitophorus
furcatus Verma
Apterae are rather elongate oval, pale brown with thick black dorsal
longitudinal stripes; BL 1.3-2.0 mm. Alatae have paired dark spinal and
marginal sclerites on each of abdominal tergites 1-6. Recorded from
undersides of leaves and twigs of Ulmus
wallichiana in India (Kashmir,
Himachal Pradesh), and on U. villosa
in Pakistan (BMNH collection, leg. M.A. Ghani). Heavy infestation of
riverside trees was reported in Kashmir in June (Verma 1970a). A record from
Algeria (Laamari et al. 2013) requires
additional confirmation. The life cycle is unknown.
One east Asian species on Geranium. The species was transferred
to Neotoxoptera by Naumann-Etienne
& Remaudière (1995), but the unusual nature of the spiracular plates and
the dorsal hairs, taken together with the wing venation of the alatae, seems
to warrant the reinstatement of the genus. Indoidiopterus
geranii (Chowdhuri, R.C. Basu,
Chakrabarti & Raychaudhuri) (Fig.30e,f) Colour of apterae in life is unrecorded;
BL 1.6-1.8 mm. The alata was described by Chakrabarti et al. (1972). On Geranium spp. in north-west India and
Pakistan. Alate males were trapped in Pakistan (Naumann-Etienne &
Remaudière 1995, as Neotoxoptera
geranii). 2n=12.
Three species in north India
and Pakistan, at least two of which have host alternation between Rhododendron and
Compositae/Asteraceae. They have characters (e.g. spinulose hind tarsi,
alatae with numerous secondary rhinaria) shared with other members of a group
of Asian genera associated with Rhododendron
(Chaetomyzus, Ericolophium, Indiaphis). Indomasonaphis
anaphalidis (A.N.
Basu) (Fig.34a, Fig.46g) Apterae are elongate oval, rather shiny
pale green to lemon yellow, BL 3.4-4.8 mm. Alatae have variably developed
black dorsal abdominal patch. Males are alate. Originally described from Anaphalis triplinervis in West Bengal,
feeding on tender shoots and leaves without causing noticeable damage, and
since recorded from various other Compositae/Asteraceae (Inula, Ageratum, Gerbera,
Senecio, Artemisia), as well as some unlikely hosts (Eurya japonica, Morus alba,
Oxalis corniculata). Heteroecious holocyclic with a sexual phase on Rhododendron spp. in northern India
(Verma 1971, as Indomasonaphis indica;
Chakrabarti et al. (1983) as Indumasonaphis tuberculatus) and
Pakistan (Naumann-Etienne & Remaudière 1995). Indomasonaphis
inulae (A.K.Ghosh &
Raychaudhuri) Apterae are
elongate-bodied, with colour in life unrecorded, probably pale; BL 2.8-3.4
mm. On Rhododendron sp. and Inula
cappa in northern India. Host alternation between these two plants (with Rhododendron as the primary host) is
believed to occur (Chakrabarti et al.
1983, Chakrabarti & Banerjee 1993a). 2n=32 (Kurl & Chauhan 1988). Indomasonaphis
rumicis (Chakrabarti &
Raychaudhuri) Apterae are green to
brown; BL c.3.8-3.9 mm. Alatae have dark dorsal abdominal cross-bands. On
undersides of leaves of an unidentified Rumex
sp., and (as 2 alate males) on Oxyria
digyna, in northern India (Uttar Pradesh). Also collected (as immature
alatae) on Gerbera (assuming that “Garbera” was a typographic error),
which is a more probable host for a species in this genus. Alate males were
found in October (original description).
Four species in east Asia with dark, somewhat
swollen siphunculi reticulated at apices. Two species have Staphyleaceae as
primary hosts. Of the other two species one was described from an unidentified plant, and the other is only
known from alatae collected on a shrub that was subsequently identified as Corchorus sp. and unlikely to be the
true host. G. Zhang & Qiao (1998b)
reviewed the genus in China. Indomegoura indica (van der Goot)
Plate 20i Apterae are orange yellow, covered in
white wax, with stout, dark siphunculi; BL
3.1-4.2 mm. On Staphylea bumalda
and Euscaphis japonica in Japan,
and recorded as common in the Tokyo area, feeding on the undersides of the
branches, which may be severely affected (Takahashi 1923, Miyazaki 1971).
