SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order) p
About
13 species mostly host-alternating between Populus and roots of Picea. The fundatrices lack wax glands and the
alate spring migrants have forewings with once-branched media. Accounts are available from Heie (1980;
Fennoscandia and Denmark), Danielsson (1976, gall generations in Sweden; and
1990a, root-feeding generations), Danielsson & Carter (1991; root-feeding
generations in UK), Blackman et al. (2019b;
the UK), G.
Zhang & Qiao (1997c; China), G.Zhang et al. (1999c, China) and Smith
(1974a; North America). Pachypappa
aigeiros Zhang Known
from only one alata and two immatures collected on Populus cathayana in July in Shanxi, China (G. Zhang & Qiao
1997c). The gall is not described. Pachypappa
marsupialis Koch
Forming large bag- or blister-like galls (fig. 131E),
open to the underside, on leaves of Populus
nigra in Europe (Austria, Estonia, Germany, Poland, Romania). The fundatrix is brownish green with white
wax. Alatae leave the gall in June
(Börner & Heinze 1957); they have a pinkish yellow abdomen and BL 3.1-3.5
mm (Szelegiewicz 1982). The secondary
host is unknown. A subspecies, P.
marsupialis lambersi, produces yellow blister-like galls on Populus spp. (laurifolia, suaveolens)
in Japan and China (Aoki 1976, G. Zhang et
al. 1985). It has dimorphic first
instar larvae, one morph being migratory and leaving the gall to invade
others (Aoki 1979). Alatae of ssp. lambersi migrate to an unknown
secondary host in July. 2n=10 (for
ssp. lambersi). Pachypappa myrtilli Börner
Apterae secreting wax; BL 1.2-1.4 mm. Described from roots of Vaccinium myrtillus in Germany. Pachpappa
pilosa (Zhang) Appearance in life is unknown, BL of
emigrant alata c. 3.2 mm. On Populus
sp. in Tibet (Zhang & Zhong 1981b; as Sigmacallis
pilosa). Pachypappa
populi (Linnaeus) Forming large pale green, yellowish to
slightly reddish blister-like galls (fig. 131F),
open to the underside, on leaves of Populus
tremula in Europe (Fennoscandia, Latvia, Poland, the Alps), Central Asia
(Kazakhstan; Kadyrbekov 2017a), and also reported (on the closely related P. davidiana) from China (Zhang &
Zhong 1985). The fundatrix is bluish
grey-black, with sparse or no wax; BL 5.1-6.3 mm. Alatae, with olive green abdomen and BL c.
3.5 mm, migrate in early July in Europe, to found large colonies in
flocculent wax on roots of Picea abies. Sexuparae appear in September, but
anholocycly on Picea roots also
occurs in southern Sweden (Danielsson 1990). Pachypappa
pseudobyrsa (Walsh) The fundatrix is wax-dusted, yellowish olive
green with a blackish head; BL 4.0-4.7 mm.
Its feeding causes a small blister-like swelling in the centre of the
underside of the leaf near the mid-rib (see influentialpoints.com/Gallery). Its numerous progeny spread out in May-July
along the main veins, developing into dusky-dark alatae, BL c. 3.5 mm. On Populus
spp. (deltoides, fremontii, tremuloides) in eastern North America, where it is widely
distributed, and also in Colorado (Palmer 1952, as Asiphum). The life cycle
is unknown; the alatae presumably migrate to an unknown secondary host,
although Smith (1974a) thought that it might be monoecious. Pachypappa
rosettei (Maxson)
Feeding by the fundatrix (BL 2.5-2.8 mm, appearance in life unknown)
causes bending of leaves and stunting of stems to form a rosette-like nest,
as in the closely-related European species P. tremulae. On Populus tremuloides in Colorado and
Utah, USA, and secondary host populations are now recorded from Washington
and Alaska (Pike et al. 2012).
Alatae (BL 2.2-2.5 mm) are produced in July.
Host alternation to roots of Picea
spp. has now been confirmed by DNA bar-coding, and the apterous exule (BL
0.8-1.4 mm) redescribed (Pike et al.
2012). Aphids on roots of Pseudotsuga
menziesii have also been tentatively assigned to P. rosettei (Danielsson 1990a). Anholocyclic overwintering may
occur on spruce roots (Pike et al.
2012). Aphids infesting the roots of potted conifers in Canada and identified
as P. tremulae may also be this
species. Pachypappa
sacculi (Gillette)
The fundatrix is wax-dusted, slate grey with brown head; BL 4.5-5.0
mm. It forms a large bag-like gall on P. tremuloides, with edges and tip of
leaf curled downward (Harper 1966; as Asiphum). In western USA (Arizona, Colorado, Utah,
Washingtom, Alaska) and across Canada (British Columbia, Alberta, Quebec,
Ontario and Nova Scotia). Progeny of
the fundatrix are all alate, with dark olive-green abdomen, BL 3-4 mm. Host alternation to Picea spp. has now been confirmed by DNA bar-coding, and a
redescription provided of the apterous exule (BL 0.9-1.8 mm) (Pike et al. 2012). Anholocyclic
overwintering may occur on spruce roots (Pike et al. 2012). 2n=10. Pachypappa
shaposhnikovi Gabrid Forming large bag-like galls from leaves
of Populus spp. (talassica, laurifolia) in Kyrgyzstan
(Gabrid 1989). BL of fundatrix c.4.8
mm, of spring migrant alata c.5 mm).
The secondary host is unknown. Pachpappa
tortuosae Zhang Only
known from a presumably vagrant sexupara collected on Salix matsudana var. tortuosa
in China (G. Zhang & Qiao 1997c), and perhaps belonging to a species
already described from Populus. Pachypappa
tremulae (Linnaeus)
The fundatrix on twigs of Populus
tremula is very large, almost globular, dirty reddish or yellowish brown,
without wax, but appearing silvery due to a covering of fine hairs; BL
5.0-6.6 mm. Its progeny move onto new
shoots and cause bending of leaves and petioles and stunting of growth of
shoot, forming a rosette-like leaf-nest (see influentialpoints.com/Gallery). Second generation are all alate,
orange-yellow to pinkish brown, wax-covered, with black head and pterothorax;
BL 3.2-4.4 mm. They migrate in June to
found colonies on roots of Picea abies
(Stroyan 1975). Apterous exules are
pale yellowish white with tufts of wax posteriorly; BL 1.3-2.1 mm. Sexuparae return to aspen in
September-October, but populations also overwinter anholocyclically on roots
of Picea. Widely distributed in the northern
palaearctic, east to China (Zhang & Zhong 1985b) and Japan (Aoki 1976; as
P. shirobamba). The numerous
records from North America on roots of Picea
spp. and Pseudotsuga menziesii
(also as Asiphum tremulae and Rhizomaria piceae; Lavallee 1987) may
all be due to confusion with P.
rosettei (Footit et al.
2006). The male genitalia were described and illustrated by Wieczorek et al. (2012). 2n=10. Pachypappa
vesicalis Koch
Forming irregular green to yellowish bag-like or blister-like galls
(see Carter & Danielsson 1993 for photograph) on leaves of Populus spp. (alba, canescens). The fundatrix is reddish-yellow to orange-red,
wax-dusted; BL 4.2-4.9 mm. All second
generation are alate, with reddish-brown abdomen, BL c. 3.3 mm, migrating in
June to found colonies on roots of Picea
spp. (abies, glauca, sitchensis),
with a record also from Pseudotsuga
menziesii in Czech Republic (Hochmut 1968). Apterous exules on spruce roots are pale
yellowish white secreting white filamentous wax. Sexuparae return to Populus in September-October; mating and oviposition on poplar
bark were described by Carter & Danielsson (1991). In northern and north-west Europe. Pachypappa
warshavensis (Nasonov) (= varsoviensis Mordvilko)
Forming loose leaf-nest galls on Populus
spp. (alba, canescens, euphratica,
pruinosa; see Carter &
Danielsson 1993 or Casiraghi et al.
2019 for photographs). Fundatrices
(feeding on twigs) are red-brown; BL 3.5-4.4 mm. Alatae from galls are yellowish orange, BL
3.0-3.8 mm, migrating in June, and apparently found colonies on roots of Populus and Salix spp., although this still needs to be confirmed by
experimental transfers. Casiraghi et
al. (2019) provide a redescription and illustrations of fundatrices and
alate fundatrigeniae. Apterae tentatively identified as this species have
been found on the roots of Salix caprea
(Szelegiewicz 1982) and S. silesiaca (BMNH
collection) in Poland, and on Populus
alba roots in Sweden (BMNH collection). Carter & Danielsson (1993)
found many adult apterae and dark yellowish alatoid nymphs associated with
wax particles in soil around roots of P.
canescens at a location in Wales in October, and collected adult
sexuparae from the rough bark of the same group of trees. They also reported
finding apterae in June on fine roots of P.
canescens in England (Lancashire). In Europe and Central Asia
(Kazakhstan: Kadyrbekov 2011a).
2n=10*.
One
or two species closely related to Pachypappa,
but the fundatrix has wax glands on the thorax and abdomen. Pachypappella
aliquiphila Zhang
Known only from the fundatrix, BL 2.2 mm, collected from an
unidentified Populus sp. in Shanxi
Province, China (G. Zhang & Qiao 1997c). The gall is not described.
Possibly this is a small specimen of P.
lactea. Pachypappella
lactea (Tullgren)
The gall is formed on Populus
tremula by folding down of the leaf lamina to form a cone, which becomes
bright orange. Fundatrices are greyish
brown with white wax; BL 4.6-5.2 mm.
All second generation are alate, with dark greyish brown abdomen, BL
2.5-3.5 mm, migrating in June-July to colonise roots of Picea abies (Danielsson 1990).
The root-feeding apterae live in nests of dense wax wool, 1.5-2.0 mm
across, usually one aphid in each (Albrecht 2015). Anholocyclic overwintering
on Picea roots, including P. sitchensis (Danielsson & Carter
1991), may be common. Northern
palaearctic and boreo-alpine in distribution, from Britain to east Siberia
(Pashchenko 1988a).
About 10 rather large, shiny brown species and subspecies
mostly in eastern Europe and western Asia, related to Macrosiphoniella, with both pre- and postsiphuncular sclerites
well-developed, and mostly associated with Echinops. The Echinops feeders
have a long wedge-shaped R IV+V. Holman (1981) provided a key and detailed
morphological comparisons, also noting the existence of a single aptera of an
undescribed species from an unknown host in Ethiopia (BMNH collection, leg.
H. Scott). Paczoskia brevipilosa Tashev Apterae
are shining dark brown; BL 1.9-2.8 mm. On leaves of Echinops
spp. in Bulgaria and Rumania (Holman 1981). Biology has not been studied; Holman
suggested that its morphology might indicate a close association with ants
and subterranean life at some stage in its life cycle. Paczoskia colchica Holman
Apterae are shiny brown, BL
2.6-3.8 mm. On undersides of leaves of
Echinops galaticus (= ossicus) in southern Russia (Krasnodar). Monoecious holocyclic, with
oviparae and apterous males in late October (Holman 1981). Paczoskia jurineicola (Bozhko)
Apterae are shining dark brown with black appendages; BL c.2.2 mm. On Jurinea stoechadifolia in Ukraine,
forming dispersed colonies on undersides of leaves (Bozhko 1976b). Paczoskia longipes (Tashev)
Apterae are shining dark brown, BL 4.3-4.7 mm. On Echinops spp. in
eastern Europe (Bulgaria, Poland, Rumania). Paczoskia major Börner Apterae are shining dark brown, BL
2.4-4.3 mm. Described from Echinops
sphaerocephalus in Germany, and since found in various east European
countries, and in Turkey and Iran (Goodarzifar et al. 2010). In Turkey it
was common on stems of E. ritro
(Remaudière & Tuatay 1963). A single aptera (without host data)
ascribable to this species was collected in 1926 at 3,000 m in Ethiopia (BMNH
collection, leg. H. Scott). Sexuales occur in Europe in September-October,
with both apterous and alate males (original description). A form on E. microcephalus and Echinops sp. in Bulgaria and Turkey is
regarded as a subspecies, P. major ssp. bulgarica Holman. Paczoskia meridionalis Holman
Apterae are shining dark brown, BL 3.1-4.2 mm. On Echinops spp. in Iran, Israel, Lebanon and former Yugoslavia.
Holman (1981) compared the morphology of different populations. Paczoskia obtecta Börner Plate 27e Apterae are shiny bronze brown, with bright
red eyes, black siphunculi and cauda, antennae black except at base of III, and
legs yellowish with dark segmental apices; BL 3.1-4.3 mm. On undersides of
lower leaves, and later on upper parts of stems, of Centaurea
spp. In Europe (France, Sweden,
Denmark, Germany, Poland, Austria, former Yugoslavia, Hungary, Czech
Republic), and in Israel. Sexual morphs are apparently unknown. 2n=12. Paczoskia paczoskii Mordvilko
Apterae are shining dark brown, BL 2.3-4.2 mm. On stems and undersides
of leaves of Echinops spp. in
Europe (France, Rumania, Ukraine).
Monoecious holocyclic; oviparae were recorded in France in
October. Nevsky (1929a) described a
subspecies from E. karatavicus in
Uzbekistan, P. paczoskii ssp. turanica,
and this subspecies is also recorded from Kazakhstan (Kadyrbekov
& Aoitzhanova 2005, Kadyrbekov 2014e) and Greece (Tsitsipis et al.
2007). Holman (1981) compared various populations and described P.
paczoskii ssp. ruthenica from
E. ritro ssp. ruthenicus in Hungary and Crimea. Kadyrbekov (2010, 2014g)
identified aphids on E. albicaulis
in Kazakhstan and on E. ritro in
Russia as ssp. ruthenica. Paczoskia wagneri (Remaudière
& Tuatay) Apterae are shining
dark brown to almost black in life; BL 2.7-3.4 mm. Described from an
unidentified Echinops sp. in Turkey
(Remaudière & Tuatay 1963, as Macrosiphoniella
subgenus Paczoskia). No other
morphs are recorded.
A
genus of one or two little-known species.
Its taxonomic position within Aphidinae is uncertain. Paducia
antennata (Patch) (fig. 106A) Only oviparae (collected October-November)
and a single alate vivipara have been described (Patch 1913, Hottes &
Frison 1931). Oviparae are dark
greenish or chocolate-brown to black dorsally, greenish yellow ventrally; BL
c. 2 mm. Alata are mainly brown, BL c.
1.4 mm. Presumably it lives without
host alternation on the roots of Salix
sp., although the evidence for this host association is entirely
circumstantial. Recorded from
Illinois, New York and Maine, USA. Paducia
aterrima Hille Ris Lambers Apterae are "probably shiny
black"; BL 2.0-2.2 mm. Other
morphs are unknown. On roots of Salix sp. in Greenland (Hille Ris
Lambers 1952). Some differences from P. antennata were given by Hille Ris
Lambers, but as different morphs were being compared the distinctness of the
two species requires verification.
Large
distinctive aphids with short antennae living on Juglans. The three described taxa are closely related and could
perhaps be geographic variants of a single species. In the past the name Callaphis was widely used for this
genus, but was suppressed following a plea by Quednau (1983), on the grounds
of potential confusion with Calaphis
Walsh. Quednau (2003) revised the genus. Panaphis
juglandis (Goetze)
All viviparae are alate, large and stout-bodied, with dark head and
thorax, yellow abdomen with dark brown transverse bands, and forewing veins
fuscous-bordered (see
influentialpoints.com/Gallery); BL 3.5-4.3 mm. On upper sides
of leaves of Juglans regia, in rows
along the veins, often ant-attended. In Europe, central Asia, Pakistan and
north-west India, and introduced (in 1928) to western USA, where populations
seemed to increase after the successful control of Chromaphis juglandicola . It appears that these two species
cannot readily coexist, because colonies of P. juglandis on upper surfaces of walnut leaves are adversely
affected by a rain of honeydew from colonies of Chromaphis on the undersides of the leaves above (Olson, 1974).
Oviparae and alate males occur in September-October. The male genitalia were described and illustrated by Wieczorek et al. (2011). Masoodi
et al. (1987) studied its
population dynamics in Kashmir. Biological control has been attempted in
California with a Praon species
from Afghanistan (Mackauer & Sullivan 1982). 2n=22. Panaphis
nepalensis (Quednau) All viviparae are alate, colour in life
unrecorded but probably like P.
juglandis (see Quednau 1973). On Juglans
regia in Nepal, Tibet and China (Yunnan; Qiao et al. 2005b). Sexuales are unrecorded. Zhang in Zhang &
Zhong (1985c) described a subspecies, P. nepalensis yunlongensis, from Juglans mandshurica in Yunnan, China.
About
22 species on Burseraceae and Combretaceae in Africa (plus one in India),
closely related to the South American genus Lizerius. Both apterous
and alate viviparae are known for most species, but there are a few species
which do not seem to produce apterae. Apterae and often alatae have
conspicuous long dorsal processes (fig. 40). Quednau
(2010) reviewed the world fauna and illustrated all available morphs. Paoliella
ayari (Eastop)
Apterae are yellow, BL c. 0.9 mm, on undersides of leaves and young
shoots of Commiphora spp. (pedunculata, angolensis). Recorded from
Nigeria, Tanzania, South Africa and Zimbabwe.
Life cycle unknown. Paoliella
browni Quednau
The single known aptera, possibly a fundatrix, was dark brownish
velvety green with white dorsal processes; BL c. 1.1 mm. Quednau (2010) has illustrated the alata
and alatoid nymph of this species (the two alatae described in Quednau (1962)
are P. echinata). On Combretum gueinzii (= C. molle) in South Africa. Van Harten (1979) described an aphid with
greenish yellow alatoid nymphs living solitarily on undersides of very old
leaves of Combretum sp. in Angola
as a subspecies, P. browni capinganae.
Alate males and apterous oviparae of this form (the latter dark
velvety green, BL 1.0-1.1 mm) were collected in early June. An alata trapped in Burundi (Remaudière
& Autrique 1985) may be this subspecies. Paoliella
capeneri Quednau
Apterae are unknown and colour of alatae in life unobserved; BL of
alata 1.0-1.4 mm. On Commiphora
angolensis in South Africa (Quednau 2010). Paoliella
chiangae van Harten & Ilharco Only alatae are described; they have head,
thorax and legs brown-black, abdomen greyish brown with dark tubercles, BL
1.2-1.6 mm (van Harten & Ilharco 1972).
Living singly or in small colonies on undersides of young leaves of Combretum sp. (?psidioides) in Angola. Paoliella
commiphorae (Doncaster) All viviparae are alate, yellowish, thickly
dusted with grey wax; BL c. 1.5 mm. On
Commiphora eminii ssp. zimmermannii in Kenya and Malawi. Similar aphids collected on C. fischeri (= mossambicensis) in
Tanzania were described as a subspecies, C. commiphorae persimilis (Eastop
1955). Alate males of this latter form
have been trapped in January-March (BMNH collection). Paoliella
commiphoroides Quednau Apterae are unknown, colour of alata in
life was not observed; BL of alata 1.2-1.5 mm. On Commiphora mildbraedii in Kenya (Quednau 2010). Oviparae and
alate males were collected in May along with alate vivparae and alatoid
nymphs. Paoliella
delottoi (Hille Ris Lambers) Apterae are brownish, BL 0.9-1.2 mm,
feeding on undersides of shoot-tips of Commiphora
africana in Eritrea and Nigeria.
The life cycle is unknown.
Aphids collected on C. dulcis
in Namibia (BMNH collection, leg. H.D. Brown) are also regarded as this
species. Paoliella
eastopi Hille Ris Lambers
Colour in life is unknown; BL of aptera 0.9-1.4 mm. Found in large
numbers on Commiphora eminii ssp. zimmermannii in Kenya (Hille Ris
Lambers 1973d). Alatae were present in January, March and May. Paoliella
echinata Eastop
Apterae are black with white dorsal processes; BL c. 0.8 mm. Immatures are yellow. On Commiphora
pilosa in Tanzania and South Africa.
The biology is unknown. Paoliella
(Lizerocallis) flava (Sousa-Silva & Ilharco) Apterae are yellowish, with dark apices to
antennae and rostrum; BL 0.6-0.8 mm. Living dispersed on undersides of leaves
of Tibouchina granulosa in Brazil
(Sousa-Silva & Ilharco 2003). Quednau (2010) illustrated apterous and
alate morphs. Paoliella
harteni Ilharco
Only alatae are known, orange-reddish appearing rose-coloured due to
fine pulverulence; BL 1.5-1.9 mm. On
both sides of leaves of Terminalia
spp. (brachystemma, sericea) in Angola (Ilharco
1971). Van Harten & Ilharco(1972)
described the alate male and apterous ovipara, collected in July. Viggiani (1987) described a new aphelinid
parasitoid from this aphid. Paoliella
hystrix Theobald Plate 7e The colour of the aptera in life is
not recorded; BL 0.5-0.8 mm. Alatae
and life cycle are unknown; only known from original collection on Commiphora savoiae (= edulis ssp. boiviniana?) in Somalia (Theobald 1928). Paoliella
(Lizerocallis) laxutuberculata Quednau Apterae are yellowish; BL c. 0.9-1.0 mm.
On Combretum laxum in Belize, and
also found on an unidentified host in Mexico. Apterae and alatae were
described an illustrated by Quednau (2010). [This aphid was wrongly
identified as Lizerius cermelii in
Blackman & Eastop (2006, p. 1189).] Paoliella
(Kenyaphis) lepidosoma Quednau Apterae are orange-coloured in life: BL
1.0-1.2 mm. On ?Commiphora sp. in
Kenya (Quednau 2010). An immature ovipara was collected in May. Paoliella
longirostris Quednau Only alatae are known, and their colour in
life is unrecorded, BL c. 1.5 mm.
Described from an alata trapped in Zimbabwe (Quednau 1974), and
subsequently collected in Burundi on Combretum
sp. (Remaudière & Autrique 1985). P.
monotuberculata van Harten & Ilharco Dense colonies of reddish-brown alatoid
nymphs together with a small number of adult alatae (BL 1.2-1.5 mm) were
found on the undersides of young leaves of Combretum sp. (?psidoides)
in Angola (van Harten & Ilharco 1972). The alata was illustrated by
Quednau (2010), and data are tabulated for comparison with P. browni capinganae by van Harten
(1979). Paoliella
nachensis (Eastop)
All viviparae are alate, with head and thorax dark, abdomen yellow
with dark dorsal processes; BL 1.1-1.4 mm.
Living singly on undersides of leaves Combretum spp. (incl. apiculatum,
?psidioides). Recorded from Angola, Burundi and
Tanzania. The life cycle is unknown. Paoliella
namaachae Ilharco
Only alatae are known, with pale yellow-orange abdomen and slightly
darker head and thorax; BL 0.9-1.2 mm.
Living solitarily on undersides of leaves of Combretum molle in Mozambique.
Ilharco (1976) discusses differences from the closely-related P. nachensis. The life cycle is unknown. Paoliella
nirmalae (David)
Apterae are bright brick red with numerous dull white dorsal processes
(see aphids
of Karnataka website); BL of aptera 1.1-1.4 mm. On Terminalia
arjuna in Mysore, India (David 1969a).
Ghosh & Quednau (1990) provided a redescription of aptera and alata,
and discussed the differences setting this Indian species apart from all the
African ones. The life cycle is unknown. Paoliella
papillata (Hall) All
viviparae are alate, very dark brown; BL 1.4-1.5 mm. Immatures are pale reddish brown with white
dorsal processes (Hall 1932). On Combretum spp. (incl. apiculatum, molle), widely distributed in Africa south of the Sahara. Doncaster (1954b) described the alate male
and apterous oviparae from amongst the type material, collected in Zimbabwe
in December. Paoliella
pteleopsidis Quednau Only alatae are described; colour in life
is unknown, BL maximally c. 1.8 mm. On
Pteleopsis anisoptera in Angola
(Quednau 1974). Paoliella
terminaliae (Hall) All viviparae are alate, brick red to very
dark but with tip of abdomen always pale; BL c. 1.5 mm (Hall 1932). On Terminalia
sericea in Nigeria and southern Africa (Angola, Malawi, South Africa,
Zimbabwe). The life cycle is
unknown. Paoliella
ufuasi (Eastop)
Only alatae are known, appearance in life unknown; BL 1.2-1.4 mm. Described from trapped alatae in Tanzania
(Eastop 1955), but apterae collected on Commiphora
pilosa in Kenya are also identified with this species (Quednau
2010). The life cycle is unknown, but
an aptera illustrated by Quednau and collected in September in Kenya might be
a fundatrix. Paoliella
wettsteini Quednau
Apterae are pale yellow with white dorsal processes; BL 1.0-1.2
mm. Alatae are very pale with head and
thorax only slightly dusky. On leaves
of Combretum gueinzii (= C. molle) in South Africa (Quednau
1964). The life cycle is unknown.
One
fern-feeding North American species, related to Macrosiphum but with unusual antennal sensoriation. Alatae often have a once-branched media. Papulaphis sleesmani (Pepper) Plate 26c (Fig.43b) Apterae are lemon yellow to lemon green; BL
1.6-2.1 mm. On Adiantum pedatum and
Cystopteris bulbifera in eastern
USA.
A
genus for one oriental species on Spiraea
with apterae having reduced eyes (triommatidia in fundatrices and
oviparae), a much shorter antennal PT than in alatae, and very long dorsal
hairs arising from dark tuberculate bases. The life cycle is unknown, but it
is believed to migrate to an unknown secondary host (see below). Formerly given its own subfamily
(Parachaitophorinae), Sugimoto (2011c) placed this genus in Drepanosiphinae
because of characters of the oviparous morph, although heteroecy is hitherto
unknown in that subfamily. Parachaitophorus spiraeae (Takahashi) (= P.
sikhotealinicus Pashchenko, = P. yamashitai Sorin; synonymies by
Sugimoto 2011c) Apterae in life are
brown with dark brown head, antennae, legs, siphunculi, cauda and dark brown
spots at bases of dorsal hairs; BL 1.2-1.7 mm. On Spiraea spp. in
Japan, Korea and east Siberia, forming compact ant-attended colonies on stems and at bases of young
offshoots, but Japanese populations disappeared in late May perhaps
indicating migration to an unknown secondary host. Alatae
(migrants?) produced in spring and autumn were compared by Sugimoto (2011c),
who also described oviparae and alate males collected on S. cantoniensis
in October-November. The
male genitalia were described and illustrated by Wieczorek et al. (2011).
About
four species resembling Forda except
for the longer outer margin of the hind coxae. Only P.
cimiciformis is known to complete the holocycle with galls on Pistacia; sexuparae of P. donisthorpei have been recorded
from P. terebinthus (Roberti 1939,
Nieto Nafría et al. 2002a), but the
gall-living generations of that species are either unknown or confused with
those of P. cimiciformis. Molecular
studies indicate a close relationship between Paracletus and Forda compared
with other genera of Fordini (Ortiz-Rivas et
al. 2009). Hille
Ris Lambers (1954b) discriminated the species on the material then available.
Paracletus bykovi (Mordvilko) Apterae are yellowish white; BL 2.4-3.5
mm. On roots of many species of Poaceae (Calamagrostis,
Festuca, Hordeum, Setaria, Poa); also Luzula (Juncaceae). In eastern Europe (Poland, Ukraine), Turkey,
Transcaucasia, Kazakhstan, and Uzbekistan, where a population was described
as P.
bykovi ssp. uzbekistanicus
Kan. Apparently mainly or entirely anholocyclic, although Mordvilko (1935) described
a sexupara collected on trunk of Pistacia
mutica in Georgia. Different from other Paracletus in several respects, and possibly deserving of
separate generic or subgeneric status (as Hemitrama).
Paracletus
cimiciformis von Heyden The galls on Pistacia (khinjuk, palaestina, terebinthus, vera) are
flat folds of the edges of the leaves, retaining the original leaf
colour. The fundatrix – globose,
yellow, with 6-segmented antennae, BL c. 1.5 mm - has never been properly
described. Second generation are all
alate (BL 1.7-2.4 mm, fig. 81N) emerging
from the gall in September-October to found colonies on roots of many species
Poaceae (e.g.
Agrostis, Brachypodium, Dactylis,
Festuca, Hordeum, Oryza, Poa, Polypogon, Triticum).
Apterae of the typical form on grass roots are shining waxy yellowish white
to pale brown, with body dorso-ventrally flattened, especially at lateral
margins, and 6-segmented antennae; BL 2.4-3.5 mm (Plate 2d).
They are always attended by ants, especially Tetramorium spp. However transfer experiments indicated the existence of a
second distinct grass root-feeding apterous morph of this species (Ortiz-Rivas
et al. 2009), and this has been confirmed by comparison of DNA sequences (Mróz &
Depa 2012). This morph has a dusky
olive- to apple-green, highly domed body, BL 1.8-3.3 mm, and 5-segmented
antennae, and was previously identified as Forda rotunda Theobald. It has been recorded from many genera of
Poaceae (Brachypodium, Dactylis, Festuca, Poa, Polypogon), and is also ant-attended
(Zwölfer 1958, as Forda skorkini). The factors influencing the
production of these two morphs and their inter-relationships have not yet
been elucidated (see Salazar et al.
2105, supporting information). However, work in Spain has now established
that the two morphs have different relationships with the ant Tetramorium semilaeve; the round morph
has a normal mutualistic relationship, whereas the flattened morph is an
aggressive mimic with cuticular hydrocarbons that resemble those of the ant
larvae, so that it is transported into ants’ nests where it sucks on ant
larval haemolymph (Salazar et al.
2015). Sexuparae return
to Pistacia in April-June; these
and the sexual morphs were fully described by Roberti (1939). P.
cimiciformis is recorded from Pistacia
in southern France, Italy, Iran, Israel, Malta (Mifsud et al. 2009) and Turkey.
Anholocyclic populations on roots of grasses, recorded as either
typical P. cimiciformis or as the morph previously identified as Forda rotunda, occur outside the range
of the primary hosts throughout Europe, in north Africa and across Asia to
Japan, Korea and China. Some regionally variant populations have been described as subspecies
(panicumi Bozhko, on Panicum miliaceum in Ukraine, and zhanhuanus Zhang &
Qiao (Zhang & Qiao, 1998d) on Triticum
aestivum in China). For fuller
information see Zwölfer (1958) and references therein. T. Zhang et al. (1987) studied
the bionomics of this aphid in China.
Álvarez et al. (2009)
studied the histology of the gall. 2n=16. Paracletus donisthorpei
Theobald Apterae are pale; BL c.2.9-3.0 mm.
Described from nests of the ant Tapinoma
nigerrimum, and since found on roots of Triticum sp. and unidentified woodland grasses (Nieto Nafría et al. 2002a). In Italy, Spain, Greece
and Algeria. The life cycle is unknown; sexuparae have been collected on Pistacia terebinthus (Roberti 1939),
but the galls have not been identified and are possibly identical to those of
P. cimiciformis (Ortiz-Rivas
et al. 2009). Paracletus subnudus Hille Ris Lambers Colour of apterae in life is unknown; BL
of last instar imm. 2.6-3.0 mm. On Hordeum sativum (= vulgare) in Israel. Recorded also now
from Turkey (Şenol
et al. 2017), but this requires
additional confirmation.
Two
species in east Asia similar to Colopha,
but the media of the forewing of alatae is unbranched as in Tetraneura, and the apterae on the
secondary hosts (bamboos) always have siphunculi, which are displaced towards
the midline. The life cycle of only
one species is known. Akimoto (1985a)
gave a full account. Paracolopha
morrisoni (Baker)
The gall on Zelkova serrata
is a club-shaped outgrowth from the upper surface of the leaf lamina between
veins, with a narrow stem and a swollen, roughened apex (fig. 125c
and http://www.agr.hokudai.ac.jp/systent/akimoto/gall/P.morrisoni.jpg
and influentialpoints.com/Gallery). Alatae (fig. 120B)
emerge in June-July and colonise roots of bamboos (Sasa spp.). Sexuparae
return to Zelkova in
September-October. Akimoto &
Narita (2002) studied the influence of alternating temperatures on the
hatchability of overwintered eggs.
Moritsu's (1958) record from Celtis
sinensis var. japonica (as Tetraneura nishiyae) requires
confirmation. Galls on Zelkova occur in Japan and southern
China (as Tetraneura zelkovisucta;
Zhang 1980b). Apterae on roots of bamboo are globular, pale yellow with
greyish legs and antennae, covered completely with white filamentous wax
(Malumphy 2012); BL 1.5-1.8mm. Presumably anholocyclic populations have been
introduced into USA, where it is recorded from the roots of various bamboos (Phyllostachys, Arundinaria) in Maryland, South Carolina and southern California
(Hille Ris Lambers 1966a). It has been recorded on bamboo roots in Britain
since at least 1998, and may be widely distributed in Europe (Malumphy
2012). This species has a complex
synonymy; commonly used names in the past have been Colopha moriokaensis (Monzen) and Dryopeia nishiyae (Matsumura 1931, nec. 1917). 2n=10*. Paracolopha takahashii Akimoto Appearance of apterae in life is unknown;
BL 1.25-1.5 mm. On basal part of Carex
breviculmis ssp. royleana in Japan. Only known from the
original collection (1956).
Two
species in New Zealand, resembling Aphis
but lacking lateral tubercles on ABD TERG 7, and often also lacking them on
ABD TERG 1. Molecular studies (von Dohlen & Teulon 2003, Foottit et al. 2008) indicate a close
relationship with indigenous New Zealand Aphis
species. Teulon et al. (2013)
summarised collection data and other the available information about both
species. Paradoxaphis
aristoteliae Sunde Apterae are reddish brown with black head
and dusky thorax; BL of aptera 1.5-2.0 mm.
On leaves of Aristotelia serrata
in New Zealand (Sunde 1987). Oviparae
and eggs were obtained in January (Teulon et
al. 2013). Podmore et al.
(2019) reported COI sequence variation between populations. 2n=8*. Paradoxaphis
plagianthi Eastop Apterae are salmon-brown to brown with a
green to dark brown longitudinal stripe; BL 1.3-2.0 mm. On leaves of Plagianthus regius in New Zealand (Eastop 2001). Kean &
Stufkens (2005) reported that Hoheria
angustifolia and the woody vine Parsonsia
heterophylla may act as “spill-over” hosts. It seems to be anholocyclic,
persisting on plump terminal buds for the short time in winter when Plagianthus is completely leafless
(Kean 2002).
Three
palaearctic species, one on Rosaceae and two on Apiaceae, resembling Myzus but with apterae having
secondary rhinaria on ANT III. Shaposhnikov & Stekolshtshikov (1989)
reviewed the genus and provided a key to species. Paramyzus heraclei Börner Plate 18e Apterae are white or yellow, shiny, pale
except for tarsi (see influentialpoints.com/Gallery);
BL 1.3-1.9 mm. On undersides of basal leaves of Heracleum spp., causing many small yellow spots and slight
vaulting of the leaves. In Japan it is recorded from Angelica spp. (Miyazaki, 1971). Monoecious holocyclic on Heracleum in Germany, with alate males;
a full account of the life cycle and sexual morphs was given by Müller
(1977b). Widely distributed in Europe
(not recorded from Scandinavia), Turkey, and in east Siberia and Japan.
Japanese populations are possibly a different race or subspecies, P.
heraclei ssp. similis Takahashi (see also Shaposhnikov
& Stekolshtshikov, 1989). Paramyzus longirostris Miyazaki
Apterae are pale yellow, yellow or green, with antennae black towards
tips; BL 1.1-1.4 mm. Alatae are undescribed. On Potentilla spp. in Japan and east Siberia (Kamchatka), and also
found on Fragaria sp. in Japan (BMNH collection,
leg. R.L.Blackman). 2n=14. Paramyzus sii Shaposhnikov & Stekolshchikov
Apterae are shiny greenish yellow with black eyes and tarsi, greenish
siphunculi and whitish cauda; BL 1.6-2.0 mm. Alatae are undescribed. On Sium suave, at leaf bases and on
inflorescences, in east Siberia (Vladivostok).
One
species on Quercus in Japan, close
to Nipponaphis but without
siphuncular pores. Paranipponaphis
takaoensis Takahashi
Apterae are nearly circular, with side of body perpendicular
("pill box-shaped"), blackish brown, BL 1.5-2.0 mm. On branches of an unidentified Quercus sp. in Japan (Takahashi
1959b). Other morphs and life cycle
are unknown.
One
species in China with Phorodon-like
digitiform process on the antennal tubercles, and dorsal cuticle densely
papillated. Paraphorodon omeishanaensis Tseng & Tao Apterae are
pale green, with dusky antennae; BL c.1.3 mm. Alatae are undescribed. On
undersides of leaves of an unidentified Rubus
sp. in China. This species has been mistakenly synonymised with Phorodon cannabis.
Seven
species in east and south-east Asia differing from Thoracaphis in the complete fusion of prosoma and abdominal
plate, different cuticular ornamentation, spine-like marginal hairs and
absence of siphunculi. Six species are
recorded from Fagaceae, although one of these was originally described from Litsea; the host of the seventh is
only tentatively recorded as Myrica.
Only apterous viviparae are known.
Some of the species may have unrecognised gall-forming generations on Distylium. The genus was reviewed and
keyed by J. Chen et al. (2016a),
and two of the Chinese species were reviewed by Qiao et al. (2018). Parathoracaphis
cheni (Takahashi)
Apterae are black, oval, flattened, strongly sclerotised, without wax:
BL c. 0.9 mm. Sedentary on undersides
of leaves of ?Myrica rubra (host is
queried in original description) in China (Chekiang). The life cycle is unknown. Parathoracaphis
elongata (Takahashi)
Apterae are black, elongate oval, strongly sclerotised, with numerous wax
filaments dorsally: BL c. 0.5 mm. On
undersides of leaves of Quercus
sp., and other unspecified Fagaceae, in Thailand (Takahashi 1941). The life cycle is unknown. Parathoracaphis
gooti (Takahashi)
Apterae are black, broadly oval, flattened, sclerotised, with much
white marginal wax: BL c. 1 mm. On
undersides of leaves of Quercus sp.
in Malaya (Takahashi 1950). The life
cycle is unknown. Parathoracaphis
kayashimai (Takahashi)
Apterae are presumably black, elongate oval, strongly sclerotised,
perhaps without wax (not stated); BL c. 0.5 mm. On undersides of leaves of Quercus sp. in Malaya (Takahashi
1950). The life cycle is unknown. Parathoracaphis
manipurense (Pramanick, Samanta &
Raychaudhuri) Apterae are broadly
oval, aleyrodiform, brown to black; BL
0.75-0.9 mm. On Litsea sebifera (= glutinosa) in Manipur, India (Pramanick et al. 1983), and also recorded from Castanopsis sp. (A.K. Ghosh
1988). Biology is unknown. (An apterous morph collected on Amaranthus in another locality was
described as the paratype fundatrix of this species, but no life cycle
observations or other evidence linking these two morphs was reported, and Amaranthus is a most unlikely primary
host for a hormaphidine.) Parathoracaphis
setigera (Takahashi)
(fig. 91F)
Apterae
are black, elongate oval, strongly sclerotised, without wax; BL 0.5-0.7
mm. In loose clusters on undersides of
leaves of Quercus spp. (glauca, myrsinaefolia) and Lithocarpus
sp. in Japan, China (Sikong) and Taiwan (Tao 1966). There are also records
from Palaquium sp. (Sapotaceae) and
Styrax suberifolia (vagrants?) in
Taiwan (Tao 1999). In Japan apterae
occur on Quercus throughout the
year (Takahashi 1958a). Parathoracaphis
spinapilosa Chen,
Jiang & Qiao Apterae are black,
with a fringe of white curved wax filaments, sparser posteriorly; BL 0.6-0.8
mm (J. Chen et al. 2016a). In large
colonies on the undersides of leaves of an unidentified Quercus sp. (and also on another unidentified species of
Fagaceae). In China (Fujian and Yunnan). The life cycle is unknown.
One
Indian species on Quercus,
apparently close to Metathoracaphis. Only apterous viviparae are known. Parathoracaphisella
indica Pramanick, Samanta & Raychaudhuri Apterae are dark brown, with margin
darker, oval: BL c. 1.1 mm. On Quercus sp. at Manipur, India
(Pramanick et al. 1983). Other
morphs and biology are unknown.
One
North American species on Fagaceae, perhaps closely related to Lachnochaitophorus, but the apterae
have separate head and prothorax. Patchia
virginiana Baker Apterae
are broadly oval, rather uniformly velvety black, secreting wax; BL 1.8-2.3
mm. Alatae are brown with a large
black dorsal abdominal patch and thickly-bordered forewing veins. On bark of Castanea spp. in USA (Florida, North Carolina, Pennsylvania,
Virginia). Apparently monoecious
holocyclic; oviparae were collected in September (BMNH collection, leg. J.O.
Pepper).
One
palaearctic species host-alternating between Tilia and Araceae. Patchiella
reaumuri (Kaltenbach)
(fig.
115B) This species forms large
leaf-nest galls on Tilia spp. in
spring by twisting and stunting the terminal growth and clumping the leaves
(see influentialpoints.com/gallery)
. Fundatrices are plump-bodied,
yellowish brown to greenish; BL c. 3.5 mm.
All second generation are alate, with abdomen green to yellow-green,
BL 2.1-2.7 mm, migrating to found colonies on roots and tubers of Arum spp. (Roberti 1939) and other Araceae (Arisaema, Colocasia). Apterae
on the secondary host are whitish, with a powdery wax coating, and flocculent
wax on the posterior abdomen; BL 1.9-2.5 mm (Stroyan 1979b). It
is recorded from Tilia in England,
Italy, Bulgaria, Hungary, Spain, Greece and Turkey. Anholocyclic populations
occur on roots of Araceae elsewhere (e.g. Hawaii, Solomon Islands), and also
in China, as Gharesia kolokasia Qiao
& Zhang is almost certainly a synonym (Qiao & Zhang 1999b) . Pashchenko (1984a) described a subspecies, P.
reaumuri orientalis, from Tilia
amurensis in east Siberia; this is presumably the insect referred to as orientalis Mordvilko in Shaposhnikov
(1955).
One
species on Madagascar related to Eonaphis
and Schoutedenia, but with two
pairs of dorsal processes on abdominal tergite 7, and a different host
association. Remaudière (1988) gave a
full account. Paulianaphis
madagascariensis Essig
Apterae are broadly oval (almost globular), their colour in life is
not recorded; BL 1.7-2.1 mm. In galls
on leaves of Apocynum (according to
label on type slide in BMNH collection, although host is given as
"Apocynaceae" in Essig (1958); host plant and locality data were
given in error under Eonaphis pauliani
(see Hille Ris Lambers 1966b). Alatae
have thickly-bordered forewing veins.
Only known from Madagascar. The
life cycle is unknown.
Two species in South
America on roots of Senecio,
related to Aphis but with reduced
siphunculi placed on abdominal tergite 6, reduced triommatidia, an unusual
distribution of lateral tubercles, and distinctive biology. One species is
known only from trapped alatae (P.
americana, previously described in the genus Seneciobium). Pehuenchaphis agilissima Mier Durante, Nieto Nafría & Ortego Apterae are dark green to brown, sometimes
wax-dusted; BL 1.5-2.4 mm. Apterae have secondary rhinaria distributed III
14-18, IV 0-10. On Senecio spp.,
living in small ant-attended groups on thick, lignified roots, and at bases
of lignified stems, moving quickly when disturbed. Monoecious holocyclic with
apterous males (Mier Durante et al.
2003).
One species in North
America distinguished by its dark sclerotic carapace, with abdominal tergites
2-6 fused together, and by the long dorsal hairs which often have furcate
apices. Peltaphis hottesi Frison & Ross Apterae are dark brown, almost black; BL
c.2.1-2.3 mm. On Carex spp., only
known from Illinois and Wisconsin, USA and Quebec, Canada. Alatae are unknown. Monoecious holocyclic,
with oviparae and apterous males in October-November (original description).
Over
70 species have been described, of which 45 are known to form galls on leaves
or twigs of Populus (subgenus Eupopulus). Alatae emerging from these galls have a
black head and pterothorax and a rather elongate, yellow-green to grey-green,
wax-dusted, abdomen. The life cycles
of 18 of these gall-formers are known; 4 are monoecious on Populus, the alate sexuparae being
produced in the galls, and in the other 14 the alate progeny of the fundatrix
migrate to various herbaceous mainly dicotyledonous secondary hosts in
Amaranthaceae, Apiaceae, Brassicaceae, Compositae/Asteraceae, Euphorbiaceae, Leguminosae/Fabaceae and Ranunculaceae, and
where they found colonies on the roots, or occasionally in woolly wax masses
above soil level. Most of the species
known only from their root-feeding generations feed on herbs in the
above-named families, but at least one species feeds on Salix roots. Many species are recognised as distinct only from the morphology of
the galls and emigrant alatae produced on Populus,
or by experimental host transfers that show specificity to particular
secondary hosts. Identification of apterae from roots
using morphological criteria is generally difficult or impossible, so host
records to species level based on apterae alone are almost always suspect.
Zwölfer (1958) provided morphometric parameters for populations of
unidentified Pemphigus spp. on
roots of Poaceae and various dicots in southern Germany. Biology and ecology
of most root-feeding populations are little-known; a Pemphigus species from Gnaphalium
in India (western Himalaya; leg. S Chakrabarti), sent to BMNH for
karyotyping in 1989 was heavily parasitised by an unidentified mermithid
parasite (Blackman& Eastop 2006, p.1261; this is the first published
record of a mermithid-aphid host association). The species
are distributed rather evenly throughout the northern hemisphere; 17 are described from
Europe, 10 from the Middle East and Central Asia, 16 from east Asia, and 21
from North America. Accounts are available for Britain
(Furk & Prior 1975); European gall-makers on Populus nigra (Lampel 1960); Fennoscandia and Denmark (Heie
1980b); the
UK (Blackman et al. 2019b); the Iberian
peninsula (Nieto Nafría et al.
2002a); India (A.K. Ghosh 1984b); East Siberia (Pashchenko 1988a; Babichev
& Kirichenko 2020); Japan (Aoki 1975); China (Zhang & Zhong 1979c, G.
Zhang & Zhong 1983b, G. Zhang et
al. 1999c); and North America (Palmer 1952, Harper 1959a, Lange 1965,
Smith 1985). Dunn (1960a) and Faith (1979) investigated gall formation. Halaj & Osiadacz (2013)
reviewed the European gall-forming species and provided an illustrated key to
their galls on Populus. Alvarez et al. (2020) compared the histology
of seven Iberian species and related it to their phylogeny. N.
Pike et al. (2007) published a
phylogeny for nine species based on mtDNA sequences. A recent study by L. Liu et al. (2014) of relationships within
and between Pemphigus and allied
genera using mitochondrial, nuclear
and symbiont DNA has shown that the genus is paraphyletic, with P. matsumurai and P. sinobursarius forming a separate clade basal to one that
includes five other Pemphigus plus Thecabius and Epipemphigus. Pemphigus
betae Doane (= balsamiferae
Williams) Galls are yellowish green
sometimes tinged with red, pocket-like, formed usually on the underside of
the poplar leaf near its base, on or adjacent to the mid-rib, with little or
no deformation of the leaf (Harper 1959a, Whitham 1978). The opening can be either on the upper side
of the leaf or underneath along the mid-rib.
Primary hosts are Populus
angustifolia, P. balsamifera,
or occasionally P. deltoides ssp. occidentalis or P. trichocarpa. Alatae (BL
1.5-2.1 mm) leave galls in late June-early August (Harper 1959b), to colonise
roots of Beta vulgaris, and also
perhaps on other Amaranthaceae (formerly Chenopodiaceae) such as Chenopodium album, and possibly Spinacia. Apterae on chenopods are yellowish aphids
secreting white wax, with BL 1.9-2.4 mm. Anholocyclic
populations can persist all year on secondary hosts (see Moran & Whitham
1988), and are widely distributed in North America. The life cycle was
studied by Harper (1963) in southern Alberta, and by Moran & Whitham
(1988) in Utah. This aphid's
evolutionary ecology has been studied in some detail, including the
territorial behaviour of first instar fundatrices in selection of galling
sites (Whitham 1979), population dynamics in relation to galling site
(Whitham 1978, 1980), genetics of resistance in the host plant (Paige &
Capman 1993), induction of sexuparae (Moran et al. 1993a), and discrimination between host trees by returning
sexuparae in autumn (Moran & Whitham 1990). However, interpretation of all this work is
problematic as Foottit
et al. (2010a) have demonstrated
that there is a complex of Pemphigus species
in western North America sharing the
same cottonwood primary hosts. Taxonomic affinities with palaearctic
populations of Pemphigus on
chenopod roots (see P. fuscicornis)
are unclear. See also Blackman & Eastop (2000). Pemphigus
birimatus Ivanoskaya
Galls on Populus laurifolia in
northern Kazakhstan are irregular in shape, with two slit-like openings, and
are formed on the leaf petiole (Ivanoskaya 1973). Also recorded on P. talassica, and from western Siberia
(Kadyrbekov 2017a) and eastern Siberia (Babichev & Kirichenko 2020). The
emigrant alata (BL 2.8 mm, with strap-like secondary rhinaria distributed III
11, IV 5, V 4, VI BASE 7), could not be included in the key on the basis of
the published description. Pemphigus
borealis Tullgren Galls
on Populus laurifolia are green,
elliptical, hazelnut-sized, usually in groups of 2-10 on stem at bases of
twigs or leaf petioles (fig. 131K). P.
laurifolia seems to be the preferred host (Dolgova 1970), although there
are also records from P. balsamifera
(planted), P. iliensis, P. koreana,
P. nigra, P. simonii and P. suaveolens. The fundatrix is grey-green with slight
yellowish tinge. Alatae (BL 1.5-2.5
mm) emerge in June-August and migrate to found colonies on the roots of Bidens spp., these colonies consisting only of immature sexuparae
(Dolgova 1970). Adult alate sexuparae are yellowish, with tufts of wax, BL
c.2 mm, and migrate back to Populus
in August-September. P. borealis occurs throughout the
northern palaearctic, from Sweden to China and Japan. A record from England
is possibly ascribable to P. trehernei
(Foster 1975). The recorded occurrence of root-feeding populations in Idaho,
USA (Griffiths et al. 1976) also
requires confirmation. In east Asia
there is probable confusion with P.
dorocola (q.v.), which is described from very similar galls on P. maximowiczii (=P. suaveolens) and is
possibly a synonym (Aoki 1975). P. yangcola, described from Populus sp. in China (Zhang &
Zhong 1979c), is also very similar and a possible synonym. 2n=20.
Pemphigus brevicornis (Hart) Apterae are pale, with wax; BL c.1.8 mm.
Described from apterae on corn (Zea)
roots in Illinois, USA, and since recorded from roots of various plants in
Illinois (Hottes & Frison 1931) and other states, but there are no good
criteria for identification. Alate sexuparae have secondary rhinaria
distributed III 5-7, IV 1-4 (according to Hottes & Frison 1931). Pemphigus
bursarius (Linnaeus)
Poplar‑Lettuce Aphid, Lettuce Root Aphid Galls on Populus spp. (most commonly P. nigra) are yellowish or reddish
when mature, purse-shaped, always formed on leaf petioles (fig. 131L,
and see
influentialpoints.com/Gallery). The fundatrix is greyish green, slightly
wax-dusted. Alatae (BL 1.6-2.5 mm)
leave the gall through a lateral opening over an extended period from late
May to September, with peak emergence in late June-July (northern
hemisphere). They found colonies mainly on roots of Compositae/Asteraceae (e.g. Cichorium, Lactuca, Lampsana, Sonchus, Taraxacum, Tussilago).
Many of the records of P. bursarius
from roots of plants in other families may be referable to other Pemphigus spp. Apterae on roots of secondary hosts are yellowish
white with a tuft of white wax on the posterior part of the abdomen; BL
1.6-2.5 mm (Plate 2g).
Alatae from root‑feeding colonies (sexuparae) have a brownish‑orange
abdomen and secondary rhinaria distributed III 5-10, IV 1-4. Return
migration of sexuparae to poplar is in October-September, but anholocyclic
overwintering on roots of secondary hosts is common. P. bursarius occurs in Europe, across Asia to eastern Siberia
(Babichev & Kirichenko 2020), northern and southern Africa, North and
South America, and (perhaps) Australia and New Zealand. There is some
uncertainty about its occurrence in Australia and New Zealand because,
although bursarius-like aphids occur in bursarius-type galls on Populus
and are also recorded from Sonchus roots,
the secondary host populations are mainly recorded from Amaranthaceae and
Polygonaceae.
Miller et al.. (2005) found
genetically distinct populations on different secondary host plants,
indicating a complex of races and/or subspecies. Dunn (1960a)
studied gall formation and (1960b) reviewed natural enemies. Alleyne & Morrison (1978a, b) gave
accounts of its biology and natural enemies in Quebec. See also Blackman
& Eastop (2000). 2n=20. Pemphigus
circellatus Zhang & Zhong Large galls are formed on the branches of Populus davidiana in China
(Yunnan). Emigrant alatae (BL c. 1.8
mm) were collected in May (Zhang & Zhong 1985a). The life cycle is unknown. Pemphigus
cylindricus Zhang Only fundatrices are described, from a gall
on an unidentified Populus sp. in
Tibet (Zhang & Zhong 1981). Pemphigus
dorocola Matsumura
The galls on twigs of Populus
maximowiczii (=P. suaveolens)
in Japan are pouch-shaped, reddish when mature, with a transverse apical slit
(Aoki 1975). Also recorded from P.ciliata, P. koreana and P. suaveolens. The fundatrix is green. Alatae (BL 1.9-2.3
mm) emerge over an extended period (late July-November). The secondary host is unknown, and
Chakrabarti (2001) suggested that it may not host-alternate. In Japan, Korea, east Siberia (Pashchenko
1988a) and northern India (Chakrabarti 2001). Aoki (1978, 1980a) described
aggressive and gall-cleaning behaviour by first instar larvae, and Aoki &
Kurosu (1991d) described an aptera producing abnormal progeny in the gall. [P.
dorocola is separated in the key to gall-formers on Populus according to the description by Aoki (1975), who does not
recognise the presence of P. borealis
in Japan. Alatae in the BMNH
collection from galls on Populus
suaveolens in Japan and Korea all have antennal sensoriation like that of
P. borealis, and further work may
show P. dorocola to be a synonym of
P. borealis, as suggested by
Takahashi (1920).] 2n=20. Pemphigus
echinochloaphaga (Zhang, Qiao & Chen)
Appearance of apterae in life is unknown; BL 1.9-2.2 mm. On “grass” (Echinochloa sp.?) in China (Zhang et al. 1999, as Desiforda). Pemphigus fatauae Shinji
Apterae are undescribed, alatae (sexuparae?) from roots of Fatoua (as Fatauna) pilosa in Japan
had secondary rhinaria distributed ANT III 6, IV 3-4, V 1. These may possibly
have been sexuparae of Epipemphigus
niisimae. Pemphigus fuscicornis (Koch) Apterae are pale yellow or yellow-green
to greyish green with wax tuft on posterior abdomen; BL 1.4-2.2 mm. On roots
of Tripleurospermum inodorum (= Matricaria inodora) in Europe (UK,
Denmark, Sweden, Germany, Finland) and Central Asia (Kazakhstan; Kadyrbekov
2017a). Originally also described from Chenopodium
roots, and this name has therefore also been applied to very similar aphids
found on Amaranthaceae (Atriplex, Beta, Chenopodium) in central and eastern Europe, and also in Iran and
Central Asia (Kadyrbekov 2017a); however, these might be an anholocyclic
population of P. betae introduced
from North America (Blackman & Eastop 2000). Anholocyclic overwintering
occurs on Tripleurospermum; alatae
sexuparae (with secondary rhinaria distributed III 5-8, IV 1-3) are produced
in autumn, but the sexual phase and primary host are unknown. Studies of
microsatellite DNA variation in P.
bursarius by Miller et al. (2005)
may have included this species. 2n=20. Pemphigus
gairi Stroyan
One of three palaearctic species producing similar pouch-shaped galls
on or near the mid-rib on the upper sides of leaves of Populus nigra (incl. var. italica),
with an opening to the underside of the leaf (the other two species are P. phenax and P. populinigrae). The fundatrix is dull green or greyish green
under a thick covering of wax (see
influentialpoints.com/Gallery). Alatae (BL 1.5-2.2 mm) emerge
in June-July and migrate to colonise the roots of Aethusa cynapium (Stroyan 1964b). Apterae of the root-feeding generation are
pinkish ochreous or pale straw-coloured; BL 1.3-2.0 mm. Sexuparae
(with
secondary rhinaria distributed III 4-7, IV 2-4) return in October.