Populations in summer occur on Hemerocallis
spp., but there is some uncertainty about the life cycle as Takahashi
(1923), in describing the biology, recorded apterous sexuparae. and both
apterous and alate males on the primary host, indicating that only part of
the population may migrate. In Japan,
Korea, China, Taiwan and India. 2n=10. Indomegoura nigrotibiae (Tao)
Colour of apterae in life is unknown; BL 2.2-2.5 mm. On Staphylea bumalda (Miyazaki 1971). In
Taiwan, Japan and Korea (Lee et al.
2002). Alatae are undescribed, and the life cycle is unknown.
Myzus-like aphids in east Asia with long dorsal hairs
like Micromyzodium but having
apterae with secondary rhinaria on ANT III and also sometimes on IV. Alatae
are undescribed. The type species I.
sensoriatus is recorded from both India (A.K. Ghosh et al. 1971f) and China (Su et
al. 2011) but in both cases the host plant was unidentified. Indomyzus glaricae Sathe & Jadhav
NOMEN NUDUM Apterae are dark
brown, shiny; BL c. 1.5 mm. Differences from the type species of the genus include
fewer secondary rhinaria, siphunculi with a much smaller flange, and more
hairs on the cauda. On Gliricidia
maculata in India (Sathe & Jadhav 2008). Other morphs and life cycle
are unknown. Unfortunately the name and description are invalid as no type
specimens were designated.
Two
little-known species described from Quercus,
related to Nipponaphis but without spinal
hairs on abdominal tergite 7, and with paired spinal and marginal tubercles
on prosoma and abdominal plate. Indonipponaphis
fulvicola Sorin
Galls on Distylium racemosum in
Japan are almost globular, up to 27 mm, in cusps of young leaves, and appear
to be very rare (Fukatsu et al.
(2000). Alatae migrate in late October-November to Quercus acuta and Q. gilva.
Apterae on Quercus are brown,
almost circular, dorso-ventrally flattened, strongly sclerotised; BL c. 2.1
mm (Sorin 1979b). Sorin (1997) gave an
account of the life cycle and described all the morphs on Distylium including sexuparae and
sexuales. Fukatsu et al. (2000)
described attacking behaviour by unmodified but long-lived 1st
instars of the gall generation. Only known from Japan; records from India in
Holman (2009) appear to be in error. Indonipponaphis
tuberculata Ghosh & Raychaudhuri Apterae are brownish, oval, strongly
sclerotised; BL 1.9-2.1 mm. On stem of
Quercus dealbata (= Q. floribunda?) in Meghalaya, India
(A.K. Ghosh & Raychaudhuri 1973).
Biology and other morphs are unknown.
One nearctic species with very
short and sparse hairs on body and appendages, a short rounded cauda, short
flangeless siphunculi, well-developed marginal tubercles and a
little-differentiated anal plate. Until recently it was known from only two
apterae collected in Iowa, USA in 1925, but Favret et al. (2004) rediscovered it in Illinois and described all morphs.
A second species described in this genus from China is also included here,
although more likely to belong to another genus, possibly Aphis (Protaphis). Iowana
frisoni Hottes Apterae are pale
green with light reddish brown head
and prothorax and brown siphunculi; BL 1.8-3.1 mm. Alatae have the media
usually once-branched and 12-22 secondary rhinaria on ANT III. In ant
shelters at bases of flowering stems and in leaf axils of Silphium laciniatum and S. terebinthinaceum (no
colonies were found on two other Silphium
spp.). The populations studied in Illinois by Favret et al. (2004) were most commonly attended by Lasius neoniger.
Monoecious holocyclic, with oviparae and apterous males produced in October.
Iowana
zhangi L. Zhang Colour of apterae in life is unrecorded;
BL c.1.6 mm. On Ophiuros exaltatus
in Heilongjiang province, China (L. Zhang 2000). Other morphs and life cycle
are unknown.
Two species on
Compositae/Asteraceae in Africa and east Asia , possibly most closely related
to Aulacorthum, but antennae of apterae as well as alatae bear numerous
secondary rhinaria on III-V, and immatures have spinulose hind tibiae (van
Harten & Ilharco 1976). The host of a third, African, species is unknown. Ipuka
dispersa (van der Goot) Plate 22e Apterae are brownish grey with ill-defined
dark transverse bars on thorax and abdomen, darker brown patches around bases
of siphunculi, dark red eyes, antennae dark except at base of III, legs
yellowish except for black knees and tarsi, siphunculi black and cauda pale
yellowish (see aphids of
Karnataka website); BL 1.6-2.2 mm. On Emilia
spp., often living concealed in leaf sheaths. In Africa, Sri Lanka, China
(Su et al. 2011), south-east Asia
and Australia. Apparently anholocyclic. Ipuka
melantherae van Harten
& Ilharco Apterae are greenish
yellow to yellowish orange, with an orange spot around siphunculi, which are
blackish, and a pale yellowish cauda; BL 1.8-2.2 mm. On undersides of leaves
of Melanthera sp. in Angola.