Described from England, and also recorded from Czech Republic, where Urban
(2004) reported on the development and natural enemies of its gall
generations on P. nigra, and from
Ukraine. There seem to be no other literature records of this species,
possibly due to confusion of its galls with those of P.
phenax and P. populinigrae, as
it is presumably more widely distributed in Europe. Pemphigus groenlandicus
(Rübsamen) Colour of apterae in life is unrecorded,
secreting wax; BL 1.5-2.1 mm. Alate sexuparae have secondary rhinaria
distributed III 3-8, IV 2-4. On roots of unidentified Poaceae in Greenland,
where it is presumably anholocyclic, although producing sexuparae (Hille Ris
Lambers 1952). Zwölfer (1958) noted that populations on roots of various
grass species in Germany (Agrostis,
Arrhenatherum, Avena, Cynosurus)
agreed well with this species. There
are subsequent records from France and Spain (Nieto Nafría et al. 2002a), the morphology agreeing
best with that of the Greenland population described as P. groenlandicus crassicornis
Hille Ris Lambers. Differences from P. similis Börner, recorded from ?Agrostis alpina, Nardus stricta and Poa laxa
in the Austrian Tyrol, and from P. vulgaris Raychaudhuri, Pal & M.R. Ghosh, described from Triticum aestivum and unidentified
grasses in Sikkim, and West Bengal, India, have not been established. Pemphigus hydrophilus Narzikulov
Apterae are yellowish green, secreting flocculent wax; BL 1.9-2.2 mm.
One alata (sexupara?) had secondary rhinaria distributed III 6, IV 5, V 2. On
the moss Cratoneurum filicinum, and
possibly also on Hygrohypnum luridum.
In Tajikistan at 1600-1700 m. Pemphigus
ignotus Habib & Ghani
Recorded as forming leaf galls on Populus
ciliata in Pakistan (Habib & Ghani 1970), but no description is
given, and this is therefore a nomen
nudum. Pemphigus
immunis Buckton (= lichtensteini Tullgren) The galls are quite large (up to 4 cm
maximum diameter), greenish brown or reddish, walnut-like in shape and
surface texture, thick-walled, with an apical opening, on twigs or branches
of Populus spp., especially nigra and euphratica (fig. 131M,
and see
influentialpoints.com/Gallery). The fundatrix is glaucous
green (Roberti 1938). Alatae (BL
2.2-2.5 mm) emerge from late April to early August and found colonies of
apterae (BL 1.2-1.8 mm) secreting copious wax on roots of annual Euphorbia spp. (helioscopia, peplus;
Hille Ris Lambers 1973b). A.K. Ghosh (1984) reviewed its biology. In Europe,
north Africa, south-west and central Asia, Pakistan, north-west India, and
China (Tseng & Tao 1936; as P.
napaeus). A record from Euphorbia
in USA (as brevicornis) may also be this species. Habib
& Ghani (1970) studied natural enemies in Pakistan, and Bhagat (1982a)
reported a parasitoid and hyperparasitoid in Kashmir. 2n=20 (or 2n=10
according to Pal & Khuda-Bukhsh, 1982). Pemphigus
iskanderkuli Narzikulov Galls on Populus dentata when mature are pinkish to red, rounded, smooth,
walnut-sized, with apical opening, situated on upper leaf surface at junction
of petiole and lamina (Narzikulov 1957).
BL of alata 2.7-3.5 mm. In
central Asia (Tajikistan, Kirgizia).
The life cycle is unknown. The
alatae seem indistinguishable from those of P. napaeus, and Doncaster (1969) suggested that the two might be
conspecific, but for reported differences in the morphology of the
fundatrices, and position and colour of the galls. Populations in Kirgizia on P. laurifolia described by Gabrid
(1989) as Pemphigus sp. aff. iskanderkuli would appear to be large
specimens of iskanderkuli, although
the galls on P. laurifolia
illustrated by Gabrid appear very irregular in shape. Pemphigus
knowltoni Stroyan
Galls are green, pouch-like, of variable shape, on undersides of
leaves of Populus angustifolia
alongside the mid-rib at the base of the lamina (Stroyan 1970). Fundatrix and immatures in gall are
blackish green. Alatae (BL 1.8-2.1 mm)
are mature in the gall in July, and migrate to an unknown secondary
host. Described from Utah, USA. Pemphigus
laurifolia Dolgova
Galls occur on Populus
laurifolia in the Altai region of Russia.
Alatae (BL c. 2.1 mm) emerge in July (Dolgova 1973). It has also been reported to occur on P. talassica in northern Tien-Shan
(Kadyrbekov 2013f) and in eastern Siberia (Babichev & Kirichenko 2020).
The original description is not available; this species is keyed on the basis
of alate specimens (paratypes?) collected by Dologova on 1.vii.71 (BMNH
collection, leg. D. Hille Ris Lambers). [Note that Pemphigus laurifoliae Holman & Szelegiewicz 1972 (nomen nudum) is P. mongolicus Holman & Szelegiewica 1974a)]. Pemphigus
longicornus Maxson Galls are elongate, thin-walled, on upper
sides of leaves of Populus deltoides. The fundatrix is pale yellowish white. Alatae (BL 1.7-2.1 mm) emerge in May-June
and migrate to an unknown secondary host (Maxson 1923). In eastern and southern USA. [According to Smith (1985), there is also
an undescribed Pemphigus sp. on P. deltoides in North Carolina that
causes elongate to slightly globular galls on the upper sides of the leaves,
and has alatae with antennae intermediate in length and antennal sensoriation
between longicornus and populivenae.] Pemphigus
mangkamensis Zhang Described from apterous viviparae from an
unidentified Populus sp. in Tibet
(Zhang & Zhong 1981b), and said to be similar to P. chomoensis. If the
specimens are not fundatrices this would indicate absence of host
alternation. Pemphigus
matsumurai Monzen
The gall is of globular or irregular shape, at base of mid-rib on
upper side of leaf, concolorous with mid-rib or slightly reddish (Aoki 1975,
Zhang & Zhong 1983b, Chakrabarti 2021).
Recorded from Populus ciliata,
P. koreana, P. laurifolia, P. nigra, P.
talassica and P. suavolens. The fundatrix is green. Alatae (BL 1.7-2.7 mm, fig.
86C)
migrate in June-July to colonise roots of Thalictrum
spp. In east and Central Asia
(Kazakhstan, northern India, Pakistan, Bhutan, Mongolia, east Siberia, China,
Japan). Also in Tajikistan, as Pemphigus
montanus, described from Populus
tadzhikistanica (Narzikulov 1957), is clearly a synonym. Aoki &
Kurosu (1989) studied the influence of parasitisation of the fundatrix on the
development of the gall and subsequent generations, and Takada et al. (2010)
studied the effects of a parasitoid and three hyperparasitoids. Bhagat's (1982a) record of a parasitoid of
P. immunis possibly should be
referred to matsumurai, as this is
the only species known to form leaf galls on P. ciliata. DNA studies by
L. Liu et al. (2014) place this
species, together with P. sinobursarius, in a basal position in a phylogeny
that includes Thecabius and Epipemphigus. 2n=12 (an unusual
chromosome number for a Pemphigus, confirmed for Japanese
samples from Thalictrum). Pemphigus
microsetosus Aoki
A large closed pouch-gall is formed as an outgrowth of the leaf of Populus suaveolens, taking in the
whole leaf when mature; lignified galls remain on the tree after leaf-fall
(Aoki 1975). The fundatrix is
yellowish green. Alatae (BL 2.1-2.5
mm) are adult from late July and leave gall through several secondary exit
holes, migrating to an unknown secondary host. In Japan, and since recorded from west
Siberia (Stekolshchikov & Novgorodova 2015) and east Siberia (Pashchenko
1988a, Babichev & Kirichenko 2020). The latter authors record the host in
eastern Siberia as P. laurifolia.
2n=22 (Blackman 1986). Pemphigus
mongolicus Holman & Szelegiewicz The gall is globular, pouch-like, on upper
side of leaf of Populus laurifolia
near the base of the mid-rib; green, becoming tinged with red when mature
(Holman & Szelegiewicz 1974a). The
fundatrix is palish green, producing a generation of dark green apterae in
the galls. Monoecious; alate sexuparae
(BL 1.6-2.0 mm) leaving galls in early August give birth to sexuales in bark
crevices on the trunk. In Mongolia and
probably in eastern Siberia (Babichev & Kirichenko 2020). ["P.
semenovi", noted by Mordvilko (1935) as monoecious on P. suaveolens in Siberia but
apparently a nomen nudum, is
probably this species.] Pemphigus
monophagus Maxson The
gall is globular, pale greenish yellow, consisting of a swelling of upper
side of base of leaf lamina with petiole broadened and somewhat twisted, and
a slit-shaped opening underneath between petiole and leaf (fig. 131N,
and Maxson 1934). Fundatrix
honey-yellow to pale olive-green, producing a generation of creamy white
apterae in the gall. Monoecious,
alatae (BL 1.4-1.9 mm) emerging from galls in August-September to deposit
sexuales on the bark of the trunk. On Populus angustifolia, P. balsamifera and P.trichocarpa in western North
America. Aoki & Kurosu (1988)
discussed the evolution of monoecy in this species. Pemphigus
mordvilkoi Cholodkovsky
Galls, formed on the twigs of Populus
ciliata, are smooth, globular, sessile, yellowish green to dark greenish
brown (Chakrabarti et al. 1985; Chakrabarti 2021). The fundatrix is pale green. Chakrabarti (2001) noted that its first
instar progeny were dimorphic, one form apparently exhibiting the
characteristics of “soldiers”. The life cycle is unusual, and requires
further study. Fundatrices have a long life (Chakrabarti & Das 2014). In
Pakistan, alatae emerge from the galls over an extended period from August
until January of the next year (BMNH collection). According to A.K. Ghosh et al. (1981), the alatae emerging
from galls just before onset of winter in north-west India hibernate in bark
crevices of the same tree, or in other sheltered places; however, these
alatae are not sexuparae, so the monoecious life cycle suggested by them
seems unlikely. Apterous exules and
sexuparae likely to be those of P.
mordvilkoi have been collected in Punjab in April on roots of Helipterum (= Acroclinium) roseum (Compositae;
BMNH collection, leg. A.S. Sohi); similar sexuparae have also been collected
on other Compositae in other parts of northern India and Nepal in March-April
(BMNH collection), and sexuparae
and sexuales were observed on P.
ciliata in Pakistan in April-May (Habib & Ghani 1970). The oviparae laid eggs which hatched in 6-9
days, and the fundatrices formed galls in May-June. Thus a two-year life cycle with alternation
between P. ciliata and Acroclinum is indicated. 2n=20. Pemphigus
nainitalensis Cholodkovsky Galls are about the size and shape of a
hazelnut, sessile, on twigs of Populus
ciliata, which are thickened in the region of the gall (Cholodkovsky
1912). Only known from the original
descriptions of the fundatrix and the alata, from galls collected in Uttar
Pradesh, India. Pemphigus
napaeus Buckton
Galls on twigs of an unidentified Populus
sp. were smooth, roundish or of irregular shape, shining green "slightly
veined with brown, and variegated with yellowish patches" (Buckton
1896); but see comments by Doncaster 1969, p. 162). In the Northern Areas of Pakistan
(Gilgit). There are no other
authenticated records, and contrary to some authors (eg. A.K. Ghosh et al. 1981), the host species is
unidentified. Reports of napaeus in Taiwan (e.g. Tseng &
Tao 1936) should probably be referred to immunis. Doncaster (1969) remounted and redescribed
the type material. See also under P. iskanderkuli. Pemphigus
nortonii Maxson
Galls on Populus deltoides
especially var. occidentalis (= monilifera) are pale yellowish green, formed
by a swelling at the base of the leaf lamina, twisted on the flattened,
broadened petiole; resembling those of P.
populicaulis (Palmer 1952), but with a long slit-like opening (Harper
1959a). Alatae emerge over an extended
period (June-Sept; Harper 1959b, Grigorick & Lange 1968), migrating to an
unknown secondary host. Apparently
widely distributed in USA, and in Alberta, Canada. Apart from the greater number of rhinaria
on ANT V and VI this species closely resembles P. junctisensoriatus.
Populations from Alberta (Harper 1959a, b) seem to be intermediate in
this respect, and the distinction between these two species needs to be
verified. Pemphigus
obesinymphae Aoki & Moran Galls are formed on the petioles of late
flushing leaves of Populus fremontii (Aoki
& Moran 1994), and P. deltoides (Abbot
2009), usually close to the leaf blade. When mature they are almost
spherical, 8-16 mm in diameter, with the lips of the transverse slit
sometimes bulging outwards, and a circular ostiole about 1.2 mm in diameter
somewhere along the slit. First instar progeny of the fundatrix in the gall
are heavily sclerotised with well-developed fore legs and attack predators
(Moran 1993). Abbott (2009) demonstrated using microsatellite DNA analysis
that there was a high level of clonal admixture within galls, and Grogan et
al. (2010) investigated its consequences. Alatae (BL
1.9-2.5 mm) fly in August-October, presumably migrating like P. populitransversus to cruciferous
hosts (N. Chen et al. 2009), on
which overwintering occurs. Sexuparae return to Populus in May of the following year, and the eggs hatch only a
short time after oviposition. Apparently widely distributed in USA (Ohio,
Tennessee, Arizona, Utah, Texas), and probably in northern Mexico. N. Chen et al. (2009) recorded molecular and
phenological differences between P.
obesinymphae and P. populitransversus on cabbage in
Texas. This species was confused for many years with P. populitransversus (q.v.), and its distribution and range of
both primary and secondary hosts requires further confirmation. Pemphigus
passeki Börner
Galls are formed on leaves of Populus
nigra (incl. var. italica) by
swelling of the basal half of the mid-rib, the swelling being broadest
towards the junction with the petiole (fig. 131O,
and Prinsen 1990). Alatae (BL c. 2.1
mm) fly in June-August to colonise roots of caraway (Carum carvi). It is very similar to P. phenax but can be separated on the basis of morphological differences
in the alate sexupara. In northern palaearctic; north-west,
north and central Europe, Russian Altai region (BMNH collection, leg. L.P.
Dolgova) and eastern Siberia (Babichev & Kiirichenko 2020). The spring migration was studied by Prinsen
(1991). 2n=22. Pemphigus
phenax Börner & Blunck One of a complex of species in Europe
producing similar mid-rib galls on upper sides of leaves of Populus nigra (incl. var. italica), with a slit-like opening
underneath the leaf. The galls of P. phenax are reddish, tend to be
larger than those of P. gairi, and
more elongate (spindle-shaped) than those of P. populinigrae, somewhat more wrinkled, and often tinged with
yellow laterally (Stroyan 1964b). The
fundatrix is dull pale green when adult.
Alatae (BL 1.5-2.4 mm) fly in June-August to found colonies on roots
of wild and cultivated carrot (Daucus
carota). Apterae on carrot are pale lemon yellow to
yellowish‑white with white wax; BL 1.7-2.6 mm. Populations may also
persist parthenogenetically on carrots remaining in the ground through the
winter. In northern
Europe (England, Denmark, France, Germany, Netherlands, Sweden), Turkey (Şenol
et al.
2014) and west Siberia. For a more detailed account see Stroyan (1964b). Pemphigus
plicatus Dolgova
Galls occur on Populus
laurifolia in the Russia (Altai region, western and eastern Siberia, and
are also recorded from P. laurifolia
and P. nigra in Kazakhstan
(Kadyrbekov & Aoitzhanova 2005).
Alatae (BL c. 1.8 mm) were collected in July. The original description is not available.
[It is included in the key on the basis of specimens collected by Dolgova on
1.vii.71 (BMNH collection, leg. L.P. Dolgova.)] Pemphigus
(Pemphiginus) populi Courchet Galls on Populus spp., especially P.
nigra (incl. var. italica) are
globular outgrowths of the mid-rib, not larger than a hazelnut, becoming
club-shaped with the basal part narrower than the apex when mature, situated
near the base on the upper side of the leaf (see influentialpoints.com/Gallery). Alatae (BL 1.4-2.5 mm, fig.
86D)
emerge in June-July through a secondary opening at the top of the gall, migrating
to found colonies on roots of Melilotus
altissima (Furk & Prior 1975), and certain other Leguminosae (Lathyrus, Medicago). Apterae on secondary
hosts are pale yellow-green, with white wax; BL 1.9-2.6 mm. Sexuparae
(with secondary rhinaria distributed III 4-7, IV 1-2) return to P. nigra in September-October. In Europe, south-west and central Asia, and
eastward to east Siberia (Babichev & Kirichenko 2020) and China (Zhang
& Zhong 1979c). Furk & Prior
(1975) reported parasitisation of fundatrices by Monoctonia pistaciaecola. Coffin & Remaudière (2001) compared
the life cycle and morphology with P.
vesicarius. N. Pike et al.
(2007) studied the phenology of the gall generations. For a general account
see Osiadacz & Halaj (2014). Pemphigus
populicaulis Fitch The gall is yellowish green to creamy,
formed by swelling of base of leaf lamina, twisted around broadened petiole,
rather like that of P. nortonii and
P. populiglobuli but with a small,
rounded exit hole on the underside (Palmer 1952, Harper 1959a). Especially associated with P. deltoides and P. deltoides ssp. occidentalis
(= monilifera), but also recorded
from other cottonwoods and P.
tremuloides. The fundatrix is pale
yellowish to whitish green. Alatae (BL
1.7-2.0 mm) emerge in July-September.
The secondary host has not definitely been established; Glendenning
(1924) reported that it was Oenanthe
sarmentosa, but did not provide evidence for this assertion. Hottes & Frison (1931) suggested that P. brevicornis Hart, described from
corn roots, was the sexupara of populicaulis. Widely distributed in North America. Setzer (1985) studied gall mortality. Sokal et
al. (1980) and Bird et al.
(1982) studied geographical variation in eastern North America using
multivariate techniques. Pemphigus
populiglobuli Fitch The gall is green, often tinged with red
and/or yellow when mature; globular, conical or bullet-shaped, formed by
swelling of base of leaf lamina and thickening of petiole, the exit hole
being a slit on the underside between leaf and petiole (Maxson & Knowlton
1929). Primary hosts are Populus angustifolia and P. balsamifera, more rarely P. trichocarpa. The fundatrix is variously reported to be
dark green (Palmer 1952) or pale yellowish white (Maxson & Knowlton
1934). Alatae (BL 1.5-2.3 mm) leave
gall in June-early July (Alberta; Harper 1959b) for an unknown secondary
host. Widely distributed in North
America. Differences from P. populicaulis and other
closely-related species are discussed by Maxson & Knowlton (1929). Pemphigus
populinigrae (Schrank) (= filaginis
Boyer de Fonscolombe) Galls on Populus spp., especially P. nigra, are pouch-shaped (fig. 131P),
similarly situated to those of P.
phenax and P. gairi, on
upperside of leaf lamina alongside mid-rib with opening along original line
of invagination on underside of leaf.
Gall formation was described by Dunn (1960). Mature galls are rounded and smooth on top,
and of a dull reddish colour without much tinge of yellow (Stroyan
1964). The fundatrix is dark
green. Alatae (BL 1.8-2.9 mm) emerge
in June-August and migrate to found colonies above ground on Filago, Logfia and Gnaphalium,
often in the tomentum of the inflorescences.
Apterae
on secondary hosts are yellow-green, with white wax-wool; BL 2.0-2.2 mm. Sexuparae
(with secondary rhinaria distributed III 7-9, IV 2-4) return to P. nigra in September-October, but
anholocyclic overwintering apparently occurs where the primary host is not
available (Mordvilko 1935). Throughout
Europe and in north Africa, south-west Asia, Central Asia and across to east
Siberia (Pashchenko 1988a). Records
from China (Tao 1958) may be due to confusion with Epipemphigus niisimae.
2n=22. Pemphigus
populiramulorum Riley Galls are formed by swelling of new growth
of twigs, or rarely at the bases of petioles; they are quite large (up to 2.5
cm diam.), rounded, green to brownish, with a transverse or vertical
slit-shaped opening which often has protruding lips. The primary host is usually Populus deltoides ssp. occidentalis (= monilifera), but it is also recorded from P. balsamifera and P.
fremontii. The fundatrix is pale
creamy yellow (Palmer 1952). Alatae
(BL 2.2-2.5 mm) leave galls in July-September (Harper 1959b), flying to an
unknown secondary host. In western
North America (Utah, Colorado, South Dakota). Pemphigus
populitransversus Riley
Plate 6c The gall is formed by swelling and bending
of petioles of the leaves of Populus
deltoides (incl. ssp. occidentalis)
and P. fremontii; it is green,
globular to ovoid, with a usually transverse slit (see influentialpoints.com/Gallery). Sometimes recorded from other Populus spp. (acuminata, angustifolia,
balsamifera, nigra, tremuloides, trichocarpa) However, some of these records,
particularly those from western cottonwoods, may apply to other species. The fundatrix is whitish to pale dirty
greenish yellow (Palmer 1952). Alatae
(BL 1.7-2.0 mm) emerge in July and found colonies of pale yellow aphids in
flocculent wax on roots of Brassicaceae (Jones & Gillette 1918), where it
can be a serious pest (Royer & Edelson 1991). Sexuparae return to cottonwood in
September-October to produce sexuales which lay overwintering eggs on the
bark. Geographic variation in eastern USA has been studied extensively using
multivariate methods (Sokal & Riska 1981). Bingham & Sokal (1986) studied host
tree effects on morphology. P. populitransversus is recorded
throughout most of North America east of the Rocky Mountains. However it is
still unclear whether the name is being applied to a single species; apart
from the confusion with P. obesinymphae
(q.v.), Bird et al. (1979) found
that two forms with different gall shape and phenology, and corresponding
morphological differences in fundatrices and emigrant alatae, occur
sympatrically on P. deltoides over
a large area of the USA east of the continental divide. It should be noted
that the morphology of their aphids from globular galls on late flushing
leaves does not correspond with that described for P. obesinymphae. Pemphigus
identified as P. populitransversus
have been found on Populus sp. in
Mexico (BMNH collection, leg. R. Peña) and Chile (BMNH collection, leg. G.
Monsalve); it has also been introduced to South Africa (Müller & Scholl
1958), where it occurs on Populus nigra
(BMNH collection, leg. E McC. Callan) and P.
deltoides, and the Azores (on Populus
sp.; BMNH collection, leg. A. van Harten).
Presumed anholocyclic populations also occur in New Zealand on Eutrema wasabi roots (BMNH collection,
leg. L.M. Emms), and aphids collected on Arabis
roots in England (BMNH collection, leg. G. Fox-Wilson) may also be this
species. 2n=20. Pemphigus
populivenae Fitch Galls are elongate pocket-shaped or cock's
comb-shaped, greenish or yellowish or pale red, arising alongside mid-rib on
upper side of leaf, with a slit-shaped opening underneath. On heavily-infested trees, galls may occur
on both sides of the leaf, and those on the underside may then be confused
with those of P. betae (Maxson
& Knowlton 1934; however see also Footit et al. 2010). Whitham (1978) reported that P. populivenae galling Populus
fremontii in Utah caused chlorosis of the leaf distal to the gall. As in
the case of P. betae, several
species of cottonwoods and their hybrids are colonised (angustifolia, balsamifera,
deltoides, fremontii and trichocarpa;
the latter species in experimental conditions only). The fundatrix is yellowish green. Alatae (BL 1.1-1.8 mm) emerge in June-July
and migrate to found colonies on roots of Amaranthaceae. Rumex
(Polygonaceae) may also be colonised. The evidence for this host
alternation is work in California by Grigorick & Lange (1962), who
transferred sexuparae from sugar beet to poplar bark under controlled
conditions and obtained fundatrices that produced galls specifically on P. trichocarpa. The alatae from these galls were identified
as P. populivenae, and colonised
roots of B. vulgaris and Chenopodium album. However, the suggested synonymy of both P. betae and P. balsamiferae with populivenae
indicates some confusion of identity, and further work seems advisable to
confirm that the correct name for populations damaging sugar beet in
California (e.g. Summers & Newton 1989) is in fact populivenae rather than betae. Foottit et
al. (2010) have found molecular evidence that at least three taxa with
similar galls occur sympatrically on cottonwoods in Alberta. Pemphigus
protospirae Lichtenstein Galls on Populus spp., especially P.
nigra var. italica, are smooth,
shiny, green mottled with red, caused by swelling, flattening and spiral
twisting of the leaf petiole, similar to those of P. spyrothecae but often thinner and with more spirals (see fig. 131R
and influential
points/Gallery). P.
protospirae, however, is heteroecious, and the alatae leave the gall in
late May-early July to found colonies usually in the sheathing leaf-bases of
aquatic Apiaceae (Berula erecta, Sium latifolium, Apium nodiflorum, Oenanthe aquatica). Apterae on secondary hosts are pale yellowish
green or yellow, with abundant white wax especially on posterior abdomen; BL
1.6-2.3 mm. The return to Populus occurs in September-October. Widely
distributed in Europe, but probably restricted by availability of secondary
host habitats. Also recorded from west
Siberia, Iran, Central Asia, eastern Siberia (where it is particularly
asociated with P. laurifolia
according to Babichev & Kirichenko 2020) and China (Zhang & Zhang
1979c). Pemphigus rubiradicis Theobald
Apterae are dirty greenish brown, with flocculent white wax on
abdomen; BL 1.7-2.2 mm. On roots of Rubus
idaeus in England (Kent). Pemphigus
saliciradicis (Börner) Apterae are yellowish white, lightly coated
with greyish wax; BL 1.6-2.4 mm. On
roots of Salix spp. (fragilis, herbacea, polaris) in
Europe (records are from England, Finland, Sweden, Switzerland), Greenland
(as salicicola Hille Ris Lambers),
Transcaucasia (Georgia; but see below), north-east Russia (Wrangel Island;
Khruleva & Stekolshchikov 2019) and Japan. Aphids on Salix roots in California, USA, and on Baffin Island in arctic
Canada, are possibly also this species (but see below). Palaearctic populations identified as P. saliciradicis are apparently
normally anholocyclic on Salix
roots. A dense colony identified as this species was found on grapevine (Vitis vinifera) roots in Georgia
(Barjadze 2010), but a subsequent morphometric study (Barjadze & Asanidze
2013) revealed differences from the willow root-feeding aphid (and see note
below). Alatae that are intermediates between virginoparae and sexuparae
occur in Japan (Aoki 1975). Lange
(1965) reported Pemphigus
populations of the saliciradicis
group on Salix roots in northern
and southern California; the southern population occurred throughout the year
on Salix roots, whereas the the
northern population produced sexuparae in autumn, but no descriptions were
published of these aphids. Richards'
(1963) record of P. groenlandicus
(a grass-root feeder) from roots of Salix
reticulata on Baffin Island may be referable to this species. [Note:
There is a strong possibility that the aphids from willow roots in Europe and
Asia, described as P. saliciradicis by
Börner, are the introduced secondary host populations of the North American
poplar-galling aphid Thecabius
populimonilis, which is believed to have a partial migration to Salix roots; Aoki et al. (1996) noted a
close resemblance between first instar nymphs of the two species, and data
for apterous exules of T. populimonilis
from Salix roots and mosses in
Washington and Alaska, confirmed by DNA barcoding as this species (Pike et al. 2012), agree closely with P. saliciradicis (S. Bardjadze, pers.
comm. 2013). If this is the case then the grapevine-feeding aphid found in
Georgia will require a new name. The identity of a Pemphigus sp. collected from grapevine roots in Queensland,
Australia, shown by Barjadze & Asanidze (2013) to differ morphologically
from the grapevine root feeder in Georgia, will also need to be
re-investigated.] Pemphigus
sinobursarius Zhang Galls on Populus simonii in China are on undersides of leaves at base of
the lamina (Zhang & Zhang 1979c).
Alatae similar to P. bursarius,
but with a shorter R IV+V (0.065 mm).
Migration occurs to an unknown secondary host. Pemphigus
siphunculatus Hille Ris Lambers Galls on twigs of Populus ciliata in Pakistan in August contained apterous progeny
of the fundatrix, colour in life unknown (presumably wax-covered); BL 1.9-3.4
mm (Hille Ris Lambers 1973b). Presence
of apterae in the gall is normally indicative of a monoecious life cycle but,
according to S. Chakrabarti (pers. comm. cited by Aoki & Kurosu 1991d), P. siphunculatus is heteroecious. The secondary host is not recorded. Pemphigus
spyrothecae Passerini Gall is green, reddish or yellowish,
smooth, formed by thickening, flattening and spiral twisting of petiole of Populus nigra (fig. 131S). Gall formation was studied by Dunn
(1960). The fundatrix is pale green,
giving rise to second generation apterae within the gall. Monoecious holocyclic; alate sexuparae
emerge in August-November, to produce sexuales on the bark. Lampel (1960) provided a detailed
account. First instars with thick
forelegs function as soldiers and defend the gall against predators (Aoki
& Kurosu 1986, Foster 1990), also repairing the gall (Pike & Foster
2004) and removing wax-coated droplets of honeydew (N. Pike et al. 2002). Urban (2002) made
detailed studies of gall populations and the impact of natural enemies in the
Czech Republic. N. Pike (2007) showed that different morphs occupy different
positions within the gall, and N. Pike (2010) reported on the amino acid
profile of this species. Widely distributed in Europe, in North Africa
(Tunisia), Iran, Kazakhstan, Pakistan, eastern Siberia (Babichev &
Kirichenko 2020), and introduced into Canada (Quebec, Alleyne & Morrison 1974; British
Columbia). 2n=20. Pemphigus
tartareus Hottes & Frison (= Pemphigus
junctisensoriatus Maxson) Described originally
from alate sexuparae collected on roots of Bidens sp. in Illinois, USA. Galls on Populus deltoides are pale yellowish
green, composed mostly of swollen leaf petiole around broadened base of leaf
lamina, with an exit hole in the form of a long slit between the petiole and
the leaf (Maxson 1934). The fundatrix
is pale yellowish white, with brownish head and tibiae. Alatae (BL 1.4-1.8
mm) emerge in June-July and migrate to Bidens
sp(p). Smith (1985) established the synonymy with P. junctisensoriatus and made successful host transfers from Populus to Dichondra repens (Convolvulaceae) as well as to Bidens. In USA and eastern Canada. Pemphigus tibetapolygoni Zhang
Apterae are presumably with wax; BL c.2.5 mm. On roots of a Polygonum sp. in Tibet. Pemphigus
tibetensis Zhang
Galls are globular, pouch-like, formed on twigs of Populus cathayana in Tibet (Zhang
& Zhong 1979c). This appears to be
a member of the borealis/dorocola group. Pemphigus
trehernei Foster
The gall induced experimentally on Populus
nigra var. italica was green
with reddish tinge around exit hole, globular, formed at base of current
year's shoot. The fundatrix is dull
pale green. Alatae (BL 1.9-2.4 mm)
emerge in June-July and found colonies on roots of Aster tripolium in salt-marsh habitats (Foster 1975), and probably specific
to this halophytic host. Apterae on the secondary host are yellowish white
to greenish white with greyish white wax forming tufts on abdomen; BL 1.3-2.4
mm. Alate sexuparae (with secondary rhinaria distributed III 3-7, IV 1-3)
are produced in September-December. Anholocyclic
overwintering on Aster roots is
common, and the species has only been found naturally on its secondary host;
the gall generations were obtained by experimental transfer of sexuparae to P. nigra. In England, Ireland and
France. Pemphigus
turritus Zhang The
galls on Populus purdomii in China
(Gansu Prov.) are green, chilli- or horn-shaped, c. 15 mm × c. 3 mm thick.
Described from three alatae (BL c.2.4 mm) collected in July (G. Zhang &
Qiao 1997c). The life cycle is
unknown. Pemphigus
venosus Habib & Ghani Recorded as forming galls on twigs of Populus ciliata in Pakistan, but no
description is given, and therefore this is a nomen nudum. Pemphigus
(Pemphiginus) vesicarius Passerini Galls on Populus nigra originate from the mid-rib at the base of the upper
side of the leaf, developing in April-May into an irregular pale green
structure with numerous tubular outgrowths, of diameter up to 4 cm, that
encompasses the whole leaf (fig. 131T
and see influentialpoints.com/Gallery)
. The fundatrix is dark slate-grey to
blue-black. Alatae, BL 1.7-3.0 mm,
leave the gall in May-June through secondary holes at apices of outgrowths
(Roberti 1938) and migrate to found colonies on the stems and basal parts of Colutea arborescens. Apterae on Colutea (for long known as P. coluteae) secrete abundant wax and
have BL 2.7-3.0 mm. Coffin & Remaudière (2001)
determined the life cycle in southern France, described alate sexuparae and
sexuales, and compared the various morphs with those of P. populi. The male genitalia were described and
illustrated by Wieczorek et al.
(2012). In southern Europe, Algeria, south-west and
central Asia, east to Pakistan and India. Pemphigus
wuduensis Zhang The
gall on Populus purdomii in China
(Gansu Prov.) is flattened, rounded, and has a secondary exit. Described from
two alatae (BL c. 2.4 mm) collected in July (G. Zhang & Qiao 1997c). The life cycle is unknown. Pemphigus
zhongi (Zhang)
The gall on an unidentified Populus
sp. is “dumpling-shaped”, formed by folding of the two halves of a leaf
together, later becoming reddish, swollen and corrugated. Emigrant alatae are
blackish green in life; BL c.2.6 mm (G. Zhang et al. 1995, as Thecabius
(Parathecabius) zhongi). In Gansu province, China. The
host is identified in Zhang (1999, p. 69) as P. cathayana. Except for the high rhinarial numbers and different
gall, the alatae seem similar to those of P.
matsumurai.
Four
species of small brownish aphids with symmetrically swollen siphunculi, the
alatae having broadly dark-banded forewings with very characteristic
venation, the radius strongly curved and almost touching or fused with the
media for part of its length, making a closed or nearly closed cell (see aphids
of Karnataka website). Foottit & Maw (2019) reviewed geographical distribution, host
preferences and molecular diversity within the genus. Pentalonia caladii van der Goot
Apterae are reddish brown to almost black, with dark-tipped antennae;
BL 1.0-1.7 mm. On plants in the families Zingiberaceae and Araceae (e.g. Alpinia, Caladium, Colocasia, Eletteria, Hedychium, Zingiber),
or occasionally on plants in other families (Musa, Heliconius).
Described from Java, distributed widely through the south Pacific, Hong Kong, Australia, Central America, and in
hothouses in the northern hemisphere. This species has been long regarded as
a synonym of P. nigronervosa, or as
a “form” of that species, but molecular and multivariate studies by Foottit et al. (2010b) have shown that it
should be treated as a distinct species. Bhadra & Agarwala (2010)
compared fitness parameters of P.
caladii and P. nigronervosa in
reciprocal transfers between their respective host plants. Pentalonia gavarri Eastop
Apterae are dull black; BL 0.7-1.0 mm. Inside unfurling new grass
blades (Centrotheca, Oplismenus). P. gavarri was originally described from alatae trapped in
Philippines. Apterae described by Martin (1987) from West Malaysia, and
specimens presumed to have been collected from Brachiaria sp. (although the host-plant is not explicitly stated)
in India (Assam), described by A.K. Ghosh (1978) as Micromyzus brachiariae, clearly
correspond well with this species.
Alatae have also been trapped in New Britain and Australia (Carver
& Hales 1983). Pentalonia
kalimpongensis (A.N. Basu) Apterae are reddish brown, with
dark-tipped appendages; BL 1.5-1.9 mm. On Zingiberaceae (Curcuma, Elleteria,
Hedychium) in
India. In transfer experiments this
species did well on other Zingiberaceae (Canna
sp., Amomum sabulatum; original
description, as Micromyzus). Monoecious holocyclic; oviparae and alate males
were described from Elleteria by
Mondal et al. (1978). 2n=12. Pentalonia nigronervosa Coquerel
Banana Aphid Plate 19f Apterae are reddish‑brown to
almost black, with black-tipped antennae, and legs pale except for distal
parts of femora (see aphids
of Karnataka website); BL 1.2-1.9 mm. P. nigronervosa
feeds almost entirely on plants in the family Musaceae, with a few
records from Heliconiaceae and Zingiberaceae (Foottit & Maw 2019). It is
often found living under the old leaf bases, usually ant-attended. It is an
important vector of banana bunchy top virus (see also Blackman & Eastop
2000). P. nigronervosa is widespread through all tropical and
subtropical parts of the world, and in hothouses in Europe, Australia and
North America. Probably it is anholocyclic almost everywhere; in the BMNH
collection there is one alate male from Nepal (leg. K.C. Sharma). The sexual
morphs recorded from Curcuma in
India by Bhanotar & L.K. Ghosh (1969) were possibly P. kalimpongensis. The complete genome sequence has been
determined (Mathers et al. 2020).
2n=14.
Four
species in southern South America and one in western USA, possibly most
closely related to Carolinaia, with
5-segmented antennae in all morphs. Four of the five species are grass
feeders, and one (P. tenuis) has
strongly modified morphology which parallels that of unrelated grass-feeding
aphids. Nieti Nafría et al. (2002) reviewed the genus. Pentamyzus acaenae (Schouteden)
Apterae are pale green; BL 1.0-2.5 mm. On leaves and shoots of Acaena splendens in southern Argentina
(Tierra del Fuego) and Chile (Magallanes; Nieto Nafría et al. 2018). Monoecious holocyclic with an abbreviated life cycle, oviparae and
apterous males appearing in January (Nieto Nafría et al. 2002b). Pentamyzus falklandicus Hille Ris Lambers Colour of apterae in life is unknown; BL
2.4-3.0 mm. On Poa flabellata in
the Falkland Islands. Pentamyzus fueginus Nieto Nafría, Mier Durante & Ortego Apterae are reddish brown to greenish
brown; BL 1.8-2.6 mm. On outer-facing surfaces of leaves of Alopecurus magellanicus and Hordeum comosum growing in moist
situations. In southern Argentina (Tierra del Fuego). Monoecious holocyclic
with an abbreviated life cycle, oviparae and apterous males occurring in
January (Nieto Nafría et al.
2002b). Pentamyzus graminis Hille Ris Lambers
Plate 12b Apterae are uniformly green; BL 1.6-2.0 mm.
On basal parts of unidentified grasses, in somewhat moist and sunless
situations, in California, USA. Monoecious holocyclic, with numerous oviparae
and some apterous males occurring with apterous viviparae in March. Pentamyzus
tenuis Brown Colour of apterae in life is unknown,
probably wax-covered; BL 2.3-2.9 mm. On Poa
alopecurus on Falkland Islands, and subsequently found on Alopecurus magellanicus on Tierra del
Fuego (Nieto Nafría et al. 2002b).
About 42
species of medium-sized to large, generally long-haired aphids, almost all
living on Acer except for a few
species on Aesculus and one on Koelreuteria. The antennal PT is usually long and the
siphunculi are stump-shaped, usually with polygonal reticulation, but often
longer than in Chaitophorus and
with a pronounced flange. The cauda is
either rounded or tongue-shaped with a slight constriction. Colonies are usually ant-attended. There is no host alternation, and the
association with the host plant is an intimate one, involving a complex
seasonal polymorphism to cope with physiological changes in the host, the
most unusual feature of which is the frequent occurrence of specialised
larval aestivating morphs (fig. 5). Periphyllus
is predominantly a palaearctic genus, (15 European, 11 north
Indian/Himalayan, 13 east Asian), with just 3 species native to North
America. Essig & Abernathy (1952) wrote
a monograph of the genus. There are
keys and/or accounts of Periphyllus
species of Europe (Hille Ris Lambers 1967a), Britain (Stroyan 1977),
Fennoscandia and Denmark (Heie 1982), Iberian peninsula (Nieto Nafría &
Mier Durante 1998), Poland (Wieczorek & Junkiert 2012), European Russia
(Mamontova 1955), India (Chakrabarti et al. 1987), Pakistan (Remaudière
2002), China (Tao 1964), east Siberia (Pashchenko 1987a, 1988a, b), east Asia
(Park & Park 1995), Japan (Sorin 1990) and Canada (Richards 1972a). Wieczorek
& Junkiert (2011) and Trela et al.
(2020) provided keys to oviparae and males of species found respectively in
Europe and the Nearctic, and Lubiarz & Mackoś-Iwaszko (2015)
provided morphometric data and keys to fundatrices of Periphyllus species living in Poland on Acer platanoides. Accounts of British species with excellent
colour photographs are available online at http://influentialpoints.com/Gallery/Periphyllus_aphids.htm.
T. Liu et
al. ( 2022) studied macroevolution of Periphyllus
and other Chaitophorinae in relation to changes in host use. Recent
molecular work indicates that Periphyllus
is sister group to a clade containing Chaitophorus
and Siphini (Wieczorek & Kajtoch 2011, Wieczorek et al. 2017b), but more recent results (T. Liu et al. 2022) suggest that the
transition of Siphini to grasses may have occurred much earlier. Aphidius aerolatus, A. setiger, Praon negundinis and Trioxys
falcatus are specialised parasitoids. Periphyllus
acericola (Walker)
Apterae are pale green or yellowish green, sometimes with dorsal
brownish markings; BL 2.4-3.5 mm.
Alatae have broad dark dorsal abdominal cross-bars scarcely separated
between segments, and paler marginal sclerites (see
influentialpoints.com/Gallery); BL 3.0-3.5 mm. On undersides of leaves, petioles and young shoots of Acer pseudoplatanus throughout most of Europe, and usually
regarded as specific to this host, but Lubiarz & Mackoś-Iwaszko
(2015) recorded fundatrices on A.
platanoides in Poland. Aestivating first instar nymphs are yellowish
white with long, pointed dorsal and marginal hairs; they aggregate in dense groups
appearing like whitish spots on undersides of leaves. Oviparae and alate males occur in
October. 2n=18*. Periphyllus
acerihabitans Zhang
Apterae are pale green; BL 1.9-2.5 mm.
The antennal terminal process is unusually short, less than twice the
length of the base of ANT VI. Alatae are without dark dorsal abdominal
sclerotisation. On undersides of leaves, petioles and young shoots of Acer buergerianum, usually
ant-attended. Aestivating nymphs have foliate marginal hairs. Oviparae occur
in October-November, and were described by Wieczorek et al. (2016a), who also provided redescriptions of the apterous
and alate viviparae; males are unknown. In China and Japan. Niu et al. (2016) compared the
mitochondrial genome of P.
acerihabitans with that of Chaetosiphella
stipae. Periphyllus
aceriphaga Chakrabarti & Mandal Appearance in life is unknown; BL 2.4-2.7
mm. On Acer spp. (caesium, caudatum) in Uttar Pradesh, India
(Chakrabarti et al. 1987), and
Pakistan. Aestivating nymphs have
foliate marginal hairs. Remaudière
(2002) recorded oviparae and alate males collected from Acer sp. at Murree, Pakistan in November. Periphyllus
aceris (Linnaeus)
Apterae are yellow, often with green flecks dorsally (see
influentialpoints.com/Gallery); BL 1.5-3.7 mm. Alatae have dorsal cross-bands more widely
separated than in P. acericola,
with equally dark marginal sclerites and pterostigma; BL 3.2-4.5 mm. On undersides of leaves, petioles and
growing shoots of Acer spp.,
especially A. platanoides, which is
clearly its normal host. There are
also records from Aesculus, but no
authentic specimens have been seen and these should possibly be referred to P. testudinaceus. Throughout most of Europe, and also
recorded from Iran (Abaii 1984). Previously stated to have been introduced to
North America, but Foottit et al.
(2006) point out that all North American specimens in USNM are
interceptions. Aestivating nymphs have
long pointed dorsal and marginal hairs and aggregate like those of P. acericola. Monoecious holocyclic; oviparae and alate
males occur in October-November. The fundatrix was described by Lubiarz
& Mackoś-Iwaszko (2015). This species has been confused in the
literature with acericola, and also with other species; e.g. Börner
(1940) had P. lyropictus, and
Wahlgren (1935) had P. testudinaceus. 2n=16 (Blackman & Eastop 1994). Periphyllus
aesculi Hille Ris Lambers Apterae are rather elongate oval, brownish
in life with black dorsal spots (A.K. Ghosh 1980); BL 2.5-3.5 mm. Siphunculi are wholly dark. Alatae have short dorsal abdominal
cross-bars and dark marginal sclerites.
On undersides of young leaves of Aesculus
indicus in India (Kashmir, Himachal Pradesh, Uttar Pradesh) and Pakistan
(Murree; Remaudière 2002). Chakrabarti
et al. (1972a) described the apterous vivipara and redescribed the alate
vivipara, L.K. Ghosh (1986) described the ovipara, and Agarwala &
Raychaudhuri (1982) described the alate male.
Aestivating morphs are not recorded.
Sexuales were collected in October-December. Periphyllus
allogenes Szelegiewicz
Apterae are elongate oval, brown with dark brown antennae and legs; BL
1.0-1.2 mm. Alatae are unknown. On leaf petioles of Acer triflorum in Korea.
Described from autumnal (late August) apterae, which may have
developed from aestivating nymphs and be very different from spring forms,
which are possibly much larger (Szelegiewicz 1981b). However apterae of P. triflorumi, also described from A. triflorum in Korea
without reference to P. allogenes and
presumably a synonym, are equally small
and were collected in July (Lee & Seo 1992). The biology is unknown. Periphyllus
americanus Baker
Apterae in life are "pale yellowish green with dark median stripe
on abdomen forming a cross in region of cornicles" (Hottes, quoted by
Palmer, 1952); BL 2.0-2.5 mm. The
alata is dark greenish with black dorsal abdominal cross-bars and marginal
sclerites. On leaves of Acer spp., especially sugar maples,
widely distributed in North America.
Aestivating nymphs have foliate marginal hairs (fig. 19A). Essig & Abernathy (1952) gave an
account of the summer generations. Fundatrices and oviparae have apparently
not been described. Trela et al.
(2020) described an alate male (leg. G.F. Knowlton), collected in Utah in
October. Periphyllus
bengalensis Ghosh & Raychaudhuri Apterae are yellow, with dark siphunculi,
pear-shaped; BL 1.8-2.2 mm. Alatae
have dark dorsal abdominal cross-bands.
Forming a colony on undersides of older leaves of an undentified Acer sp. in West Bengal, India (A.K.
Ghosh 1980). Aestivating and sexual
morphs are unknown. Periphyllus
brevisetosus Sorin Appearance in life is unknown; only late
summer apterae (BL 1.2-1.3 mm) are described, collected in Japan on Acer pictum var. typicum (= mono) (Sorin
1990b). Closely related to P. kuwanaii. Periphyllus
brevispinosus Gillette & Palmer Apterae are brown with dark and pale mottling,
and pale siphunculi; BL 2.4-2.9 mm.
Alatae are brown with black dorsal abdomnal cross-bands and marginal
sclerites. In undersides of leaves of Acer glabrum in western North
America. Aestivating nymphs have
foliate marginal hairs with serrate margins (Essig & Abernathy
1952). Monoecious holocyclic; ovipara
and alate males collected in mid September were described by Trela et al. (2020). Periphyllus
bulgaricus Tashev
Apterae are shining black, with pale legs and antennae; BL 2.0-2.8
mm. Apterae and alatae both have
variably-developed black dorsal abdominal cross-bars (see Nieto Nafría &
Mier Durante 1998). On shoot tips and
undersides of leaves, usually at base of mid-rib. Described from Acer hyrcanum in Bulgaria (Tashev 1964c); also collected from A, hyrcanum in Turkey, from A. opalus and A. ibericum in Iran, from A.
hermaneum in Lebanon (BMNH collection, leg. D. Hille Ris Lambers), and
from A. granatense and A. monspessulanum in Spain. Nieto Nafria & Gonzales Funes (1990)
described fundatrices, oviparae and alate males, and discussed the very
difficult separation of this species from P.
rhenanus. Aestivating nymphs have
not been recorded. Periphyllus
caesium Chakrabarti & Saha Appearance in life is not recorded, probably
rather pale with dark siphunculi; BL of aptera 2.3-2.4 mm. Alatae have dark dorsal abdominal
cross-bands. On Acer caesium in Uttar Pradesh, India (Chakrabarti et al. 1987). An alate male ascribed to this species was
collected in October. Longer haired
than P. aceriphaga, and with a
shorter HT II, but otherwise apparently very similar. Periphyllus
californiensis (Shinji) Apterae are dark olive-green to brown, with
dark brown head and prothorax, and irregular dark dorsal segmental spots or
cross-bands (see
influentialpoints.com/Gallery); BL 2.3-3.5 mm. Alatae have dorsal abdominal cross-bands
which with a hand lens appear darker than the pterostigma of the forewing
(cf. aceris). On Acer
spp. in east Asia (China, Japan, east Siberia, Korea, Taiwan, Thailand, West
Bengal), living on undersides of leaves along veins; Pashchenko (1987) gave A. pseudosieboldianum as the principal
food plant. Also recorded from Aesculus turbinata in Japan (as Chaitophorinella acerifoliae;
Takahashi 1919a). Introduced to
Europe, North America, Australia and New Zealand, where it is most commonly
found on planted Asian ornamental maples, but sometimes also colonises native
species; there is one record from Aesculus
californica in California.
Aestivating nymphs have foliate marginal hairs. Essig & Abernathy (1952) fully
described the life cycle in California, USA, where oviparae and alate males occur
in October-November. Population
dynamics of P. californiensis in
Japan has been studied in relation to host phenology (Hashimoto & Furuta
1988, Wang 2006), mortality of aestivating
nymphs (Furuta 1985), host preferences (Furuta 1986) and natural enemies
(Furuta & Mao 1986, Kan & Sasakawa 1986). 2n=18 (Blackman 1986). Periphyllus
changlai Remaudière
Apterae are pale-coloured, long-bodied; BL 3.2-3.8 mm. Alatae are
without dorsal abdominal sclerotisation. On Acer caesium in Pakistan (Remaudière 2002). An ovipara and alate
males were collected at the same time as the parthenogenetic morphs in
October. Periphyllus
coracinus (Koch)
Apterae are variable in colour, yellow, light or dark green, or brown,
or mottled with green or brown. On
leaf petioles and twigs of Acer platanoidis
in western, central and eastern Europe (Netherlands, Sweden, Finland,
Germany, Austria, Poland, Hungary, Moldova, Ukraine, Turkey), and also
recorded from Kazakhstan (Kadyrbekov 2017a).
Oviparae and alate males occur in October. The male genitalia were described and illustrated by Wieczorek et al. (2011). The fundatrix was
described by Lubiarz & Mackoś-Iwaszko (2015).
Apparently no specialised aestivating nymphs are produced. Periphyllus
formosanus Takahashi
Apterae are reddish brown, with thorax darker; BL c. 2.5 mm. Alatae have blackish dorsal abdominal
cross-bands. On undersides of leaves
of unidentified Acer sp(p). in
Taiwan and China. Aestivating nymphs
have foliate marginal hairs (Takahashi 1933a). Probable fundatrices were observed by
Takahashi in February, and Essig & Abernathy (1952) described the
oviparae and alate males, collected in November. Periphyllus
garwhalensis Chakrabarti & Mandal Apterae are undescribed. Alatae (BL 3.0-3.6 mm) have dark siphunculi
but no dorsal abdominal cross-bands.
On Acer caudatum in Uttar
Pradesh. India. Aestivating nymphs
have foliate marginal hairs with serrate margins (Chakrabarti et al. 1987). Periphyllus
ginnalae Paik ex Szelegiewicz (= P.
loricatus Pashchenko) Apterae and
alatae are shining dark brown to black; BL 1.6-2.6 mm. Alatae have secondary rhinaria distributed
ANT III 10-19, IV 0-4, V 0-2. On Acer
ginnala, living in early spring on leaf petioles at tips of young branches,
and later forming dense colonies on the undersides of the leaves. Aestivating nymphs are green to brown with
foliate marginal hairs, living singly on both sides of leaves; but most of
the population apparently does not aestivate (Pashchenko 1988b, as loricatus). Apterous males and oviparae occur in
September (Pashchenko 1987a). In Korea
and east Siberia, and possibly also in Japan, as P. takahashii Sorin (1990b), described from A. ginnala in Japan, seem likely to be small specimens of this
species. [Park & Park (1995) questioned the synonymy of ginnalae, loricatus and takahashii proposed
in Blackman & Eastop (1994). Although they appear to be correct in
claiming validity for the name ginnalae,
which had been listed as a nomen nudum
(Eastop & Hille Ris Lambers 1976, Remaudière & Remaudière 1997),
specimens in the BMNH collection from A.
ginnala in both North and South Korea determined as ginnalae by Paik and by Szelegiewicz agree well with the
description of P. loricatus (Pashchenko
1987), in particular having 7 hairs on first tarsal segments and R IV+V
0.7-0.8 × HT II. We therefore remain of the opinion that these are all likely
to be one species.] Periphyllus
himalayensis Chakrabarti Apterae are undescribed. Alatae have dark dorsal abdominal cross-bands,
colour in life unknown, BL 3.1-3.5 mm (Chakrabarti 1977). On unidentified Acer sp(p). in northern India (Himachal Pradesh, West
Bengal). Biology is unknown. Perhaps these are large specimens of P. bengalensis. Periphyllus
hirticornis (Walker) (fig. 23B, fig. 25A)
Apterae
are light green, with red eyes, without any dark dorsal markings (see
influentialpoints.com/Gallery); BL 2.0-3.0 mm. The abdomen of
the alata has variably developed spinal sclerites, but not cross-bands. On undersides
of leaves, leaf petioles and fruits of Acer
campestre, often with attendant ants.
Aestivating nymphs have foliate marginal hairs (see Stroyan 1977).
Widely distributed in Europe. Oviparae and alate males are produced in
October-November. 2n=18*. Periphyllus
hokkaidensis Sorin Apterae are dark reddish to blackish
brown; BL c. 2 mm. On Acer miyabei and A. mono var. subtrifidum
in Hokkaido, Japan (Sorin 1990b). Other morphs and biology are unknown. It seems to be closely related to P. kuwanaii. Periphyllus
karatavicus Kadyrbekov Apterae pale green; BL c.1.5-1.6 mm. On
upper and undersides of young leaves of Acer
semenovii in Kazakhstan (Kadyrbekov 1999b). Periphyllus
koelreuteriae (Takahashi) Apterae are variably pigmented, yellow or
black, with wholly black tibiae, and often with paired dark dorsal spots on
abdominal tergites; BL 1.6-2.5 mm.
Alatae have dark dorsal abdominal cross-bars, and 25-38 secondary
rhinaria on ANT III (cf. alatae of californiensis,
with 7-25 secondary rhinaria on III).
On Koelreuteria paniculata
in China, Korea and Japan. Aestivating
nymphs have foliate marginal hairs (Takahashi 1933a, Essig & Abernathy
1952). The life cycle in Taiwan was
studied by Y.C. Liu et al.
(1999)and the sexual morphs were described by Junkiert & Wieczorek
(2019). The recorded occurrence on Aesculus
turbinata is apparently casual, being based on a single alata (Higuchi
1972). DNA studies in southern China
(Q. Li et al. 2020, 2021) indicate
that there may be geographic races or cryptic species. Sorin's (1990b)
apterae from A turbinata keyed as P. koelreuteriae do not seem to be
this species. 2n=18 (Blackman &
Eastop 1994). Periphyllus
kuwanaii (Takahashi)
Apterae and alatae are shining black; BL 1.4-2.4 mm. Principally on Acer mono; Pashchenko (1988 a,b) recorded it from several other Acer spp., but such visits may be
casual. In China, east Siberia, Japan
and Korea. Aestivating nymphs have
foliate marginal hairs (Takahashi 1933a); but part of the population does not
aestivate (Pashchenko 1988b). Sexual
morphs are apparently undescribed. Periphyllus diacerivorus Zhang (in
Zhang & Zhong 1982c) seems to be a synonym. 2n=18 (Blackman & Eastop 1994). Periphyllus
lyropictus (Kessler)
(fig. 23D)
Apterae
are yellowish with variably developed brown or green dorsal markings, usually
comprising a broad spinal stripe on head and thorax and a large V-shaped mark
on the dorsal abdomen (see influentialpoints.com/Gallery); BL 1.9-3.0 mm. Alatae have dark marginal sclerites, but
other dorsal sclerotisation is limited to the spinal area, not forming
cross-bands. On undersides of leaves
of Acer platanoides, often forming
large colonies producing much honeydew and visited by ants and other
insects. This species does not produce
aestivating nymphs. In Europe, Iran
(Rezwani et al. 1994), and
introduced on Norway maple to North America, where it is also
widespread. A. platanoides is clearly the favoured host, and records from
other Acer are mostly due to
confusion with P. aceris and P. testudinaceus, particularly in the
North American literature (see Essig & Abernathy 1952). Oviparae and alate males are produced in
October-November. Periphyllus
mamontovae Narzikulov
Apterae are green; BL 2.1-2.9 mm.