One species only known from Iran, apparently related to Hydaphias but with first tarsal chaetotaxy 3,3,3, much shorter siphunculi and cauda, and apterae with numerous, long pointed dorsal hairs (Remaudière & Davatchi 1959). Iranaphias dehbani (Remaudière & Davatchi) Apterae are very broadly oval, dark-coloured with slight pruinosity; BL 1.0-1.4 mm. Alatae have secondary rhinaria distributed III 16-24, IV 0-3. On Galium sp. in Iran (Izadbar), at 2,600 m altitude. Life cycle is unknown.
A small, distinctive and probably rather ancient genus of grass-feeding aphids found only in the Mediterranean region and western Iberia. The life cycle is very unusual, and related to the Mediterranean climate. The parthenogenetic phase exploits short-lived annual grasses with a winter growth period following autumnal rains, sexuales being produced and diapausing eggs laid in spring. Alatae are apparently very rare and only known for one species, so dispersal is very limited. Barbagallo & Patti (1999) reviewed the genus and discussed its possible affinities. Nieto Nafría & Mier Durante (1998) reviewed the Iberian species. Israelaphis alistana Mier Durante Apterae are dark green, with rather darker appendages than I. carmini; BL 2.1-2.5 mm. On grasses (Anthoxanthum, Vulpia) in Spain (Zamora). Monoecious holocyclic with oviparae and apterous males in May-June (Mier Durante 1978, as I. tavaresi ssp. alistana). Oviparae are dark green (cf. carmini). Slide-mounted specimens are not reliably separable from I. carmini, and in view of the low dispersal ability of these aphids (no alatae are known), it seems likely that the original subspecies designation of alistana was correct, and that it is a local isolated variant population of carmini. 2n=18. Israelaphis carmini Essig Apterae are greenish yellow, with slight wax covering, and pale appendages; BL 1.9-3.0 mm. On grasses (Avena, Bromus, Hordeum, Koeleria) in hilly regions of Israel, Sicily and Portugal. Monoecious holocyclic, with oviparae and apterous males in March-May (Ilharco 1965, as I. tavaresi; Barbagallo & Patti 1999). Oviparae are brown (cf. alistana). 2n=18. Israelaphis ilharcoi Barbagallo & Patti
Apterae are grass-green or yellowish green, with slight whitish wax
secretion, head brownish, appendages dark-tipped; BL 2.3-3.0 mm. Alatae are unknown. On grasses, especially Bromus subg. Anisantha, in Sicily and Portugal. Monoecious holocyclic, with
eggs hatching in October, apterous viviparae developing during the winter
months and giving rise to oviparae and apterous males in spring (Barbagallo
& Patti 1999). Israelaphis lambersi Ilharco Plate 5a Apterae are greenish yellow to yellow; BL 2.0-2.9 mm. On upper surfaces of leaves of grasses (Avena, Bromus, Phalaris, Phleum) in Portugal and southern Spain. Alatae are very rare and have abnormal wing venation (Ilharco 1966b). Monoecious holocyclic; apterous males are produced in early spring (original description). 2n=16.