On leaves of Acer turkestanicum
and (more rarely) A. regelii in
Tajikistan (Narzikulov 1957).
Aestivating nymphs are apparently not produced. Life cycle and sexuales have not been
described. Periphyllus
mandshuricus Pashchenko Apterae are apparently not produced in
spring; reddish brown to dark brown fundatrices on shoot tips produce only
dark brown alatae (BL 2.5-3.4 mm), which live on undersides of leaves and
produce aestivating nymphs with foliate marginal hairs. On Acer
mandshuricum in maritime region of east Siberia. Oviparae and alate males ocur in October
(Pashchenko 1988b). Periphyllus
minutus Shaposhnikov
Apterae are shining brown or greenish yellow, with dark brown dorsal
markings; BL 1.6-2.5 mm. Alatae have
dark dorsal abdominal cross-bands. On Acer tataricum, colonising young
shoots, leaf petioles, undersides of leaves, fruit stalks and fruits. This species is also recorded from A. ginnala (Ripka et al. 1998) and A.
campestre (Vereshchagin et al.
1985). Aestivating nymphs have long pointed dorsal and marginal hairs. In eastern Europe, south-west and central Asia
(Bulgaria, Hungary, Moldovia, Ukraine, Iran, Kazakhstan). Sexuales have not been described. Described as a subspecies of aceris by Shaposhnikov (1952); Mamontova (1955) provided a fuller
description and gave it full species status. Periphyllus
negundinis (Thomas)
Apterae are pale yellow-green to apple green with dark dorsal markings
(see influentialpoints.com/Gallery);
BL 2.0-2.5 mm. Alatae have variable
dark green dorsal abdominal markings, but no distinct cross-bands. On Acer
negundo, feeding on young growth in or near fruit clusters in spring,
then on undersides of leaves. Widely
distributed in North America, including Mexico. Aestivating nymphs have
foliate marginal hairs. Apterous males and oviparae occur in late September
to November. Essig & Abernathy
(1952) provided detailed descriptions of the life cycle, and Trela et al. (2020) redescribed the sexual
morphs. 2n=20. Periphyllus
nevskyii Mamontova
Apterae are shining grey-black, with a central black sclerite
extending across ABD TERG 4-6; BL 1.3-2.0 mm (Nevsky 1929a; as Chaitophorinella negundinis; Mamontova
1955). On Acer regelii in central Asia (Tajikistan, Kazakhstan,
Uzbekistan). Also recorded from A. semenovii (Kadyrbekov 2002), A. turkestanica and A. pseudoplatanus (Gabrid 1989). The life cycle has not been studied. Periphyllus
obscurus Mamontova
(fig. 23C, fig.
24B)
Apterae
are blackish green (see influentialpoints.com/Gallery); BL 1.8-2.6 mm. In ant-attended colonies on young shoots,
leaf petioles and undersides of leaves of Acer
campestre in Europe (Bulgaria, Czechoslovakia, England, Netherlands,
Germany, Hungary, Poland, Turkey, Ukraine) and Iran (Abaii 1984). Aestivating nymphs are apparently not
produced. Sexual morphs have not been
described. Periphyllus
pakistanicus Remaudière Apterae pale-coloured with dark apices to
antennal segments, dark siphunculi, and dark spots at bases of dorsal hairs;
BL 2.1-2.5 mm. Alatae are without dorsal abdominal cross-bands. In spring
populations on Acer oblongus in
Pakistan (Remaudière 2002). Periphyllus
pallidus Chakrabarti & Saha Appearance in life not recorded; BL of
aptera 2.4-2.7 mm. On an unidentified Acer sp. in Uttar Pradesh, India
(Chakrabarti et al. 1987). Aestivating
nymphs have foliate marginal hairs.
Other morphs and life cycle are unknown. Periphyllus
rhenanus (Börner)
Apterae are shining, dark; BL 2.5-3.0 mm. On undersides of leaves, extending onto
petioles and twigs. Alatae have broad
black dorsal abdominal cross-bars and equally dark marginal sclerites. On Acer
monspessulanum and A. obtusatum
in southern and central Europe (Bulgaria, France, Germany, Greece, Italy,
Switzerland). Aestivating nymphs have
very long marginal hairs and form dense clusters (Barbagallo & Stroyan
1982). Apterous males and oviparae
occur in October-November (Hille Ris Lambers 1947d; as P. lichtensteini). Nieto
Nafría & Gonzales Funes (1990) discussed differences from the
closely-related P. bulgaricus. Periphyllus
singeri (Börner)
Apterae are reddish brown, in spring bearing dark dorsal abdominal cross-bands
like the alatae; BL 2.4-3.5 mm. On
young shoots or branches of young trees of Acer pseudoplatanus, or on old wounds on the bark of large tree
trunks, sometimes in large colonies, attended by ants (Lasius fuliginosus). In
Europe (Austria, Denmark, Germany, Lithuania, Netherlands, Ukraine), but
apparently rare or sporadic in occurrence.
Aestivating nymphs are apparently not produced. Oviparae and alate males occur in October-November,
and were fully described and illustrated by Wieczorek & Junkiert (2011). The male genitalia were described and
illustrated by Wieczorek et al.
(2011). Periphyllus
steveni Mamontova
Apterae are elongate oval, shining brown or dark green; BL 1.8-2.4
mm. On young growth and leaves of Acer stevenii in the Crimea (Mamontova-Solukha
1962). One of the group of species
with long black dorsal hairs (Nieto Nafria & Gonzales Funes 1990). Aestivating nymphs are yellow, with long
pointed hairs. Life cycle and sexual
morphs have not been described. Periphyllus
tegmentosus Pashchenko The fundatrix is green with brown markings;
her progeny are all alate, having shining greenish yellow abdomen with dark
dorsal adbominal cross-bars and margunal sclerites. BL of alata 2.2-2.9 mm. On Acer
tegmentosum in maritime east Siberia.
Aestivating nymphs yellow-green with foliate marginal hairs. Oviparae and alate males occur in October
(Pashchenko 1988b). Periphyllus
testudinaceus (Ferni) Plate 12c, d (fig. 23A, fig. 24C)
Apterae
are dirty dark green to dark brown or blackish (see
influentialpoints.com/Gallery); BL 2.0-3.7 mm. Alatae have dark dorsal abdominal
cross-bands, darker than pterostigma, and equally pigmented marginal
sclerites. On young growth, leaves and
leaf petioles of numerous Acer spp.
(but especially common on A. campestre),
and sometimes on Aesculus spp. Throughout Europe, in Iran (Hodjat 1993),
in Korea (needs confirmation), and introduced to Tasmania, New Zealand and
North America - where it is widely distributed. Aestivating nymphs with foliate marginal
hairs are the only morph occurring from June to September (fig. 5). Oviparae and alate males occur in
October-November. Essig &
Abernathy (1952) provided a detailed account of the life cycle and
polymorphism. Warrington &
Whittaker (1985) showed that ant attendance caused a large increase in sap
ingestion by this species. 2n=18. Periphyllus
tokyoensis Sorin
Appearance in life is not recorded.
BL of aptera only c. 1.1 mm. On
an unidentified Acer sp. in Japan
(Sorin 1990b). Aestivating nymphs have
foliate marginal hairs. Other morphs
and life cycle are unknown. Possibly
these are dwarf specimens of a previously known species. Periphyllus
unmoonsanensis Park & Park Apterae are “generally brown”; BL 2.6-2.8
mm. Alatae are yellowish green. On Acer
palmatum in Korea (Park & Park 1996). Periphyllus
vandenboschi Hille Ris Lambers Colour of aptera in life was not recorded
but probably pale green, with contrastingly black siphunculi; BL 1.4-2.0
mm. Alata has pale brown abdomen with
rather small and indistinct spinopleural sclerites on ABD TERG 1-6 and
cross-bands on ABD TERG 7 and 8 (Remaudière 2002). Collected on leaf petioles of an
unidentified Acer in Pakistan
(Hille Ris Lambers 1966c), subsequently found on A. caesium and ornamental A.
oblongum (Naumann-Etienne & Remaudière 1995), and also recorded from
Himachal Pradesh, India (see L.K. Ghosh 1986). Aestivating nymphs and other morphs,
including sexuales, have not been recorded. Chakrabarti et al. (1972a) had P.
villosii. Periphyllus
venetianus Hille Ris Lambers
(fig. 24A, fig. 25B)
Apterae
are shining blackish green with pale antennae and legs; BL 1.1-2.0 mm. Alatae are undescribed. On undersides of leaves and leaf petioles
of Acer campestre, attended by
ants. Aestivating nymphs have foliate
marginal hairs. In southern Europe
(Austria, Czechoslovakia, France, Italy, Yugoslavia). Barbagallo & Stroyan (1982) recorded
immature oviparae in Sicily in September. Periphyllus
villosii Chakrabarti
Aptera are elongate oval, large, probably pale; BL c. 3.5 mm. The abdomen of the alata is without dark
dorsal cross-bands. On Acer spp. (caesium, caudatum, villosum) in Uttar Pradesh,
India. Aestivating nymphs have foliate
marginal hairs (Chakrabarti et al.
1987). Oviparae and alate males occur
(on A. villosum) in October
(Chakrabarti 1977). Remaudière (2002) reported this species from Acer caesium in Pakistan, but noted
differences from the original description that might indicate the existence
of two distinct species. Periphyllus
viridis (Matsumura)
Apterae (?fundatrices) are green; BL 2.9-3.9 mm. Alatae (in spring) are without dorsal
abdominal cross-bands; according to Pashchenko (1987), all progeny of the fundatrix
are alate. On young growth, leaves and
leaf petioles of Acer mono in
Japan, Korea and maritime east Siberia.
Aestivating nymphs have foliate marginal hairs. Oviparae and alate males occur in October;
Pashchenko (1987) gave a full account of the life cycle. Sorin (1990b) described some large alate
specimens in Japan as a subspecies, P.
viridis osugiensis.
Perhaps
only one species, unique in morphology and in that the only alatae are
sexuales. Parthenogenetic forms are all
apterous, with fused head and prothorax and 3-faceted eyes. Phloeomyzus
passerinii Signoret Plate 2e, f Apterae are green, covered with dirty
white wax wool; BL 1.2-2.2 mm. On bark
and in crevices on trunks of Populus
spp., heaviest infestations occurring on 6 to 8-year-old trees. There is no host alternation. Oviparae and males are produced in
September-October in the northern hemisphere, the oviparae each laying two
eggs (Theobald 1929). The male genitalia were described and
illustrated by Wieczorek et al.
(2011). Anholocyclic overwintering by apterous
viviparae is also common, and may predominate; no morph recognisable as a
fundatrix has been described. In
Europe, north Africa (Egypt, Morocco), south-west, central and east Asia
(incl. Japan; BMNH collection, leg. S.Aoki); USA (only record is from Maine)
and South America (Argentina, Chile, Colombia). Arzone & Vidano (1984) studied the life
cycle in Italy. The European P. nigra-feeding form was described as
a separate species, Ph. redelei
Hille Ris Lambers 1931 (see Doon & Hille Ris Lambers 1962), as also were
large specimens found on P. tremuloides
in Maine, USA (as Ph. dearborni;
Smith 1974b). Zhang (in Zhang &
Zhong 1982) described a subspecies on P.
lasiocarpa in China, P. passerinii
zhangwuensis, which was redescribed by Qiao et al. (2018). It seems
likely, however, that a single, somewhat variable species is involved. 2n=10.
A
small genus of Myzus-like species
with finger-like processes on antennal tubercles and a secondary host
association with Cannabaceae (Cannabis,
Humulus). The host-alternating species have Prunus as primary hosts. See also Blackman & Eastop 2000, p.
321-2. Phorodon (Diphorodon) cannabis Passerini Cannabis Aphid Plate 19h Apterae are shiny yellowish green with three
green longitudinal stripes, or pale to mid-green or pale pink without stripes
(see
influentialpoints.com/Gallery); BL 1.5-2.7 mm. Alatae
have a dark dorsal abdominal patch, and secondary rhinaria distributed III
15-30, IV 2-13, V 0-2. On undersides of leaves of and on flower stems of Cannabis spp. In central, eastern and
southern Europe, across Asia to Japan, in North Africa, and now recognised to
occur widely in North America (Cranshaw et
al. 2018). Monoecious holocyclic, with oviparae and alate males in
September-November (Müller & Karl 1976). 2n=12. Phorodon
humuli (Schrank) Damson-Hop Aphid Plate 19i Apterae on Prunus in spring are pale green to
yellowish green, rather shiny, with three darker green dorsal longitudinal
stripes (see
influentialpoints.com/Gallery); BL 2.0-2.6 mm. Alatae have a black dorsal abdominal patch and secondary rhinaria
distributed III 23-35, IV 1-11, V 0. Spring colonies occur on Prunus insititia and related species (domestica, mahaleb, spinosa), causing only slight leaf
curl. It is a vector of plum pox
virus. Migration occurs in May-June to
hops (Humulus lupulus), on which
its importance as a pest has increased with development of insecticide
resistance (Hrdy et al. 1986).
Apterae feed on undersides
of leaves and on flowers and fruits of hops and are pale green with darker
green medial and lateral longitudinal stripes; BL 1.0-2.1 mm. Return
migration to Prunus (studied by
Eppler 1988) is in September-October.
In Europe, north Africa (Ethiopia), south-west Asia (Iran, Turkey),
Central Asia; and introduced to North America, where it is widespread, and
New Zealand (BMNH collection, leg. P.L. Robertson). Morphometric variation was studied by
Hampson & Madge (1986 a,b), and allozyme variation by Eggers Schumacher
& Sander (1988). The photoperiodic determination of sexual morphs was
studied by Campbell & Tregidga (2005). See also Blackman & Eastop
2000, p.321). 2n=12. Phorodon humulifoliae Tseng & Tao
Apterae are pale yellowish green or green with dark head, ANT, legs,
and cauda; BL c.1.8 mm. Siphunculi are
strongly curved outward, and brownish towards their apices. On leaves of Humulus japonicus in China (Chekiang, Szechuan). Presumably there is host
alternation to Prunus spp. as
primary hosts, as there are records from north-west China of this species
occurring on P. armeniaca and P. salicina (G. Zhang 1999). The
record of P. humulifoliae from Ulmus sp. (Tao 1963) is probably in
error. 2n=12 (Chen & Zhang, 1985b). Phorodon
japonensis Takahashi Apterae
and alatae resemble those of P. humuli
(Takahashi 1965c); BL 1.4-1.8 mm. On Prunus spp. (mume, salicina) in
Japan, Korea, east Siberia and Taiwan.
Migration occurs to Humulus
spp. (japonicus, scandens). It was described as a subspecies of humuli, to which it is closely
related. 2n=12. Phorodon viburni Matsumura
Apterae are undescribed, alatae are brownish, BL c.1.5 mm. On
undersides of leaves of a Viburnum sp.
in Japan. This species has apparently not been found since its original
description in 1918.
A
genus with two or three species on Fagus
with well-developed dorsal wax glands, a very short processus terminalis and
pore-like siphunculi. Takahashi (1960)
distinguished it from related genera with a key, and Stroyan (1977) provided
a generic diagnosis. Quednau (2010) reviewed the genus and provided
illustrations of all available morphs. Trioxys
phyllaphidis is a specialised parasitoid. Phyllaphis
fagi
(Linnaeus) Plate
8 e,f Apterae are elongate
oval, pale yellowish green, covered with wax wool (see
influentialpoints.com/Gallery); BL usually 2.0-3.2 mm (but
summer dwarfs down to 1.1 mm). Alatae
have abdomen wax-covered, with variably-developed dark dorsal
cross-bars. On undersides of mostly
young leaves of Fagus spp., causing
them to curl downwards on both sides of the mid-rib, and often to wither and
die prematurely. Throughout Europe,
east to Turkey and Caucasus, more recently reported from China (Qiao &
Zhang 2004a) and Korea (Quednau & Lee 2001), and introduced to Australia,
New Zealand and North America. Records
from Japan possibly all apply to P.
fagifoliae. Monoecious holocyclic,
with dwarf apterae living through summer months to produce apterous oviparae
and alate males in October-November.
Oviparae have variably developed dark dorsal cross-bars (cf. fagifoliae and grandifoliae). Wieczorek
& Świątek (2008) studied morphology and
ultrastructure of the male reproductive system. Chandler (1968) studied its
association with a syrphid predator.
2n=16. Phyllaphis
fagifoliae Takahashi
Apterae are pale, presumably with wax; BL 1.2-2.5 mm. Alatae have much less pigmentation than P. fagi (Takahashi 1937). On Fagus
spp. (crenata, japonica) in Japan. A
record from Turkey (Görür et al.
2011a) is
doubtful and requires confirmation. Sexuales occur in October (Higuchi 1972;
as P. fagi). Oviparae are pale without dorsal abdominal
cross-bars (Quednau 2010). 2n=26 (after Shinji; see Blackman 1986). [This
species was synonymised with P. fagi by
Higuchi, but the difference in chromosome number coupled with morphological
differences indicate that it is a distinct species.] Phyllaphis
grandifoliae Richards Apterae are pale greenish, pale yellow or
somewhat cream-coloured (Richards 1973), usually exuding filamentous wax (see
influentialpoints.com/Gallery);
BL c. 1.8 mm. On Fagus grandifolia in western North America. Alatae have not been
recorded. Oviparae have pale siphunculi and lack dorsal abdominal cross-bars
(Smith & Parron 1978, Quednau 2010).
Phyllaphis
nigra Ashmead
Apterae are shiny black; BL c. 1.3 mm.
On tender shoots of Quercus
phellos var. laurifoliae (= Q. laurifolia) in Florida, USA
(Ashmead 1881). It has not been
recorded since its original description, and almost certainly does not belong
in this genus; it is perhaps a Lachnochaitophorus. The life cycle is not known.
A genus
for one east Asian narrow-bodied species living on bamboos. Quednau (2003)
illustrated the available morphs. Phyllaphoides
bambusicola Takahashi Alatae are white with pale yellow
mesothorax, secreting white cottony wax (see Martin & Lau 2011 for photograph);
BL 1.5-2.5 mm. Apterae are also
reported to occur (Takahashi 1923), but do not seem to have been
described. On undersides of leaves of Bambusa stenostachya (Takahashi 1923)
and Phyllostachys spp. (Liao 1976)
in Taiwan and China (Tao 1964, Martin & Lau 2011). Quednau (2003) illustrated an ovipara
collected on Phyllostachys in
December in China. In Taiwan it overwinters in the parthenogenetic phase
(Takahashi 1923). There are
conflicting reports as to whether this species does (Liao 1976), or does not
(Takahashi 1921), jump when disturbed.
A
genus containing nominally about 60 small pear-shaped phylloxerids living on
Juglandaceae and/or Fagaceae. More than
half the nominal species were described from galls on Carya in North America (Shimer 1869, as Dactylosphaera; Pergande 1904; Stoetzel & Tedders 1981). Only the pecan-feeding species have been
worked on taxonomically since Pergande's studies, and the status of many of
the other Carya gall-makers is
therefore uncertain. Although many
species are clearly monoecious on either Carya
or Fagaceae, host alternation has now been demonstrated for at least two
species (Stoetzel 1985c). Accounts of
the oak- and chestnut-feeding species are available for Europe (Heinze 1962,
Barson & Carter 1972) and North America (Duncan 1922). Fagaceae-feeding species previously placed
in the genus Moritziella Börner are
now included in Phylloxera (Favret et al. 2016). Grassi (1912) fully
reviewed all information on the group then available. Binazzi & Covassi
(1997) compared life cycles of oak-feeding species in Italy. Hedin et
al. (1985) studied the biochemistry of gall formation on pecan. Stoetzel (1985a) described the development
of the sexuales, through a rapid series of moults of pupiform larvae. The
nomenclatural problems associated with species described in Phylloxera and related genera are
discussed by Favret et al. (2016);
future work may make it necessary to formalise a distinction between most of
the Palaearctic species, which have spiracles on ABD TERG 1-5, and the mainly
Nearctic species, including the numerous Carya
gall-makers, which only ever have them on ABD TERG 1. Phylloxera
bipunctata Lichtenstein
Apterae are yellow with two orange spots, living solitarily on leaves
of Quercus sp(p). in southern
France and Switzerland (Lichtenstein 1874).
Alatae are unknown, and the species has not been recognised since its
original (brief) description. Phylloxera
caryaeavellana Riley Galls projecting underneath leaf are
globular, pale greenish, like a small hazelnut (5-10 mm diam.), above leaf
flat or slightly concave. Galls mature
in May-June, emergent alatae have orange abdomen, BL 0.9-1.2 mm (Pergande
1904). On Carya tomentosa in Washington, D.C. The life cycle is unknown. Phylloxera
caryaecaulis (Fitch) Galls are on young twigs, on petioles or at
bases of leaflet main veins, sometimes in clusters; globular, diam. 5-25 mm,
pale yellowish green tinted with red before opening (see influentialpoints.com/Gallery),
afterwards becoming leathery and black. Typically on Carya glabra in eastern USA.
There is no host alternation.
Alate sexuparae emerging from galls have dusky wings, and in life
there are apparently distinctive dorsal prothoracic markings and a pair of
orange or blackish spots on the posterior abdomen; BL 0.8-1.8 mm (small
alatae produce mostly males, larger ones mostly females). Morgan (1909, 1912, 1915) studied the
cytogenetics and chromosome cycle of this species (for summary see Blackman
1995). Shimer (1869) described three
other closely-related phylloxerid species producing somewhat different twig
or petiole galls on other species of Carya,
which may or may not be synonymous with P.
caryaecaulis: P. spinosa, producing galls covered with fleshy spines on Carya sp, in Illinois (the immature
inhabitants of these galls were described by Shimer as having long spiny
tubercles); P. caryaemagna, producing velvety galls on petioles or at base
of mid-rib on C. amara (= cordiformis) in Illinois and Missouri;
and P. subelliptica, producing ellipsoid mottled green galls on
side of twig of C. alba (= ovata) in Illinois (see also Pergande
1904). Caldwell & Schuder (1979)
studied the life cycle of the form on C.
ovata in Indiana. Shimer (1869)
reported on natural enemies of galls of P.
caryaemagna in Illinois. Phylloxera
caryaefallax Riley Galls on leaves of Carya alba (= ovata)
are thin-walled, 1-5 mm diam., on upper side of leaf and convex with a
central nipple-like opening below leaf.
Alatae with dark-veined wings and orange-yellow abdomen, BL 2.3-2.8
mm, emerge in May-early June (Pergande 1904).
In Missouri, Illinois and states bordering the Mississippi at least as
far south as Tenessee. The life cycle
is unknown. [Morgan's (1912,1915)
cytogenetic study of Phylloxera “fallax” cannot apply to this species,
as Morgan's species was monoecious, with fundatrices that produced apterous
sexuparae; its identity is still in doubt.] Phylloxera
caryaefoliae Fitch Galls on leaves of Carya glabra project above leaf as pale green slender cones with
pale rings at their bases, c. 5 mm diam., almost flat on underside of leaf
with a central nipple. Alatae, with
distinctively marked prothorax and orange abdomen, BL 0.8-1.0 mm, emerge from
the upper side of the gall (Pergande 1904) in early June. In north-eastern USA (New York, Illinois,
Iowa, Virginia). The life cycle is
unknown. Phylloxera
caryaeglobuli Walsh Galls according to original description
are spherical, on leaf lamina between veins, opening in June to release
alatae through a simple longitudinal slit on underside of leaf. On shagbark hickory (= Carya ovata) in Illinois and Missouri. Pergande (1904) recorded the available
information, and suggested that P.
hemispherica Shimer might be a synonym.
The life cycle is unknown. Phylloxera
caryaegummosa Riley Galls are spindle-shaped, suspended from
the underside of leaf of Carya alba
(= ovata) by a slender petiole;
they are white or yellowish, pubescent and sticky. In Illinois and Missouri, USA. Alatae are undescribed and the life cycle
is unknown. Phylloxera
caryaeren Riley
Galls develop in clusters along leaf petioles and mid-ribs of leaflets
of Carya glabra; they are reniform,
pale green, 2-15 mm max. diam., placed transversely to axis of petiole,
deeply divided by a transverse slit and opening into two halves (rather like
a bivalve mollusc) when mature (Pergande 1904). The phylloxerid itself is
still undescribed, and the life cycle is unknown. In Illinois and Missouri, USA. Phylloxera
caryaescissa Riley Galls are globular, diam. 8-14 mm, pale
greenish or dirty yellowish green, clothed with short fine yellowish hairs,
projecting about eqally on both sides of leaf lamina, opening underneath when
mature (mid-May to mid-June) by a transverse slit (Pergande 1904). Alatae have orange abdomen, BL 1.0-1.2
mm. On Carya tomentosa in eastern USA (Florida, D.C.). The life cycle is unknown. Phylloxera
caryaesemen Walsh Galls are rounded, small (diam. 0.3-2.0
mm) and very numerous, yellowish green to brownish yellow, rather tough and
leathery, projecting more above leaf than below, with a more-or-less
depressed centre above and opening (in July-early August) by a central
nipple-like opening underneath. No
alatae are produced; according to Walsh (1868, cited by Pergande 1904), first
instar larvae emerge to found new galls.
On Carya glabra in Illinois
and Missouri, USA; Shimer (1869) also recorded it from C. amara (= cordiformis)
in Illinois. Phylloxera
caryaesepta (Shimer) Galls of Carya alba (= ovata)
have conical projections both above and below leaf, with nipple-like apices
fringed with filaments when mature, the structure on both sides of the leaf
somewhat resembling a miniature sea-anemone (diam. 5-12 mm). Colour of gall varies from pale yellowish
green to red. Alatae with dark orange
abdomen, BL 0.8-1.2 mm, emerge in late May-June from upper side of gall
(Pergande 1904). In Illinois and
Missouri, USA. On Carya glabra further east (New York, Virginia), the same or a
closely-related species forms galls with a narrower conical structure above
and below leaf (P. caryaesepta var.
perforans; Pergande 1904). The life cycle is unknown. Phylloxera
caryaevenae Fitch Galls on Carya tomentosa and several other Carya spp. are distinctive, usually pale yellowish green,
elongate folds along the main veins on the underside of the leaflet, with
corresponding ridges or carinae above.
The opening is a slit on the underside. There is no host alternation. Galls are mostly dry and empty by
mid-August, but no alatae are known; the sexuales are produced from late July
by apterous sexuparae in free-living colonies in deep bark fissures on the
trunk and branches (Pergande 1904).
Widely distributed in eastern USA, from New York to Florida and as far
west as Illinois and Missouri. Phylloxera
castaneae (Haldeman)
Apterae are pale yellow, BL c. 1 mm, on both sides of leaves,
especially along the mid-ribs, of Castanea
spp. (dentata, mollissima, pumila, sativa) in north-eastern USA (New
York, Maryland, D.C., West Virginia).
Infested leaves become much distorted and desiccated. Stoetzel (1985b) demonstrated host
alternation in West Virginia from galls on Carya tomentosa. The galls
and gall generations have not yet been described; the galls are similar to
those of P. intermedia (M.B.
Stoetzel, pers. comm.). Alatae from
these galls gave rise to large populations on Castanea spp., causing yellow discoloration along the leaf veins.
Pergande (1904) drew attention to the fact that most of the immatures on the
upper sides of the chestnut leaves were bright orange (as opposed to pale
yellow), and had longer tubercles than those on the undersides; his name spinifera is available for this form
should it prove to be specifically distinct. Phylloxera
coccinea (von Heyden)
Fundatrices in spring cause crinkling and discoloration of margins of
young oak leaves. Later generations of
apterae (yellowish orange, BL 0.8-1.2 mm) lay eggs in compact circles and
cause necrotic spots on undersides of leaves (see Börner & Heinze
1957). Living without host alternation
on Quercus spp. (especially robur) throughout most of continental
Europe, and east to Kazakhstan. Both
apterous and alate sexuparae are produced (Börner & Heinze 1957). Anholocyclic overwintering may occur;
Lubiarz (2007) observed overwintering of immature stages of P. coccinea (and/or P. glabra) in bark crevices in Poland.
P. rutila Dreyfus is a synonym
(Börner 1952). New Zealand records
apply to Moritziella corticalis
(Sunde 1974). Phylloxera
confusa (Grassi)
Fundatrices feed from veins on undersides of oak leaves, causing the
lobe of the leaf distal to the feeding point to fold inwards. Later generations of apterae (BL 1.5-.2.0
mm) are yellowish to orange-yellow and develop in small groups on young
twigs, on leaf scars or under loose bark (Grassi 1912). There is no host alternation; sexuparae are
apterous. On Quercus robur, only recorded from central Italy. Phylloxera castaneivora
(Miyazaki) Apterae are pear-shaped,
broadest anteriorly, yellowish brown to purplish brown; BL 0.9-1.3 mm. Feeding on fruits of Castanea crenata in Japan (Miyazaki 1968b, as Moritziella). Yanagibashi & Nakagaki (1985) studied
the influence of temperature on development of the apterae. Other morphs and life cycle are unknown. This species was recently reported as an
important pest of chestnuts in China (X. Wang et al. 2009, as Moritziella). Phylloxera
conica (Shimer)
Galls are almost globular, projecting on both sides of the leaf,
diameter 3-7 mm in plane of leaf, deep purplish brown, with a roughened or
granular surface texture, becoming pubescent and covered with a sticky
secretion when mature. Alatae, BL
0.8-1.2 mm with a pale yellow abdomen, emerge when the gall splits open
underneath in June, and presumably fly to an unknown secondary host. Originally described from Carya alba (= ovata) in Illinois, USA (Shimer 1869), but the description above
is of galls on C, amara (= cordiformis) in Missouri identified as
this species by Pergande (1904). Riley
(1875) suggested synonymy with P.
depressa, but this does not accord with Pergande's interpretation of the
species. Phylloxera corticalis
Kaltenbach Apterae are pear-shaped,
broadest anteriorly, green or brownish, with dark brown dorsal tubercles; BL
0.6-0.8 mm. On Quercus robur and Q.
petraea, colonising bark of previous year's growth (Heinze 1962), and
sometimes causing serious dieback of shoots (Barson & Carter 1972, as Moritziella). In Europe, and introduced to New Zealand
(Sunde 1973, as Moritziella). Life cycle unclear, but probably mainly
anholocyclic; alate sexuparae may be produced in summer and autumn, but no
sexuales have been recorded (Barson & Carter 1972). M.
corticalis is barely distinct from some little-known North American
species described in the genus Phylloxera
(e.g. davidsoni, querceti), and may be an introduced
North American species that has lost its primary host (Carya; see Stoetzel 1985). Phylloxera
davidsoni Duncan
Apterae light orange to yellowish brown; BL c. 0.7 mm. Immatures are pale yellow, alatae orange
with black thorax. On undersides of
tomentous young leaves of Quercus engelmanni
in California, USA (Duncan 1922). The
life cycle is unknown. Phylloxera
deplanata Pergande
Galls on leaves of Carya
tomentosa are circular, diameter 1-5 mm, slightly convex above leaf and
more prominent beneath, with an oval orifice fringed with fine hairs. Their colour is reddish to yellowish or
greenish yellow above leaf and purplish to dull greenish yellow beneath. There is no host alternation; fundatrices
produce sexuales in the galls in May, but some alatae may also be
produced. Galls are empty, brown and
dry by late June (Pergande 1904). Only
known from D.C., USA. Phylloxera
depressa (Shimer)
Galls on leaves of Carya alba
(= ovata) in Illinois, USA are
numerous, smooth, diameter 4-6 mm, convex and pale yellowish green above
leaf, whiter beneath with a central nipple-like projection and a round
opening. Alatae, BL c. 1 mm with a
yellow abdomen, are present in the gall in late June (Shimer 1869). The life cycle is unknown. According to Riley (1875), conifera Shimer is a synonym. Phylloxera
devastatrix Pergande Clusters of galls occur on young twigs,
leaf petioles or flower stalks of Carya
illinoensis in southern and south-eastern USA (Arkansas, Louisiana,
Mississippi, Oklahoma, Texas), often causing serious economic damage. The galls are globular, of irregular shape,
smooth, green or yellowish green with a reddish tinge before maturity,
diameter 3-15 mm. They open in
May-early June by means of a transverse split into four or more broad bracts,
releasing alatae that immediately start to oviposit on neighbouring twigs,
etc. There is no host alternation. The
eggs develop into dwarf sexuales that mate and lay overwintering eggs under
bark or at the base of the tree. H.
Baker (1935) described the life cycle in Louisiana, and reported a high
incidence of parasitism of galls.
Carpenter et al. (1979) and
Calcote (1985) studied varietal and clonal resistance to P. devastatrix in pecan, Neel & Hedin (1985) studied its
effects on nut production, and Hedin et
al. (1985) studied biochemical aspects of gall formation. Mitchell et al. (1984) studied the gall-associated fauna. Phylloxera
foae Börner
Living without host alternation on deciduous oaks, with a life cycle
similar to that of Ph. coccinea,
but Q. petraea is the usual host
plant. Fundatrices usually in small
leaf-edge folds; subsequent generations of apterae yellowish orange, BL
0.7-0.9 mm, causing necrotic spots on undersides of leaves. Sexuparae are alate. Closely related to Ph. coccinea; Heinze (1962) provided some distinguishing
characters. In Italy, Germany,
Ukraine. Phylloxera
foveata (Shimer)
Galls on leaves of Carya amara
(= cordiformis) are small (1.5-8.4 mm diam.), with a
dimpled central depression above, opening beneath leaf by a rounded downy
orifice. Alatae were plentiful in late
June. In Illinois, USA (Shimer
1869). The same leaves carried galls
of P. minima and P. caryaesemen. Phylloxera
foveola Pergande
Galls on leaves of Carya glabra
in Virginia, USA are circular, yellow-green tinged with pink or red, 3-5 mm
diam., convex and rather prominent above leaf with a deep central dimple or
depression which is pinkish in colour, part below leaf convex or depressed
with a small conical central nipple.
Alatae (BL c. 1.2 mm, with slender, orange abdomen) are mature in
galls in late May and most had emerged before June, indicating that there is
probably host alternation, but the secondary host is unknown. Possibly it is a synonym of P. foveata; the emergence time was the
principal reason given for separating the two (Pergande 1904), but this might
vary according to weather conditions or host plant. Phylloxera
georgiana Pergande
Galls occurred singly or in clusters on tender twigs and petioles of
an unidentified Carya sp.,
presumably at Georgiana, Florida (Pergande 1904). They were similar to those of Ph. devastatrix; more-or-less
globular, with a short nipple, greenish yellow (sometimes reddened on one
side), or wholly dark red, 5-10 mm in diameter. Alatae (BL 0.8-1.0 mm, with orange abdomen)
differ from P. devastatrix in
characters of the wings and antennae (see Pergande 1904). Phylloxera
glabra (von Heyden)
Plate 2a Living without host alternation mainly on Quercus robur. Fundatrices feed in spring from main veins,
sometimes causing inward curling of leaf-edge as in Ph. confusa. Subsequent
generations include yellowish orange apterae (BL 0.7-0.85 mm), which lay
circular clusters of eggs on the underside of the leaf, and both apterous and
(later) alate sexuparae (Grassi 1912).
Feeding causes necrotic spots on leaves. In Europe (England, France, Germany,
Italy), and introduced to New Zealand, where it is recorded from Q. dentata (Sunde 1984). A record from Turkey (Şenol et al. 2014c) requires additional
confirmation. Anholocyclic overwintering may occur; Lubiarz
(2007) observed overwintering of immature stages of P. glabra (and/or P.
coccinea) in bark crevices in Poland. P.
glabra is regularly preyed upon by the coccinellid Pullus auritus and the coniopterygid Conwentzia psociformis. Phylloxera
globosa (Shimer)
Galls on leaves of Carya amara
(= cordiformis) in Illinois are
mostly rather large, globular and projecting equally on both sides of leaf, or
conical and projecting only below leaf, with flat circle above ("form conifera"); smooth, leathery,
pale yellowish green to dark green, with a small central opening below leaf
surrounded by small bracts. [According
to Pergande (1904), the smaller galls included in Shimer's (1867) original
description were those of Ph.
caryeaesemen.] Alatae (BL c. 0.6
mm, with orange-yellow abdomen), emerge very late in year, in late
September-October. The life cycle is
unknown. Phylloxera
ilicis Grassi Plate 1e, f Anholocyclic on Quercus ilex in central Italy, and alatae have apparently been
trapped in Tunisia (Boukhris-Bouhachem et al. 2007). Several
generations of yellow apterae occur through spring and summer feeding mainly
on current and 1-year-old twigs, and causing small tuberosities; but
sometimes also on foliage causing necrotic spots like P. quercus. Alate
virginoparae occur in autumn generations (Grassi 1912). Phylloxera
intermedia Pergande
Galls on leaves of Carya alba
(= ovata) are circular, reddish or
yellowish, 3-6 mm in diameter, convex to conical on upper side of leaf
without any trace of a central nipple, paler and almost flat with a small
closed nipple on underside. Alatae (BL
0.8-1.3 mm, with pale orange abdomen) emerge in late may-June, mainly from a
jagged opening formed by splitting of the upper surface of the gall (Pergande
1904). Locality not stated; probably in vicinity of Washington, D.C. The life
cycle is unknown. Phylloxera
italica Grassi
Living without host alternation on Quercus
robur, and only known from central Italy.
Fundatrices develop in April-May and the next two generations consist
only of apterae (whitish, Bl c. 1.5 mm), which feed on the undersides of
leaves causing small, sparse, whitish spots.
Sexuparae are mostly apterous (Grassi 1912). Phylloxera
kunugi Shinji
Apterae (fundatrices?) are bright yellow, BL 2.3-2.5 mm. They feed in spring from mid-rib on
undersides of young leaves of Quercus
spp. (acutissima, variabilis) in Japan, causing the
infested leaves to curl downward. All
their progeny are alate (BL 1.8-2.5 mm), and apparently migrate to another
host; the full life cycle remains to be established (Miyazaki & Teramoto
1991). Phylloxera
minima (Shimer)
Galls on Carya amara (= cordiformis) in Illinois USA are
small, round (diameter 1.1-5.0 mm) and numerous, with part protruding above
leaf shining brightly, and a small opening beneath. Alatae, present in the galls in early July,
were translucent, yellowish white with minute dark spots on anterior part of
body (Shimer 1869). The life cycle is
unknown. Phylloxera
notabilis Pergande
Galls on leaves of Carya
illinoiensis are ovoid to spherical, bladder-like, very variable in size
(diameter 2-18 mm), situated close to mid-rib, projecting on both sides of
leaf, green on top with a central nipple, and reddish-green beneath covered
with dense short yellowish hairs.
Alatae (BL 0.6-0.8 mm, with dusky wings and yellow-orange abdomen),
emerge from the top of the gall, which generally splits into long slender
bract-like segments in June-July.
There is no host alternation. Several generations of galls are
produced each year; the smaller, secondary galls occurring mainly on seedling
trees (Stoetzel & Tedders 1981).
Brachypterous and migratory alatae, sexuparae and sexuales are all
produced in the primary galls.
Sexuales lay overwintering eggs on bark (Whitehead & Eastep 1937). In eastern and southern USA. Andersen & Mizell (1987) studied the
physiological effects of galling on the host. Calcote (1983) studied
resistance of pecan clones to P.
notabilis. Medina et al. (2011)
demonstrated that populations identified as P. notabilis collected from water hickory (C. aquatica) in east
Texas always lacked two species of
symbiotic bacteria that were present in populations on pecan. There were also
found to be distinct differences in DNA (AFLP) profiles, and in host
preferences and performance of alatae in reciprocal transfers between the two
Carya species (Dickey & Medina
2010, 2011), all of which indicate that the phylloxerid on water hickory is a
different, presumably undescribed, species. Phylloxera
perniciosa Pergande
Galls on young twigs, leaf petioles and along midribs of leaflets of Carya tomentosa are elliptical, with a
blunt tooth-like or thorn-like projection.
In Maryland, Virginia, D.C. and Pennsylvania; large gall populations
in Virginia caused defoliation of trees (Pergande 1904). Also reported from C. illinoensis in Louisiana and Mississippi (Carpenter et al. 1979), although the identity of
these populations with Pergande's species needs confirmation. Living all year around on Carya, with three generations per
year; fundatrices, alate sexuparae and sexuales. Alate sexuparae emerging from galls in
May-early June laid eggs indiscriminately on trees and surrounding
vegetation. Sexuales laid
overwintering eggs in June, mostly high in trees on twigs that had been
previously galled. The eggs were
preyed upon through the summer by various natural enemies. including mites of
six different genera. Surviving eggs
hatched the following April (Pergande 1904). Phylloxera
picta Pergande
Galls on leaflets of Carya
tomentosa resemble those of Ph.
intermedia, but have thinner walls, adn a central nipple above as well as
below the leaf; colour variable, diam. 3-6 mm. Alatae (BL 0.8-1.3 mm, with pale orange
abdomen) emerge from an opening on the upper side in late May-June (Pergande
1904). Locality not stated; probably
in vicinity of Washington, D.C. The
life cycle is unknown. Phylloxera
pilosula Pergande
Galls on leaflets of Carya
glabra near Washington, D.C. are flat or with a central depression above
leaf and projecting with a well-developed central nipple below leaf; diam.
3-6 mm, pale green to whitish, both sides densely covered with long, soft,
glistening, yellowish white hairs.
Alatae (BL c. 1.2 mm, with orange abdomen) emerge from the underside
of the gall in early June (Pergande 1904).
The life cycle is unknown. Phylloxera
querceti Pergande
Apterae (= fundatrices?) are yellow to orange, BL c. 0.5 mm. On undersides of leaves of Quercus spp. (alba, dentata(?), macrocarpa) in the vicinity of
Washington, D.C. (Pergande 1904), Duncan 1922). Affected leaves are speckled with small
yellow spots, especially near mid-ribs and along smaller veins. Alatae of BL 0.6-1.0 mm, with orange
abdomen. The life cycle is unknown. Phylloxera
quercina Ferrari (=
spinulosa Targiani-Tozzetti) Apterae are pale yellow, with dorsal
processes whitish (but darker in spring and autumn generations); BL 0.9-1.5
mm. On undersides of leaves of Quercus
spp. (cerris, ithaburensis, suber) in
the Mediterranean region (Italy, Israel, Portugal), causing yellow
spots. Phylloxerids identified as this
species have also been recorded from Slovenia, Turkey and Iran. Living
without host alternation on Q. cerris
in central Italy, 3-6 parthenogenetic generations occurring annually, with
the proportion of alate sexuparae increasing in successive generations
(Grassi 1912, as Acanthaphis spinulosa). Phylloxera
quercus Boyer de Fonscolombe (= florentina
Targiani-Tozzetti) Apterae are
greenish yellow to orange-yellow; BL 0.7-0.9 mm. On undersides of leaves of Quercus spp., causing yellowish
spots. In southern Europe and
south-west Asia, and heavy infestations on Q. ilex were recently reported from Algeria (Laamari 2016).
(Reports from northern Europe are possibly all based on misidentification of P. glabra.) The life cycle is complex
(Grassi 1912). Overwintering usually
occurs as eggs on evergreen oaks, Q.
coccifera or Q. ilex. Progeny of the fundatrices include alatae
which fly in May to colonise deciduous oaks for the summer months (petraea, pubescens, robur), and
also apterae which continue generations on Q. ilex, etc. In both cases
alate sexuparae appear in the fourth and fifth generations, and those
produced on deciduous oaks mostly return to evergreens to give rise to sexual
morphs. Overwintering as young larvae
may also occur (see Grassi 1912 and Antonelli 1989 for further details). Mohammed & Abdulla (1985) studied the
ecology of populations identified as P.
quercus in Iraq. Phylloxera
reticulata Duncan
Apterae are bright orange-yellow, with reddish eyes; BL c. 0.8
mm. On undersides of leaves of Quercus kelloggii in California, USA,
dispersed along veins. Only apterae
are described (Duncan 1922), and they have not been recorded since. Phylloxera
rileyi Riley
Apterae are dark brown with dorsal tubercles almost black (but paler
forms mat occur); BL c. 0.6 mm. On
undersides of leaves of Quercus alba
and Q. obtusiloba (= stellata) in Missouri, causing white
or yellow circular spots. Apparently
there is no host alternation; alatae are produced from late July to October, and
those of later generations are presumably sexuparae, because sexuales are
recorded laying overwintering eggs on twigs (Riley 1875, Pergande 1904,
Duncan 1922). Phylloxera
rimosalis Pergande
Galls on Carya tomentosa in
eastern USA (Washington, D.C.) are fairly large and conspicuous (diam. 3-11
mm), pale green sometimes tinged with red, usually on terminal leaves of
young shoots; flat to slightly convex above leaf, more prominently convex
with a central nipple beneath. Alatae
(BL c. 1 mm, with pale orange abdomen) emerge in late May-June through a
round opening on the underside (Pergande 1904). The life cycle is unknown. Phylloxera
russellae Stoetzel
Galls on Carya illinoensis
in southern USA are circular, flattened, diam. 3.2-7.2 mm, with a reticulate
surface pattern. They project from
both sides of a leaflet with an opening on the underside marked by white
pubescence. Galls mature and break
open in May. No alatae are known. There is no host alternation; apterous
sexuparae produce sexuales which develop in the gall in May and produce
overwintering eggs in galls, on twigs, etc. (Stoetzel 1981). Heavy populations of galls can cause
defoliation. Calcote (1985) studied
resistance of pecan clones to P.
russellae. Phylloxera
similans Duncan
Apterae of BL c. 0.7 mm, appearance in life unrecorded. Collected on Quercus macrocarpa in New York State, USA (Duncan 1922), and not
recorded since the original description.
Its biology is unknown. Phylloxera
spinuloides Pergande Galls on petioles or at bases of main
veins of leaflets, of an undetermined Carya
species in Florida, USA, were globular (diam. 5-15 mm), with a dense covering
of long fleshy filamentous outgrowths; usually encircling the petiole and
causing it to bend or curve. Colour
uniformly greenish yellow, or brownish yellow on one side. Galls opened in late March, releasing
alatae (BL 0.6-1.8 mm, with brownish wings and orange abdomen). Life cycle is unknown, although the early
departure of alatae suggests host alternation. Phylloxera
stanfordiana Ferris Apterae are pale yellow, BL c. 0.75
mm. On twigs of Quercus douglasii in California, USA, in September,
"concealed beneath any chance covering, especially numerous under the
twig-encircling egg masses of a moth" (Ferris 1919). Other morphs and life cycle are unknown. Phylloxera
stellata Duncan
Apterae (BL c. 0.5 mm, colour unrecorded) caused brown spots on
undersides of leaves of a Quercus
sp., margaretta (= stellata or alba, in mid-August in Texas, USA (Duncan 1922). Other morphs and
life cycle are unknown. Phylloxera
symmetrica Pergande
Galls on leaves of Carya
tomentosa and other Carya spp.
are variable; typical form is circular and almost flat above leaf, protruding
as a thick-walled pale cone below leaf, basal diam. 2-5 mm. Alatae (BL 0.9-1.2 mm, with orange abdomen)
emerge in late May-early June through a small opening in the apex of the
cone, after which the galls become dry and brown. In vicinity of Washington D.C., USA. Life cycle is unknown. Pergande (1904) described variants from the
typical form of gall ("vasculosa"
and "purpurea") differing
in colour, size and shape, which he considered to be due to differences in
hatching time of the fundatrices and developmental stage of the leaf at onset
of feeding. Phylloxera
texana Stoetzel
Galls on leaves of Carya
illinoensis are globular, projecting mainly on upper side of leaf, with a
small point below; diameter 4-6 mm (Stoetzel 1981). Alatae (BL 1.0-1.6 mm, with golden yellow
abdomen) emerge in mid to late May through a large opening in the top of the
gall, which splits into bracts. In
Texas, USA. Host-alternating; alatae
from galls colonised undersides of leaves of Quercus spp. (falcata, virginiana), these colonies producing
both alatae and apterae in June (Stoetzel 1985b). The return migration to pecan has not been
observed. Phylloxera
tuberculifera Duncan Apterae (BL c. 0.5 mm, colour unrecorded)
caused brown spots on undersides of leaves of Quercus havardii in Texas, USA, in August (Duncan 1922). Other morphs and life cycle are unknown. Phylloxera
sp. Apterae are pear-shaped, dirty
yellow to dark brown; BL 0.7-1.0 mm.
On Quercus virginiana in
Cuba, colonising apical parts of young shoots, mainly on the stem but also on
young leaves which may be slightly curled or rolled. Perhaps an anholocyclic race of a North
American species described in another genus (Holman 1974, as Moritziella).
About
nine species mostly living without host alternation on Salicaceae, with one
on Nyssa (Cornaceae). Apterae are
usually covered in dense white wax. No
alatae have ever been recorded for this genus. The species on Salix in Germany were studied by
Iglisch (1965). Phylloxerina
capreae Börner Plate 2b Apterae
are yellow to brownish yellow, under dense coat of white wax filaments; BL
0.6-0.9 mm. On Salix caprea stems
and branches, in bark crevices or under scales. In Europe (Germany; although
a phylloxerid on Salix lanata in
England, BMNH collection, leg. V.F. Eastop, has been provisionally assigned
to this species), and also reported and redescribed from Japan (Miyazaki
1972b). Living without host alternation on Salix; see Iglisch (1965) for life cycle details. Phylloxerina
daphnoidis Iglisch
Apterae are light brown, under dense coat of white wax filaments; BL
0.6-0.9 mm. Living without host
alternation on Salix daphnoides and
S. elaeagnos in Germany, in bark
crevices, etc. Iglisch (1965) gave
life cycle details and compared P.
daphnoidis with the very similar P.
capreae. The two species cannot be
reliably distinguished on morphology alone. Phylloxerina
nyssae (Pergande)
Galls are usually crescent-shaped, yellowish-white, formed in spring
by rolling, scalloping and swelling of edges of leaves of Nyssa sylvatica (Stoetzel 1993). Later
in the year (September), lemon-yellow apterae, BL 1.0-1.3 mm, completely
covered in white wax wool, occur together with sexuales in bark crevices of
young trees, particularly in cavities caused by dropping out of dead twigs
about 2 m above ground level. Apparently there is no host alternation. In
Maryland and Washington D.C., USA (Pergande 1904). Phylloxerina
popularia (Pergande)
Apterae are pale greenish yellow, covered with white cottony wax, BL
c. 1.4 mm, living in deserted galls of Pemphigus
populitransversus and Pemphigus
populicaulis, in bark crevices or on branches and twigs, of North
American cottonwoods (Populus deltoides,
P. fremontii and P. trichocarpa) in Louisiana, Michigan,
Missouri, South Dakota, Texas and California (Pergande 1904, Davidson
1915). There is no host alternation;
apterous sexuparae and sexuales were found in old Pemphigus galls in October (Pergande). Phylloxerina
populi (Del Guercio)
Apterae are pale yellow to ochre, covered in wax wool; BL 0.5-0.7
mm. Alatae have not been recorded.
Living without host alternation on cortical tissues of Populus spp. (alba, ×canescens).
Described originally from Italy, and redescribed in detail from Hungary
(Ripka 2011), who found it on branches of P.
×canescens living under scales of dead females of
Chionaspis salicis (Diaspididae)
along with several mite species, and also on P. alba in woody galls caused by an eriophyid mite. Bonfigli
(1909; as Pseudochermes populi) and
Ripka (2011) have both compared it with P.
salicis. Phylloxerina
prolifera (Oestlund)
Apterae are very pale lemon yellow, found in autumn in empty galls of Pemphigus populicaulis in Minnesota,
USA (Oestlund 1887). Not recognised
since original description, and possibly not distinct from P. popularia, despite the apparent
differences noted by Pergande (1904). Phylloxerina
salicola (Pergande)
Apterae are yellow to brownish yellow, covered with white wax wool; BL
0.6-0.9 mm. Originally described from
bark crevices or other cracks in stems and twigs, and sometimes also on undersides of leaves, of an unidentified Salix species (either discolor or humilis) in eastern USA (Pergande 1904). Sexuales are present in September. Davidson (1915) recorded this species from Salix lasiolepis in California,
usually living in bud axils on twigs but also on one occasion heavily
infesting bared roots on a stream bank.
These Californian populations appeared to be anholocyclic, and could
possibly have been introduced European P.
salicis. Phylloxerina
salicis (Lichtenstein)
Apterae are pale green, yellow or brownish, secreting profuse white
wax wool; BL 0.7-1.2 mm. Living in
bark crevices on trunk and branches of Salix
alba and related species in western, southern and central Europe, and
across temperate Asia to China (G. Zhang et
al. 1985) and east Siberia (Pashchenko 1988a). Introduced to Victoria, Australia (Eastop
1966), and perhaps to North America (Börner & Heinze 1957). Sexuales occur in September (Iglisch 1965). Phylloxerina sp. Holman (1964) described (but did not name)
a species of Phylloxerina on trunks
of Salix longipes in Cuba. The life cycle is unknown. Similar to European P. capreae (but not compared with the North American species).
Clearly
differentiated from the other adelgid group, Adelges, by their biology - e.g. gall structure, secondary host
association with Pinus - and by
various morphological features. The most
consistent character that applies to all morphs is the reduction in the
number of abdominal spiracles to only four evident pairs. The galls, usually on shoot tips of Picea, are generally less compact than
those of Adelges and have
intercommunicating chambers. Adult
apterous morphs on secondary hosts (sistens and progrediens) are very broadly
pear-shaped - often almost globular - and have a fused and pigmented head and
prothoracic shield. Pineus
species may be holocyclic and host-alternating between Picea and Pinus,
completing a cycle with seven morphs in two years as in holocyclic Adelges; or they may be anholocyclic
on either Picea or Pinus (except for Pineus abietinus, which is anholocyclic on Abies). There is also one
species, Pineus floccus, that is
now known to have an anholocyclic, annual host alternation between Picea and Pinus. Knowledge of the
life cycle of many species is still incomplete. Several anholocyclic species on Pinus produce alate sexuparae which
appear to be non-functional, either because they are unable to produce males
or because they fail to find a suitable Picea
species to act as primary host. Systematic
accounts of Pineus are available
for Britain (Carter 1971), Europe (Börner & Heinze 1957, Heinze 1962),
India (A.K. Ghosh 1984a), Japan (Inouye 1953) and North America (Annand
1928). Favret et al.
(2015b) provided a taxonomic and nomenclatural catalogue of the world fauna. Havill et
al. (2007) studied the evolution of host relationships based on molecular
data. Havelka et al. (2019)
analysed mitochondrian and nuclear DNA sequnces of the species in Lithuania. Pineus
abietinus Underwood & Balch (fig. 10A) Only the first instar and adult progrediens
(aestivoprogrediens) are described; adults are very small (about 0.6 mm),
light reddish-brown with dark cephaloprothoracic shield, secreting white
wax-wool, in bark crevices on stems and branches of Abies amabilis and A.
grandis in the north-west coastal region of North America (Washington,
southern British Columbia). This is
the only known Pineus associated
with Abies. Wax pores are circular and
discontiguous. Apparently anholocyclic
and monomorphic, probably with two or three progrediens generations per year
(Underwood & Balch 1964). Pineus
armandicola Zhang, Zhong & Zhang Galls on Picea likiangensis in Yunnan, China are cone-like or torch-like
with multiple cavities (Li et al.