About 12 species of squat-bodied, distinctively marked sedge-feeding aphids, leaping from host when disturbed. Alatae have banded wing veins. Six species are nearctic, five palaearctic and one holarctic. The taxonomy of Iziphya is difficult due to seasonal polymorphism and environmental variation in important diagnostic characters (body pigmentation, antennal length, shape of dorsal hairs. Quednau (2010) reviewed the genus, keyed the apterae and alatae of all species and illustrated all available morphs. He sank several names in synonymy and also concluded that some species were less host specific than originally believed. Iziphya albipes Richards Apterae have dorsum mostly black with a pale area on anterior abdomen, a yellowish median area on head, and yellowish tibiae and tarsi; BL c.1.5-1.8 mm. In eastern North America; recorded from Carex pennsylvanica in Minnesota, USA (Quednau 2010), and on Carex sp. growing on rocky hillsides in Quebec, Canada (original description). Iziphya americana (Baker) (Fig.19d) Apterae are yellowish or greyish, usually with a pair of large dark dorsal abdominal patches anterior to siphunculi, not fused medially; BL c.2.1-2.4 mm. On Carex spp. (lepidocarpa, tribuloides) in bogs and wet meadows of northern USA and Canada. Oviparae have been swept from C. tribuloides in October in Oregon (BMNH collection, leg. D. Voegtlin). Iziphya bufo (Walker) Plate 4d Apterae are yellowish or greenish yellow with variable blackish grey markings, consisting usually of a pair of large dark areas on the metathorax and a dark arc joining siphunculi, but sometimes with dorsum almost completely black; BL 1.4-2.1 mm. Alatae have a dark central dorsal abdominal patch and broadly banded wing veins. On various Carex spp., especially C. arenaria, and also on Cyperus rotundus (BMNH collection, leg. R. van den Bosch), with odd records from Juncus (possibly misidentified Juncobia leegei) and grasses such as Cynodon dactylon (as I. maculata); often found in drier situations such as sandhills, drained forest soil and steppe regions, but also sometimes in bogs. Oviparae and apterous males occur in September-October. I. bufo is widespread in Europe, and in south-west and central Asia. This is a very variable species, particularly with regard to the shape of the abdominal and tibial hairs, and its morphological discrimination from I. memorialis is not at all clear. We here follow Quednau (2010) in regarding I. maculata Nevsky, I. austriaca Börner and I. oettingenii Quednau as synonyms of I. bufo. Iziphya eurymelaina Quednau Colour in life not observed, but dorsum with extensive dark pigmentation; BL 1.2-1.7 mm. Vacuum-trapped, possibly from Carex, in Illinois, USA (Quednau 2010). Iziphya flabella (Sanborn) Apterae are yellowish green with variable dark dorsal markings usually consisting of paired dark areas on head, metathorax and abdomen, leaving a clear spinal stripe, but early and late season individuals may be much more extensively pigmented (see Quednau 2010); BL 1.5-2.0 mm. On leaves of various Carex spp., especially on rocky hillsides or in dry forest soil. Widely distributed in North America. Oviparae and apterous males occur in September to early October. Quednau (2010) reported one incidental record from Germany, on C. pilulifera. Iziphya ingegardae Hille Ris Lambers (Fig.19c) Apterae have the dorsum strongly arched and are yellowish with dark dorsal markings, including an irregular dark band forming an arc between the siphunculi, fused at least partially in midline; BL 1.8-2.2 mm. On Carex canescens and C. leporina in bogs in northern Europe (Sweden, Germany, Poland). Oviparae and apterous males occur in Sweden in early September (Ossiannilsson 1959). Iziphya mackaueri Quednau Apterae pale ochreous dorsally except for intersegmental muscle plates and dark spots at hair bases (Quednau 2010); BL c.2.3 mm. On Carex sp. in a wet bog in Ontario, Canada. Iziphya memorialis Börner Apterae are probably yellowish with dark dorsal markings; BL 1.6-1.9 mm. On Carex spp. (praecox, stellulata, possibly montana) in both wet and dry habitats in Europe (Czech Republic, Austria, Germany, Poland and Sweden). Sexuales appeared in Sweden in early September (Ossiannilsson 1959). Iziphya mordvilkoi Quednau & Shaposhnikov Apterae are yellowish green with dark dorsal markings; BL c. 2.1 mm. On Carex sp. in east Siberia. Iziphya spenceri Richards Colour of aptera in life was not observed, probably yellowish with dark dorsal markings; BL c. 1.7-1.8 mm. Described from an unknown host in British Columbia, Canada, and also recorded from Carex sp(p). in western USA from Washington to New Mexico (aphidtrek.org), but it seems to have a holarctic distribution, occurring on Carex sp. in dry steppe country in Mongolia (Quednau 1969; as mongolica), in Kazakhstan (Kadyrbekov 2011b), and also found on C. liparicarpos in Hungary (Szelegiewicz 1981a). Quednau (2010) illustrated an ovipara collected in Oregon in late September. Iziphya variabilis Quednau Colour of apterae in life was not observed, probably yellowish, with variable dark markings; BL c.1.7-1.8 mm. On Carex sp. in wet sites in mountainous steppe country in Mongolia and east Siberia (east Baikal region). Oviparae occur in October (Quednau 2010). Iziphya vittata Richards Apterae are rather elongate-bodied, yellowish or greyish with variable black dorsal markings, often consisting mainly of spots at hair bases, with antennae black except for basal part of segment III; BL 1.8-2.3 mm. On leaves of Carex spp. in wet places, open soil near lake shores, sometimes on road sides, widely distributed in North America. Oviparae occur in late July-August in Quebec (Hille Ris Lambers 1960, as I. punctata; Quednau 2010). |