2002). Migration occurs to Pinus
armandii. Apterae on Pinus are red-brown, with wax threads;
BL 0.8-1.4 mm (G. Zhang et al.
1992c). Pineus
boerneri Annand
Apterae are small, reddish brown, laying eggs on pine twigs in
abundant greyish-white wax wool. Described from Pinus radiata in California (Annand 1928), but possibly of east
Asian origin, where it has mainly been recorded under the name P. laevis (McClure 1984b). It is now
also in north-eastern USA, Hawaii, South America (originally as P. havrylenkoi Blanchard), Australia,
New Zealand and east Africa (as P. pini).
P. simmondsi Yaseen & Ghani, on
Pinus roxburghii in Pakistan
(Yaseen & Ghani 1971), could also be this species. P. boerneri is anholocyclic everywhere, and the introduced
populations attack a range of exotic pines from both Europe (e.g. P. halepensis) and North America (e.g.
P. radiata, P. caribaea). In Australia and New Zealand the name laevis has been applied to both boerneri and pini (q.v.), which are difficult to differentiate except by using
multivariate morphometrics and karyotype analysis (Blackman et al. 1995). Biological control of boerneri and/or pini has been successful using Leucopis sp. (Chamaemyidae) in New Zealand, Chile and Hawaii
(Mills 1990). Dynamics of east African populations was studied by Mailu et al. (1980, as pini), and in the rather different conditions of Pinus resinosa plantations in
Connecticut, USA by McClure (1989b, 1990).
Cardoso (2007) has made a detailed study of its morphology, ecology
and feeding behaviour in Brazil. 2n=16 (Hawaii) or 17 (California, Africa,
Australia, New Zealand). Pineus
boycei Annand (fig.
64B) This species
forms 2-4 cm long galls on Picea
engelmanni, green with a reddish tint, at the bases of the needles,
usually at the shoot tips. It is only
recorded from Montana and Oregon, USA.
Alate gallicolae emerging from the galls in the second half of July
are 1.7-2.5 mm, with abundant wax posteriorly on the reddish brown abdomen
(Annand 1928). Their destination is
unknown. No other morphs of life cycle
details are known (although it seems possible that alate specimens described
by Annand under the name Pineus
engelmanni are the sexuparae of this species). Pineus
cembrae (Cholodkovsky) (= sibiricus Cholodkovsky)
Host-alternating between Picea
spp. (abies, glehnii, obovata) and
stone pines, Pinus spp. of
subsection Cembrae (cembra, pumila, koraiensis, sibirica);
of boreo-alpine distribution in Europe, and extending across Asia to
Japan. Other pines are not usually
attacked (Potopova 1984), except that P.
parviflora is recorded as a host in Japan. Galls are 4-5 cm long, often asymmetrical,
distorting shoot tips, with needles concentrated on one side. Gallicolae are dark red and secrete abundant
wax, migrating to Pinus in June in
Germany (Börner & Heinze 1957), but in August in Japan (Inouye
1953). The neosistens overwinter
mainly near ends of twigs at the bases of last year's pine needles, giving
rise the following year to alate sexuparae and one or two generations of
apterous progredientes living in dense wax-wool on new shoots. Inouye (1953) describes and illustrates all
morphs. A form collected on the
needles of P. koraiensis in
Heilongjiang Province, China is described as a subspecies, P. cembrae pinikoreanus (G. Zhang
& Fang 1981). 2n=18. Pineus
cladogenous Fang & Sun Anholocyclic on Pinus koraiensis in Heilongjiang, China, overwintering in
fissures in the bark. The adult
hiemosistentes (collected in April) are small and ladybird-shaped; BL about
0.6 mm (Fang & Sun 1985). Perhaps
closely related to P. harukawai. Pineus
coloradensis (Gillette) Anholocyclic on numerous Pinus spp. in USA and Canada. Apterous progredientes feeding on the
needles secrete abundant wax, which in summer becomes covered with a dark
mould. One hiemoprogrediens and two aestivoprogrediens generations
occur in New England, where this adelgid has been injurious to plantations of
P.resinosa (McClure 1989b). Alate sexuparae, presumed non-functional,
are recorded from Colorado, but not from California or Connecticut. Descriptions of morphology and biology in
comparison with related species are given by Annand (1928), Doane (1961) and
McClure (1989b). McClure (1984a)
studied interactions between this adelgid and Pineus boerneri on Pinus
resinosa, and Huff & McDonald (1977) studied differences in its
susceptibility to numerous white pine species. Pineus
cortecicola Fang & Sun Anholocyclic on Pinus koraiensis in Heilongjiang, China, overwintering under bark
in only a little wax (Fang & Sun 1985).
BL of adult hiemosistens about 0.85 mm. From the description this species appears
similar to P. hosayai, but differs
in overwintering site and amount of wax. Pineus
floccus (Patch)
"Host-alternating" between Picea spp. (rubens, mariana) and Pinus strobus in eastern USA, but the host alternation revealed
by life history studies in Virginia is remarkable because it occurs on an
annual basis, and only involves parthenogenetic morphs; there are no
fundatrices, sexuparae or sexuales (Walton 1980). There is a single generation (gallicolae)
on Picea, and a series of
progrediens generations on pine.
Overwintering apterous progredientes on P. strobus produce a spring generation of both apterous and alate
progredientes, and the latter fly to spruce buds in May. Galls are induced by the developing
gallicolae. They are usually terminal,
1.8-4.5 cm long, resembling those of Adelges
cooleyi but shorter, stouter and looser; green, but acquiring a purplish
or reddish tinge as they mature, the protruding ends of the needles remaining
green and of normal length. Adult
gallicolae (fig. 64D),
0.9-2.0 mm with abundant wax on head and thorax and a yellow-brown abdomen,
migrate in July to both old and new needles of Pinus strobus. Apterous
adults of the second generation on pine overwinter at the bases of buds,
needles or stem scales, moving to growing points in spring. In New York, however, where heavy galling
of Picea rubens in plantations is
recorded, Deboo & Weidhaas (1967) believed that gallicolae were
recolonising spruce directly and not migrating to pine. Pineus
ghanii Yaseen & Ghani Anholocyclic on Pinus griffithii in the Galis and Kaghan valleys, Pakistan,
attacking stems and twigs of mature and old trees (Yaseen & Ghani
1971). There are two generations per
year, a hiemosistens becoming adult in April, and an aestivosistens becoming
adult in August-September. Adults are
light reddish-brown, 0.9-1.2 mm. Pineus
harukawai Inouye
Anholocyclic on white pines (Pinus
parviflora, P. strobus) in
Japan (Hokkaido, Honshu, Kyushu), on stems or older branches of young pines,
or at the bases of the needles on young branches of older pines. Only the adult of the overwintering sistens
- about 0.9 mm and orange to brownish-red with abundant wax, laying large
numbers of eggs in April to May - is described; other details of the life
cycle are unknown. This species is
sometimes injurious in pine nurseries in Hokkaido (Inouye 1953). Pineus
hosoyai Inouye
Only the apterous rogrediens and immature sexupara are described,
collected from Pinus koraiensis in
June in Korea, and from Pinus sp.
in May in China (Inouye 1953). The
overwintering sistentes were found in February in Korea, living in abundant
wax on shoots of the previous year's growth.
The life cycle is unknown. Pineus
matsumurai Inouye
Apterous progredientes and alate sexuparae develop in June at needle
bases on young branches of Pinus
thunbergii and P. densiflora in
Japan (Inouye 1953). The apterous
progrediens is small (about 0.8 mm), dark reddish-brown, covered in dense
fine wax-wool. The destination of the
sexuparae and the rest of the life cycle is unknown. This species seems to be close to Pineus strobi; differences are
discussed by Inouye (1953). Pineus
orientalis (Dreyfus)
(fig. 64C) Host-alternating between
certain Picea spp. (orientalis, polita, jezoensis, but
not abies) and various Pinus spp. of the subsection (densiflora, mugo, sylvestris, thunbergii) from western Europe across
Asia to Japan. Galls are terminal,
shorter and usually straighter than those of P. cembrae, with protruding needle tips evenly distributed around
the gall. Gallicolae (Plate 1d, fig.
64E) fly
to Pinus in late June to early July
and lay egg-masses in wax wool on the previous year's needles. There are numerous acounts of the biology
of this adelgid, which is not differentiated from its anholocyclic close
relative P. pini in some of the
earlier literature (e.g. in Europe: Börner 1908, Marchal 1913, van der Goot 1915,
Mordvilko 1935; and in Japan: Inouye 1953). 2n=20. Pineus
pineoides (Cholodkovsky)
(fig. 64A) Anholocyclic, with one sumer
and one winter generation per year, both of apterae which live on well-shaded
parts of the bark of the trunk and larger branches of Picea spp. (abies, obovata, omorika, rubens). The insects are yellowish-grey to dark
red-brown, dorso-ventrally flattened, with the cephaloprothoracic shield of strongly
pigmented specimens heavily sclerotised; BL 0.6-0.8 mm. Each lays its eggs under a separate tuft of
wax wool. The overwintering generation becomes adult in May. The adults of
the summer generation, similar except that they have rather fewer wax pores,
may be found in July. In Europe,
Turkey (Şenol et
al.
2014), Canada, and also recorded from bonsai spruce in Australia (New South
Wales; BMNH collection, leg. D. Koreshoff).
For details of biology see Steffan (1963, in Germany) and Underwood
(1963, in Canada). Pineus konowashiyai, on Picea abies in Japan, has a similar habitat and life cycle to P. pineoides, but differs in having
large discontiguous wax pores (Inouye 1953). 2n=22 (Steffan, 1968) or 17
(Blackman & Eastop 1994). Pineus pini
(Macquart) Plate
1c, fig. 72B
Anholocyclic
on Pinus sylvestris and Pinus mugo in western, northern and
central Europe, with an overwintering (hiemosistens) generation on the twigs
and two or more overlapping generations of progredientes attacking the
current year's shoots. Apterae are
dark brown or dark red, BL 1.0-1.2 mm, secreting copious wax-wool, in which
the eggs are laid (see
influentialpoints.com/Gallery). The second progrediens
generation in May-June includes alatae, but first instar crawlers dispersed by
wind may be a more common means of infection of young seedling pines (C.I.
Carter, pers. comm.). The morphs of P. pini are all virtually indistiguishable from the equivalent
secondary host generations of P.
orientalis, and it seems probable that the anholocyclic P. pini arose from the holocyclic P. orientalis by loss of a functional
sexual generation, as suggested by Marchal (1913), and supported by
mitochondrial and nuclear DNA sequence analysis (Havelka et al. 2019). P. pini in Europe seems to restrict
its attacks mainly to Pinus sylvestris
and P. mugo, but anholocyclic
populations introduced to the southern hemisphere can be injurious to a wide
range of pine species. In the Australian literature populations of P. pini group have commonly been given
the name laevis Maskell and
confused with P. boerneri, which
also occurs on a range of pines in both Australia and New Zealand. P. pini has also been recorded from
pine plantations in east Africa but these records should all be referred to P. boerneri (Blackman et al. 1995). 2n=20, 21 (Blackman et al. 1995) or 2n=22 (Steffan 1968). Pineus
pinifoliae (Fitch) (fig.
72A) Host-alternating
between Picea spp. (engelmanni, glauca, mariana, pungens, rubens, sitchensis) and
white pines (Pinus strobus, P. monticola) in North America. The galls on spruce are conical, 1.8-4.3 cm
long, with their constituent needles characteristically flattened and
broadened to form "scales".
They are purple and green at first, becoming reddish-brown at the time
of opening in mid-june to early July, when the needle-scales flare
outwards. The gallicolae, 1.6-2.3 mm,
fly to needles of the previous year's growth of white pine; they invariably
face the base of the needles and deposit their eggs beneath their wings,
dying after oviposition with the wings still protecting the eggs. The first instar sistentes (neosistentes)
move to the axes of new shoots to overwinter.
There are two apterous progrediens generations on pine, alate
sexuparae being produced in May to fly to old needles of spruce. For further life cycle details see Balch
& Underwood (1950). Dynamics of
the gall populations on spruce was studied by Howse & Dimond (1965), and
of the populations on white pine by Dimond & Allen (1974). The histology of feeding damage to white
pine shoots has also been studied (Allen & Dimond 1968). 2n=22 according
to Steffan (1968). Pineus
piniyunnanensis Zhang, Zhong & Zhang Apterae red with black head and prothorax,
presumably secreting wax in life; BL c. 0.8 mm. On Pinus yunnanensis in Yunnan, China (G. Zhang et al. 1992c). The life cycle is unknown. Pineus
sichuananus Zhang Described from Picea likiangensis and Picea
purpurea in Sichuan, China. The
first instar and adult sistens, which one would normally expect to occur on a
Pinus species, were recorded along
with galls and gallicolae on spruce, so the life cycle needs
clarification. P. sichuananus is considered to be closely related to P. cembrae; the differences given all
involve the wax glands (G. Zhang et al.
1980). Pineus
similis (Gillette)
Anholocyclic, living without host alternation on a wide range of Picea spp. in North America (abies, glauca, engelmanni, mariana, pungens, rubens), and
introduced into Britain where it also colonises Picea sitchensis (Carter 1975).
The unusual life cycle of this adelgid was investigated in Canada by
Cumming (1962a). Pseudofundatrices
(not true fundatrices, because they hatch from unfertilised eggs) overwinter
in the second or third instar under bud scales. Their feeding in spring causes rather loose
galls 1-4 cm long, with the constituent needles much paler than the normal
new growth (see bladmineerders.nl/gallen). As the gall matures the shoot and needles
become brown and twisted giving it a ragged appearance. Uniquely, both alate and apterous
gallicolae are produced in the galls.
Apterous gallicolae oviposit within the galls, but apterae may also be
found outside the galls, laying eggs in small tufts of wax on the stem. Alate gallicolae, 1.0-2.1 mm and dark
reddish-brown, fly in June-July to recolonise spruce needles, laying eggs in
white wax wool. These apparently
develop into the overwintering pseudofundatrix generation. 2n=22 according to Steffan (1968). Pineus
strobi (Hartig)
Anholocyclic on Pinus strobus
where this pine species is native in eastern North America, and introduced on
this pine into Europe. In England, Carter (1971) has recorded it attacking
one species of European white pine, Pinus
peuce, and in the BMNH collection there are also specimens
tentatively identified as this species from P. cembra, P. pinea and
P. sylvestris. It has also now been recorded from western
Siberia (Altai Republic) on Pinus
sibirica (Stekolshchikov & Novgorodova 2013). Recently it
is also reported from western North America (Darr et al. 2018). Raske & Hodson (1964) describe the life cycle
and biology in Minnesota, USA, and their observations largely agree with
those made in Europe (e.g. Marchal 1913, Carter 1971). Overlapping generations of sistentes and
progredientes feed through the spring and summer on shaded parts of the bark
of the trunk and undersides of branches, secreting copious white wax
wool. On seedling trees, parts of the
new growth or last year's growth wher the needles are densest are
colonised. Overwintering occurs as
immaure apterae (progrediens or sistens) on the trunk, or under scales of
dwarf-shoots. Alate sexuparae are often produced in May-June, and fly to Picea mariana to lay eggs on the
needles. However, the sexual phase is
abortive, either because the sexuales hatch but fail to develop (Raske &
Hodson 1964) or due to the absence of any males (Marchal 1913). No galls or gallicolae of P. strobi have been found on Picea mariana or any other
spruce. Sloss & Foote (1973)
studied predation by Leucopis pinicola
(Chamaemyiidae) and other natural enemies of P. strobi in Ohio, USA, and Wantuch et al. (2019) investigated its predator complex in Virginia,
USA. 2n=22 (Steffan 1968) or 20
(Blackman & Eastop 1994). Pineus
sylvestris Annand
Apparently anholocyclic in Pinus
spp. (radiata, sylvestris) in
California, USA (Annand 1928). Inouye
(1953) records what appears to be the same species from Pinus thunbergii in Hokkaido, Japan. Reddish-brown apterous progredientes,
1.0-1.1 mm with rather sparse wax, were found in May-June in California
feeding inconspicuously at the needle bases near male cones. Alate sexuparae appeared early in May, but
their fate is unknown, no generations on spruce having been identified. The overwintering stages on Pinus, mainly second and third instar
progredientes, were found under stem scales and behind lateral buds near the
ends of branches. Pineus
wallichiana Yaseen & Ghani Anholocyclic on Pinus griffithii (= wallichiana)
in the Galis and Swat Hills, Pakistan, attacking stems and twigs of mature
and old trees (Yaseen & Ghani 1971).
Adult sistentes are about 0.9-1.0 mm, light brown with a thick
covering of wax. There are only one or
two sistens generations per year, and no alatae are known. Morphology and life cycle are apparently
very like Pineus ghanii, and this
is possibly a highland form of that species.
One
species on Fagus in Japan. Only the very specialized apterous vivipara
is known, and its phylogenetic position is uncertain. Takahashi (1957) compared it with Dasyaphis, but Shaposhnikov &
Gabrid (1987) suggested that it belongs in the Hormaphidinae. Platyaphis
fagi
Takahashi (fig. 47A) Apterae are broadly ovate, flattened, green
with pale antennae and legs; BL 1.8-2.0 mm.
On undersides of leaves, along mid-ribs, on Fagus crenata in Japan (Takahashi 1957). Life cycle and other morphs are unknown.
About 60 species of mainly
pale green to yellow aphids bearing numerous short capitate hairs, and with
slender, often elongate appendages.
With only a few exceptions the species live on Compositae, mostly on
Anthemideae (29 spp., of which 24 are on
Artemisia and 4 on Achillea)
and Astereae (19 species, of which 15 are on Chrysothamnus or Ericameria),
with a few species on Inuleae (Gnaphalium,
Helichrysum). They are specialised feeders on plants with
dense glandular hairs. The genus is predominantly American (49 species), with
seven European and only three central Asiatic and one east Asian species
known. Many of the species were
originally described in Capitophorus.
Daniyarova & Narzikulov (1984) reviewed the genus, and regional accounts
are available for Europe (Holman 1965a, Leclant 1968a, Heie 1994, Blackman
2010) and America (Hille Ris Lambers 1969, Corpuz Raros & Cook 1974).
Kadyrbekov (2016) provided a key to the palaearctic species. Pleotrichophorus
achilleae Holman Apterae are greenish or pinkish white,
with dusky to dark appendages; BL 1.6-2.3 mm. On undersides of leaves of Achillea pectinata (= kitaibeliana) in eastern Europe
(Slovakia, Hungary). Monoecious holocyclic with oviparae and apterous males
in October (Holman 1965a). Pleotrichophorus ambrosiae Hille Ris Lambers
(Fig.8a) Apterae are pale green to whitish green
with dusky apices to antennae and legs, BL 1.8-2.3 mm. On undersides of leaves of Ambrosia spp., widely distributed in
USA , and in Mexico and Cuba (Holman 1974b). Alatae have no central dorsal
abdominal patch (cf. Capitophorus
spp.). Monoecious holocyclic in USA (e.g. Colorado), with alate males
(Corpuz-Raros & Cook 1974). Pleotrichophorus amsinckii Richards
Apterae are pale green, appearing “weakly pulverulent” due to dense
dorsal hairs; BL 1.6-1.8 mm. Described from Amsinckia intermedia in British Columbia, Canada and subsequently
recorded from Washington, USA on Eriogonum
sp., Phacelia hastata and Eriophyllum lanatum, which makes the true host relationship uncertain. Pleotrichophorus antennarius Corpuz-Raros & Cook Colour of apterae in life is unknown; BL
1.1-1.6 mm. On Antennaria
plantaginifolia and Antennaria sp.
in Washington D.C., USA. Monoecious holocyclic with oviparae and apterous
males in October (Corpuz-Raros & Cook 1974). Pleotrichophorus artemisicola (Williams)
Apterae are pale green with pulverulent appearance due to dense,
capitate hairs, with mainly dark antennae and legs, black siphunculi and
white cauda; BL 1.7-1.9 mm. On upper part of stem and leaves of Artemisia cana. This specie is noted
to be very active. Described from Nebraska, USA, and there are also records
from California, Oregon and Kansas. Pleotrichophorus asterifoliae (Strom)
Apterae are shiny dirty white to pale yellow with antennae and legs
distally dark; BL 1.8-2.4 mm. On Aster sp. in northern USA and Canada,
forming large colonies on undersides of lower leaves of plants growing in
shady areas. Oviparae were found in
autumn in Wisconsin, USA (original description). Pleotrichophorus brevinectarius (Gillette & Palmer) Apterae are mid-green, with frosted
appearance due to dorsal hairs; BL 1.3-2.0 mm. On leaves and stems of Artemisia longifolia, and and subsequently recorded from several other Artemisia spp. In
western USA. Pleotrichophorus
chrysanthemi (Theobald) Plate 16f Apterae are pale green to yellowish (see
aphids
of Karnataka website): BL 1.8-2.4 mm. On undersides of
leaves of florists’ chrysanthemums, (Dendranthema
spp.), often forming large colonies on yellowing leaves. Described from
South Africa, and since recorded from Europe, Egypt, Zimbabwe, Iran, India,
Nepal, Korea, Japan, Australia, North and South America. Capitophorus formosanus Takahashi, recorded from Japan and Taiwan
on Chrysanthemum, is possibly this
species. Apparently it is entirely
anholocyclic; no sexual morphs are known.
[P. chrysanthemi is
difficult to distinguish from P.
glandulosus and may not deserve separate species status. Alatae from Dendranthema generally have fewer
secondary rhinaria (9-20 on ANT III and none on IV) than those from Artemisia vulgaris, which have 14-23
on ANT III and 0-8 on IV (most with 3 or more on IV). Best discriminant for
apterae is perhaps the number of hairs on abdominal tergite 8; P. chrysanthemi has 5-11 (64% with
7-8), and P. glandulosus has 8-13
(65% with 9-10). Populations of this species group on Artemisia
apiacea in China (BMNH collection, leg. V.F. Eastop) resemble chrysanthemi rather than glandulosus, and Lee et al. (2002c) also identify
populations on Artemisia princeps var.
orientalis in Korea as chrysanthemi; it seems likely that chrysanthemi may have arisen quite recently in east Asia as an
anholocyclic form or clone able to feed on Dendrathema, and subsequently dispersed widely on florists’
chrysanthemums.] Pleotrichophorus
decampus (Knowlton & Smith)
(Fig.14k) Apterae are bluish green to apple-green
with “whitish cast” due to dorsal hairs; BL 1.3-2.0 mm. On Artemisia californica and A. tridentata in western USA and
Mexico. Records from other Artemisia spp. are mostly assignable
to P. pseudoglandulosus, with which there is much confusion in the
literature. Oviparae and alate males of P.
decampus occur on A. tridentata in
Utah in October (Corpuz-Raros & Cook 1974). Pleotrichophorus deviatus Müller
Apterae are dull whitish; BL 1.8-2.4 mm. On undersides of leaves of Hieracium spp. in continental Europe.
Oviparae and apterous males occur in September (original description). Pleotrichophorus diutius Corpuz-Raros & Cook Apterae are green, BL 1.6-2.2mm. On
leaves of Artemisia dracunculoides
and Artemisia sp. in USA (New Mexico, Colorado, Oregon, Minnesota). This
species is confused with P. wasatchii in
Palmer (1952); see Corpuz-Raros & Cook (1974). Pleotrichophorus duponti Hille Ris Lambers Apterae are dull greyish green with
green transverse stripes; BL 1.6-2.1 mm. On lower leaves of Achillea spp., falling when disturbed.
In western, northern, central and eastern Europe. Ossiannilsson (1959)
recorded it from Leucanthemum vulgare. Monoecious holocyclic on Achillea with oviparae and apterous
males in September-October (Hille Ris Lambers 1953). 2n=14. Pleotrichophorus elongatus (Knowlton)
(Fig.21f) Apterae are shining apple green, slenderly
spindle-shaped; BL 1.7-2.9 mm. On leaves and apical parts of Chrysothamnus
and Ericameria spp. in western
USA. As suggested by Palmer (1952) and Hille Ris Lambers (1969), P. palmerae (Knowlton) is probably a
synonym; the characters given to distinguish it from elongatus by Knowlton, and by Corpuz-Raros & Cook (1974), are
those likely to be found in alatiform specimens. Pleotrichophorus filaginis (Schouteden)
Apterae are purplish grey, BL 1.5-2.2 mm. Alatae have dark brown
abdominal pleural intersegmental sclerites. On Filago and Gnaphalium
spp., and also recorded from Helichrysum
arenarium. They feed solitarily or in small single-parent families on
upper- and undersides of the leaves.
Monoecious holocyclic, producing oviparae and apterous males in
October (Hille Ris Lambers 1953). Pleotrichophorus filifoliae (Palmer)
Apterae are small, mid-green , with frosted appearance due to dorsal
hairs; BL 1.0-1.3 mm. On leaves of Artemisia
filifolia in Colorado, USA. Recently it has been collected from Artemisia sp(p). in New Mexico, Utah,
Idaho and Oregon, including oviparae found on A. tripartita in October in Oregon (A. Jensen, aphidtrek.org). Pleotrichophorus glandulosus (Kaltenbach)
(Fig.14e) Apterae are yellowish white, sometimes greenish,
occasionally with a pale green median stripe; BL 1.4-2.6 mm. On undersides of
lower leaves of Artemisia vulgaris, and sometimes on other Artemisia spp. Records from other genera are probably of
vagrants, or based on misidentifications. In Europe and across Asia to Japan,
China, Korea and Taiwan, and introduced to eastern North America; records
from western North America are possibly all due to confusion with other
species (e.g. decampus, gnaphalodes,
pseudoglandulosus). Also in
Argentina (Ortego et al. 2004) and
Uruguay (BMNH collection, leg. V.F. Eastop); but alatae from A. verlotiorum in Uruguay resemble P. chrysanthemi (q.v.) in lacking
secondary rhinaria on ANT IV. Oviparae
and apterous males of P. glandulosus occur
on Artemisia vulgaris in September-October
(Heie 1994, Corpuz-Raros & Cook 1974). Nevsky’s (1929) subspecies in
Central Asia, P. glandulosus ssp. ispharinus, is probably just a
variant population. However another variant population with fundatrix,
apterae and oviparae all found in July on Tanacetum
bipinnatum in north-west Russia (Nenets region) and described as P. sp. aff. glandulosus by Stekolshchikov (2017) is possibly a distinct
species. 2n=14. Pleotrichophorus gnaphalodes (Palmer) (Fig.14f,h) Apterae are pale green with frosted
appearance; BL 1.3-2.2 mm. On undersides of leaves of Artemisia
douglasiana, A. dracunculus, A. ludoviciana and A. tridentata in western USA, and in Mexico on A.
mexicana (BMNH collection, leg. G. Remaudière). Pleotrichophorus gregarius (Knowlton)
(Fig.21b,e) Apterae are brown to greenish brown in
summer and autumn, with antennae black at apices of III and IV and dark on
V-VI, and siphunculi pale at bases but dark distally; BL 1.8-3.2 mm. Spring
generations are dark brown to chocolate brown. On growing points and leaves
of Chrysothamnus and Ericameria spp., with especially
large, dense colonies developing on rabbit brush (E. nauseosa), in western USA. Oviparae and alate males occur in
October-November (original description). Pleotrichophorus helichrysi Bozhko (Fig.31 l,n) Apterae are transparent white or pale
greenish, with dull greyish appearance due to the numerous fan-shaped dorsal
hairs; BL 1.5-2.1 mm. Living
well-camouflaged on undersides of leaves of Helichrysum arenarium. In
Denmark, Poland, Czech Republic, southern Russia and Kazakhstan (Bozhko
1976b, Heie 1994, Kadyrbekov 2017a). Pleotrichophorus heterohirsutus (Gillette & Palmer) (Fig.14g) Apterae are pale green with frosted
appearance due to dorsal hairs; BL 1.0-1.8 mm. On leaves and stems of Artemisia tridentata, forming crowded
colonies on young plants, in western USA (Colorado, Idaho, Utah, Wyoming).
Oviparae and apterous males occur in September (Palmer 1952). Pleotrichophorus hottesi Hille Ris Lambers Apterae are greyish green; BL 1.9-2.3 mm.
On Achillea lanulosa (= millefolium?) in North America (Colorado, North
Carolina, Manitoba). Monoecious holocyclic with apterous males in September
(original description). Pleotrichophorus infrequenus (Knowlton & Smith) (Fig.14a) Apterae are pale (colour unrecorded), BL
c.1.3-1.4 mm. On leaves of Artemisia tridentata in western USA
(Idaho, Utah, Oregon) and Mexico (BMNH collection, leg. G. Remaudière). Specimens
collected from Artemisia rigida (Oregon:
BMNH collection) have a rather larger cauda, and may represent a different
taxon. Pleotrichophorus knowltoni Corpuz-Raros & Cook Colour in life is unknown, probably pale
with frosted appearance; BL c. 2.5 mm. Described from an aptera and oviparae
collected in July on an unidentified host in Alaska, and subsequently found
on Artemisia spp. including A. tilesii in north-east Siberia
(Chukotka) by Stekolshchikov & Khruleva (2015), who provided detailed
descriptions of fundatrices and apterous males collected in July-August. This
species is very similar to P.
glandulosus, but with more numerous dorsal hairs. Pleotrichophorus lagacei Hille Ris Lambers Colour of apterae in life is unknown,
probably very pale with frosted appearance; BL 1.8-2.5 mm. On Ericameria bloomeri at an altitude of
2,600m in California, and also recorded from Oregon (A. Jensen, aphidtrek.org). Pleotrichophorus longinectarius (Gillette & Palmer) Apterae are pea-green, with frosted
appearance; BL 2.0-2.5 mm. On leaves and stems of Artemisia longifolia and Artemisia
sp. in western USA (Utah, Wyoming, New Mexico) and Canada (Alberta,
Manitoba). Pleotrichophorus longipes (Gillette & Palmer) (Fig.14b) Apterae are bluish apple green, and
apparently wax-dusted; BL 2.4-3.0 mm. Living singly and well-camouflaged on
leaves of Artemisia tridentata in
western USA (California, Colorado, Idaho, Utah, Wyoming). Oviparae and
apterous males occur in September (Palmer 1952). Pleotrichophorus longirostris Hille Ris Lambers Colour in life is unknown; BL 1.6-1.9 mm.
On Eriophyllum stoechadifolium in
California, USA, and also on Eriophyllum
sp(p). in Washington and Oregon (A. Jensen, aphidtrek.org). This species has
fan-shaped dorsal hairs and a long stiletto-shaped R IV+V (0.17-0.19 mm),
similar in length to the short thin siphunculi. Pleotrichophorus magnautensus (Knowlton & Smith) (Fig.21c) Apterae are green, with short black
siphunculi; BL 2.3-2.7 mm. On Chrysothamnus
viscidiflorus in Utah, Wyoming and Colorado, USA. Alatae are undescribed. Pleotrichophorus narzikulovi Umarov
Apterae are green, apparently dusted with grey wax; BL 2.5-3.0 mm.
Living in small colonies on Artemisia
dracunculus in Central Asia (Kyrgyzstan). Alatae are unknown. Pleotrichophorus obscuratus Hille Ris Lambers Apterae are greyish green, apparently
waxy due to dorsal hairs, with dark antennae, legs, apices of siphunculi,
cauda and anal plate, and sometimes conspicuously dark subgenital plate; BL
1.3-1.8 mm. Described from Artemisia californica in California, and subsequently collected on
several other Artemisia spp. in
Oregon, Utah, Nevada and New Mexico (A. Jensen, aphidtrek.org). Alatae are undescribed. Pleotrichophorus oestlundi (Knowlton)
(Fig.21h) Apterae are bluish green to apple green,
with apparent white pruinosity due to dense dorsal covering of fan-shaped
hairs; BL 1.5-2.3 mm. On Chrysothamnus and
Ericameria spp. in western USA.
Oviparae (red to brownish red in life) and alate males occur in October
(original description). Pleotrichophorus ohioensis Smith
Apterae are greenish with greyish white overcast due to fan-shaped
hairs; BL 2.1-2.4 mm. On undersides of leaves of Helianthus sp. in Ohio, USA. Oviparae and alate males were
collected in mid-October (original description). Pleotrichophorus packi (Knowlton)
(Fig.21g) Apterae are rather slenderly
spindle-shaped, bluish green to apple green or yellow, appearing rather shiny
or frosted due to fan-shaped dorsal hairs, with long dark siphunculi, curved
outward apically; BL 1.9-3.5 mm. On leaves of
Chrysothamnus and Ericameria spp. in western USA.
Oviparae and alate males occur in late October-November (Palmer 1952).
Corpuz-Raros & Cook (1974) described one sample with shorter siphunculi
and a more hairy cauda from E. nauseosa
as a subspecies, P. packi ssp. brevis, although from their description
this seems closer to P. sporadicus.
Pleotrichophorus parilis Corpuz-Raros & Cook Apterae are pale, colour unknown, BL
1.4-2.0 mm. On Artemisia dracunculoides
in Minnesota, USA, and recently found on Iva
axillaris in Oregon (A. Jensen, pers. com., 2016). Oviparae and alate
males occur in September (Corpuz-Raros & Cook 1974). Pleotrichophorus patonkus (Hottes & Frison) (Fig.5c) Apterae are apple green with darker green
spinal and lateral stripes, with hoary appearance due to the numerous
capitate dorsal hairs; BL c.1.6-1.9 mm.
On Achillea millefolium in
central USA (Illinois, Missouri, Tennessee, Kentucky), and apparently widely
distributed in North America, as it is also recorded from Achillea spp. in many western states and from Quebec, Canada (A. Jensen, aphidtrek.org
and http://www.flickr.com/photos/sandnine/5680760834/in/set-72157626451091245).
Aphids collected from Erigeron
philadelphicus in Illinois (BMNH collection, leg. V.F. Eastop) may also
be this species. Records prior to 1974 including the original description are
confused with P. pseudoplatonkus (q.v.);
the distinction between these two species still needs further confirmation. Hottes
& Frison (1931) described oviparae and alate males collected in
October-November, but these may have at least partly been P. pseudopatonkus. Pleotrichophorus patonkusellus Corpuz-Raros & Cook
Colour of apterae in life is unknown, probably pale green with hoary
appearance due to the dense capitate hairs; BL 1.1-1.6 mm. On Achillea
millefolium and Achillea sp. in Utah, USA (Corpuz-Raros & Cook
1974). Very similar to P. patonkus, apart from the denser and somewhat
shorter dorsal hairs, and possibly just a local variant of that species. Pleotrichophorus persimilis Börner
Apterae are pale yellowish red to brownish, appearing wax-powdered; BL
1.9-2.6 mm. On Artemisia spp., especially
A. campestris and A. scoparia, in Europe, and east to Iran, Kazakhstan and Pakistan. Oviparae
and apterous males occur on A.
campestris in September in Germany (Hille Ris Lambers 1953). Green aphids
with rather shorter appendages found in Afghanistan on A. turanica are regarded as a subspecies, A. persimilis ssp. afghanensis Narzikulov &
Umarov (Daniyarova & Narzikulov 1984), and aphids identified as
this subspecies have also now been recorded (but on other Artemisia spp.) from Kazakhstan
(Kadyrbekov 2016d, 2017a). Pleotrichophorus pseudoglandulosus (Palmer) (Fig.14j) Apterae are bluish green, with frosted
appearance due to dorsal hairs; BL 1.1-2.0 mm. They live singly on leaves and
stems of Artemisia spp. (douglasiana,
frigida, ludoviciana, vulgaris) in western USA, and in Canada (Alberta,
Manitoba, Saskatchewan). There has been considerable confusion with other
species in the literature; records of P.
glandulosus from native American Artemisia spp. in western USA are likely to be this
species, and most records from Artemisia
tridentata are probably referable to the very similar P. decampus. P. intermedius, described
from A. frigida in Minnesota (Corpuz-Raros & Cook 1974) is also
very similar and possibly a local variant. Oviparae and apterous males (cf. decampus)
occur in October-November in Colorado (Palmer 1952) and in September in
Manitoba (BMNH collection, leg. A.G. Robinson). Pleotrichophorus pseudopatonkus Corpuz-Raros & Cook Apterae are whitish green to green, BL
1.3-1.9 mm. On Achillea millefolium and
Achillea sp. in north, central and
eastern USA, western Canada (New Brunswick) and Mexico (BMNH collection, leg.
G. Remaudière). This species is very
similar to P. patonkus, but has a
shorter, less acute R IV+V and shorter siphunculi (Corpuz-Raros & Cook
1974). Oviparae and alate males are
possibly included in the original description of P. patonkus (Hottes & Frison 1931). Pleotrichophorus pullus (Gillette & Palmer) (= P. rusticatus (Knowlton & Smith)) (Fig.14d) Apterae are brown with appendages mainly
dark brown to black; BL 1.7-2.0 mm. On leaves of Artemisia spp. (cana, longifolia, tridentata) in western USA (Palmer 1952). P.
rusticatus (Knowlton & Smith) was regarded as a synonym by Hille Ris
Lambers (1969), but Corpuz-Raros &
Cook (1974) resurrected it as a good species on the basis of differences in
shape of the frontal hairs. After examining types of both, we think this is
probably only local variation. Pleotrichophorus pycnorhysus (Knowlton & Smith) (Fig.21a) Apterae are pale green to deep apple
green, somewhat shiny, with long, dark-tipped siphunculi; BL 1.6-2.3 mm. On Chrysothamnus
viscidiflorus and Gutierrezia sp.
in Utah, USA; also recorded from Idaho, Nevada, Oregon, Washington and
Montana, and from Ericameria nauseosa.
There is also a record from Artemisia
tridentata, but this is unlikely to be a true host plant. A fundatrix was
found in May on C. viscidiflorus by
A. Jensen (see photo at http://www.flickr.com/photos/sandnine/5692282948/in/set-72157626451091245). Pleotrichophorus quadritrichus (Knowlton & Smith) (Fig.14c,i) Apterae are bluish green, with greyish
cast due to dorsal hairs, and usually with dark siphunculi and cauda (see aphidtrek.org); BL 1.1-2.0 mm. On
Artemisia spp. in western USA. The
variation in this species was discussed by Corpuz-Raros & Cook (1974),
who erected subspecies P. quadritrichus ssp. vulgaris and P. quadtritrichus ssp. pallidus; the latter is a pale
form that is keyed separately and resembles P. gnaphalodes. The main
distinguishing features for the quadritrichus
group are the large, constricted cauda (a feature shared with P. gnaphalodes), and the often
strap-like apices of the frontal hairs.
Oviparae and alate males occur in October. Pleotrichophorus remaudierei Leclant
Apterae are pale green with a darker spinal stripe; BL 1.2-1.6 mm.
Living singly on undersides of leaves of Erigeron
acer in the French Alps. Monoecious holocyclic; fundatrices were
collected in April (Leclant 1968) . Pleotrichophorus spatulavillus (Knowlton & Smith) Apterae are pale green, with white
frosted appearance; BL 1.4-1.7 mm. On Artemisia
tridentata and Artemisia sp. in
Idaho and Utah, USA. Pleotrichophorus sporadicus (Knowlton)
Apterae are spindle-shaped, apple green, described as “heavily
pruinose above” (presumably an effect of the fan-shaped/spatulate dorsal
hairs), with mainly dark siphunculi; BL 2.1-4.1 mm. On leaves and stems
of Ericameria
nauseosa in western USA. Oviparae were found in October. This is one of a
complex of forms described from this host, and distinguished mainly by
differences in number and form of dorsal hairs, which are likely to be influenced
by environment and by degree of alatiformity. P. neosporadicus Corpuz-Raros
and P. packi ssp. brevis are both very close to P. sporadicus, and we could not find
reliable distinguishing characters for either of them. Pleotrichophorus stroudi (Knowlton)
Colour of apterae in life was not noted, presumably pale with frosted
appearance due to fan-shaped dorsal hairs, and with antennae and siphunculi
dark except at their bases; BL 2.0-2.5 mm. Hosts are Ericameria nauseosa and Chrysothamnus
viscidiflorus (A. Jensen, aphidtrek.org). In western USA. It belongs to a
taxonomically-difficult group of closely-related forms feeding on E. nauseosa (rabbitbrush) in western
USA (see packi, sporadicus). Pleotrichophorus tetradymiae Smith & Knowlton Apterae are “black to blackish green”
(?), with black siphunculi; BL 1.5-2.1 mm. On Tetradymia canescens in western USA (Idaho, Oregon). Pleotrichophorus triangulatus Corpuz-Raros & Cook Apterae are green; BL c.1.7 mm. On Agoseris sp. in western USA
(California, Colorado). Pleotrichophorus tuberculatus Stekolshchikov & Khruleva Apterae are dirty brown-yellow
with intensive wax pulverulence both dorsally and ventrally, and dark
siphunculi; BL 1.7-2.5 mm. On upper and undersides of apical leaves and on
shoot apices of Artemisia tilesii in
north-east Siberia (Chukotka). A fundatrix was collected in early July, and
oviparae and apterous males were present in early August (Stekolshchikov &
Khruleva 2015). Pleotrichophorus ugamicus Kadyrbekov
Apterae are whitish; BL 2.2-2.8 mm. On Artemisia santolinifolia in Kazakhstan (Kadyrbekov 2016a). A
member of the glandulosus group. Pleotrichophorus utensis (Pack & Knowlton) ( = Capitophorus
acanthovillus Knowlton & Smith)
(Fig.21d) Apterae are green with black siphunculi;
BL 1.4-1.9 mm. On Chrysothamnus
viscidiflorus and Gutierrezia spp.
in western USA (Utah, Nevada,Oregon). P.
acanthovillus has been distinguished from P. utensis occurring on the same hosts solely on the basis of
differences in shape of dorsal hairs in apterae, and in particular the
subapical caudal hair(s), which are said to be pointed in apterae of acanthovillus, and flattened apically
in utensis (Corpuz-Raros and Cook,
1974). This character is unreliable as it is related to the degree of
alatiformity, alatae having longer and less expanded hairs than apterae. Pleotrichophorus villosae Robinson
Apterae are very pale green, almost silvery (due to dense covering of
fan-shaped dorsal hairs); BL 1.6-2.1 mm. On Chrysopsis (= Heterotheca)
villosa in Manitoba, Canada. Alatae
and other morphs are undescribed. Pleotrichophorus wasatchii (Knowlton) Apterae are
greenish, with apparent pruinosity due to dense covering of fan-shaped hairs;
BL 1.6-2.1 mm. On Chrysothamnus and Ericameria spp. in western
USA. Records from other host plants and from eastern USA are probably all due
to confusion with other species; see Corpuz-Raros & Cook (1974). Pleotrichophorus xerozoous (Knowlton & Smith) Apterae are green, with antennae dark
distally, and siphunculi pale basally and dusky on distal half; BL 1.7-2.5
mm. Alatae are unknown. On Chrysothamnus
and Ericameria spp. in Idaho
and Utah, USA. This species is closely related to the much commoner P. gregarius, and is possible a
seasonal variant, differing in colour in life as well as in characters given
in key. Pleotrichophorus zoomontanus (Knowlton & Smith) Apterae are greyish green, BL 1.7-2.4 mm.
On Artemisia tridentata and Artemisia spp. in western USA. It
resembles P. longinectarius,
especially in having similarly protruding compound eyes, but has a shorter R
IV+V and differences in frontal and
dorsal hairs (Corpuz-Raros & Cook, 1974). Alatae are undescribed.
A
small genus with a rather evenly holarctic distribution, living without host
alternation on Salix. They are related to Pterocomma, but less hairy, and have flangeless siphunculi. Most species secrete flocculent wax, and
they are not attended by ants.
Richards (1966b) revised the genus (which in his concept included Fullawaya) and Wojciechowski (2003) reviewed
the palaearctic species. Tang et al.
(2015) investigated the use of DNA barcodes for species identification. Plocamaphis
amerinae (Hartig)
Apterae are yellowish, greenish or brownish, wax-covered in life, with
dark head, antennae and legs; BL 3.0-4.5 mm.
On young growth of Salix
spp. (alba, purpurea, viminalis),
often in large colonies and sometimes causing damage in osier
plantations. In north-west and north
Europe, west Siberia, and reported also from north-east Siberia (Chukotka;
Stekolshchikov & Khruleva 2015) and Kazakhstan (Kadyrbekov 2017a). Possibly also in Canada, if P. martini Richards (q.v.) is this species. Apterous males
and oviparae occur in October-November.
Populations with very short siphunculi and no dorsal dark markings on
a Salix species (probably lapponum) in Sweden were described as
a subspecies (P. amerinae borealis)
by Ossiannilsson (1959); such forms are also reported from S. viminalis in Poland (Szelegiewicz
1965). Plocamaphis
coreana (Okamoto & Takahashi) (fig. 106C) Apterae are dark yellowish green, covered
with white wax; BL c. 3.5 mm.
Described attacking branch and stem of an unidentified Salix sp. in Korea (Okamoto &
Takahashi 1927; Paik 1965, as Clavigerus
salijaponicus). Pashchenko (1988b)
recorded it from S. crassijulis and
S. parallelinervis in east Russia
(Kamchatca). The life cycle is
unknown. Plocamaphis
flocculosa (Weed) (figs 106E,F) Apterae are grey or yellowish grey,
densely covered with wax powder; BL 3.1-5.0 mm. Head, legs and antennae are darker, and
siphunculi are orange. On trunk and
branches of Salix spp. in North
America (discolor, lasiolepis) and Europe (borealis, caprea, cinerea, nigricans, repens). Also recorded from Iran (Rezwani et al. 1994) and China. Three subspecies have been distinguished in
the very variable European populations (Ossiannilsson 1959), but further work
is required to determine whether these are valid taxonomic categories (Heie
1986). The European form known as P. flocculosa ssp. brachysiphon produces oviparae and
alate males in October (Stroyan 1955; as P.
goernitzi). Kanturski et al. (2016b) described the fundatrix
from Poland. P. assetacea Zhang,
described from Salix sp. in Tibet
(Zhang & Zhong 1981b), may be a synonym (Tang et al. 2015). Plocamaphis
gyirongensis Zhang
Apterae and alatae were described from Tibet (Zhang & Zhong 1981b)
as a new genus (Radisectaphis), but
were transferred to Plocamaphis by
Remaudière & Remaudière (1997). The recorded host plant is Malus; presumably this is either an
error, or the aphids were vagrants. It
is very similar to P. flocculosa,
and Wojciechowski (2003) suggested that is should be regarded as a subspecies
of that species. Plocamaphis
martini Richards (fig. 106D) Apterae light brown to pink (no wax
mentioned); BL 2.8-3.5 mm. On roots of
Salix sp(p). in arctic and
subarctic Canada (Baffin Island, Quebec). Apterous males and oviparae occur
in mid-August (Richards 1963).
Possibly conspecific with P.
amerinae.
Two east Asian species on Illiciaceae,
with numerous dorsal hairs and thick, tapering siphunculi resembling Indiaphis, but first tarsal segments have two hairs displaced from apex,
the tergum is membranous and the eyes have distinct ocular tubercles. Alatae
have dark-bordered wing veins and secondary rhinaria on ANT III-IV or III-V. Polytrichaphis
fragilis Miyazaki Apterae are yellow to pale-yellowish
green with dark brown siphunculi and a pale cauda; BL c.1.6-1.8 mm. On undersides
of leaves of Illicium anisatum in
Japan (Miyazaki 1971). Polytrichaphis
illicius Qiao & Zhang Apterae are black; BL c.1.6-1.8 mm. On Illicium verum in China (Qiao & Zhang 2001d).
About 45 species ranging in
body size from rather small to very large, usually lacking siphunculi and
with well-developed wax glands, even in fundatrices. Their host plant associations are unusual,
in that they have apparently transferred their sexual generation in
relatively recent times from Populus
to various more advanced "primary" hosts, e.g. in Rosaceae,
Caprifoliaceae and Oleaceae, while mostly retaining conifer roots as
secondary hosts for the parthenogenetic generations. The species are fairly evenly distributed
through the northern hemisphere; about 20 in North America, 7 in Europe, 3 in
central Asia and 15 in east Asia. The
life cycles of many species are unknown.
Accounts are available for north-west Europe (Heie 1980b), the UK (Blackman et al. 2019b), Spain
and Portugal (Nieto Nafría et al.
2002a), Central Asia (Narzikulov 1964), India (A.K. Ghosh 1984), Japan
(Inouye 1956), China (Tao 1970, G. Zhang et
al. 1993c, G. Zhang et al.
1999c) and North America (Smith 1974a).
Our concept of Prociphilus
includes Stagona, Paraprociphilus (with migration from Acer to Alnus) and we also include as subgenera Pulvius, Neoparacletus
and Meliarhizophagus, each with a
single species, which all appear to be relatively recent biological
deviations from the main evolutionary lineage of Prociphilus. Prociphilus
americanus (Walker)
Host-alternating between Fraxinus
spp. and roots of Abies (balsamea,
grandis, procera) in North
America. The type specimen is an alate
sexupara from Nova Scotia (Doncaster 1961); prior to Doncaster's
redescription this aphid was called P.
fraxinidipetalae and/or P.
venafuscus. Early records from
roots of Pseudotsuga are probably
in error (Smith 1969), and records from other trees (eg. Aesculus) are probably of vagrant individuals. Fundatrices on ash are dark brown to black,
globose, forming early spring colonies on in curled terminal leaves of new
growth (Essig 1911) or on twigs (Patch 1909); populations may also develop on
suckers and at base of trunk "at quite a depth underground"
(Essig). Emigrant alatae maturing from late April to early June are very dark
green to black with dusky, bluish-tinged wings and long bluish white wax
flocks extending posteriorly; BL 2.8-4.8 mm.
They found wax-covered colonies on Abies
roots; a redescription of the apterous exule (BL 1.4-2.3 mm) was provided by
Pike et al. (2012). Sexuparae return to produce dwarf sexual
morphs on trunk of Fraxinus in late
August-October (Smith 1969). Anholocyclic overwintering may also occur on Abies roots (Zak 1965). Patch (1909)
described the sexual morphs (as venafuscus)
and recorded that they were also deposited in large numbers on Syringa, which may be an alternative
primary host. This is probably the aphid referred to as “Prociphilus californicus” in some western North American
publications (see Halbert & Blackman 2004). Prociphilus
aurus Zhang & Qiao
Only the fundatrix is described, and its appearance in life is
unknown; BL c. 2.9 mm. On Malus
sieversii in Xinjiang, China (G. Zhang & Qiao 1997c). Prociphilus
(Paraprociphilus) baicalensis
(Cholodkovsky) Apterae brownish,
secreting copious white wax wool: BL c. 3.1 mm. On undersides of leaves and shoots of Alnus spp., across Asia from the Urals
to Japan. Apparently entirely
anholocyclic on Alnus, although
sexuparae are produced in autumn (Mordvilko 1935). 2n=12*. Prociphilus
bumeliae (Schrank)
Host-alternating between Oleaceae (usually Fraxinus excelsior, sometimes Ligustrum
vulgare, Syringa vulgaris) and
roots of Abies. Fundatrices are brown, covered with white
wax-wool, forming loose "leaf-nests" on primary host in
spring. Spring colonies also form on
bark of stems and branches (see bladmineerders.nl). All their progeny are alate, with blackish
brown head and thorax spotted with white wax, and light brown to yellowish
red abdomen (Heie 1980b); BL 3.8-5.5 mm.
They migrate in May-June to found wax-covered colonies on Abies roots. (There is a possibility
that they also live attended by ants on roots of Picea abies –see under P.
fraxini). Sexuparae return to Fraxinus
in late August-September (Mordvilko 1935).
Widely distributed in north-west, north, central and eastern
Europe. Also recorded from east Asia,
but its presence there needs confirmation as most of these records should
probably be referred to P. oriens. [Alatae of Prociphilus "sp. B" on Fraxinus floribunda in northern India (A.K. Ghosh et al. 1970) key to bumeliae, except that they lack
discernible dorsal cephalic wax glands and have many more secondary rhinaria
(45-48, compared with 16-24) on ANT III.] Prociphilus
carolinensis Smith Apterae are oval, presumably wax-covered
in life, with protruding dark cauda and anal plate; BL c. 2.5 mm. They live on the roots of Pinus spp. (including P. taeda), 3-4 feet below ground,
attended by ants. Sexuparae (BL
2.8-3.3 mm) emerge in October-December, migrating to an unknown primary
host. Sexuales are deposited and eggs
laid on the bark of various trees, especially Malus, but apparently do not hatch (Smith 1969). In USA (North Carolina) and Canada
(Manitoba). Prociphilus
caryae caryae (Fitch) (=alnifoliae Williams)
Spring populations (fundatrices + immature alatae) cup or curl the
leaves of Amelanchier spp. and turn
them yellow or whitish. (Records from Crataegus are probably due to
misidentification, particularly confusion with P. corrugatans.) Emigrant
alatae (BL 2.7-3.3 mm) fly in June to colonise roots of Pinus sp(p), sexuparae returning to Amelanchier in September-October.
In western USA and Canada (British Columbia). (No field observations have been made of
these western populations on pine roots, but Smith (1969) obtained colonies
readily on Pinus strobus in the
laboratory, and sexuparae collected on P.
monticola in British Columbia (BMNH collection, leg G. Shrimpton) appear to be this
species.) Similar aphids in eastern
USA are regarded as a subspecies:- Prociphilus
caryae ssp. fitchii Baker &
Davidson Spring populations
(fundatrix + immature alatae) cause cupping and curling of leaves of Malus spp. (domestica, angustifolia)
and Amelanchier spp. (see influentialpoints.com/Gallery).
Emigrant alatae (BL 1.8-3.3 mm) migrate in June to found ant-attended
colonies on roots of Pinus strobus. Sexuparae return to the primary hosts in
October-November where they deposit sexuales in bark crevices near base of
tree or in nearby leaf litter (Smith 1969).
Widely distributed in eastern North America. Parthenogenetic
populations also persist throughout the year on white pine roots. A closely-related form, P.
caryae ssp. arbutifoliae,
forms spring populations on Photinia (=
Heteromeles) arbutifolia in California, from which alatae (BL 2.9-4.1 mm)
migrate to an unknown secondary host, probably pine roots. [Records of P. caryae fitchii on Lonicera
in India (A.K. Ghosh et al. 1981,
A.K. Ghosh 1984b) apply to P.
himalayensis.] Prociphilus chaenomelis
Lee,
Seo & Hwang Apterae
(fundatrices) are undescribed, alatae are dark brown producing copious wax;
BL 2.1-2.8 mm. In rolled leaves of Chaenomeles
sinensis in Korea. Presumably heteroecious holocyclic, migrating to an
unknown secondary host. Prociphilus clerodendri Okamoto & Takahashi Apterae are not described. Alatae have head, eyes, antennae and
mesothorax black, legs black and
reddish brown, and abdomen yellowish green with a reddish tinge,
secreting white cottony wax; BL c. 4.5 mm. On Clerodendron trichotomum in Korea. Prociphilus
cornifoliae Singh, Das & Raychaudhuri Apterae (BL 4.2 mm, colour in life unrecorded)
were collected on roots of Cornus
sp. in April in Manipur, India (Singh et al. 1977; habitat information from
A.K. Ghosh 1984b). Alate specimens and
immatures were also collected, both from the same host, locality and date,
and from an unidentified plant in Nepal.
Life cycle is unknown. Prociphilus
(Neoparacletus) corrugatans (Sirrine) Spring populations cause curling and
cupping of leaves of Crataegus
spp.; infested leaves turn pale whitish, often with a reddish tinge at the
edge. Fundatrices are yellowish green
with flocculent wax, with greenish immatures that develop into emigrant
alatae (BL 1.9-2.9 mm, with brownish abdomen). Migration occurs in May-June to roots of
Cyperaceae, where the apterous exules are milky white, sometimes tinged with pale brown,
entirely covered with wax powder; BL 2.9-3.8 mm. Large colonies are formed on
Juncus
spp. and populations may persist throughout the year. It is also recorded
from Carex (Strom 1942, as Neoparacletus caricis). Sexuparae returning to Crataegus in autumn often deposit
sexuales on Malus, but apparently
these do not produce spring populations (Smith 1972a). Records from Amelanchier are possibly all due to confusion with P. caryae. Widely distributed in North America. [Dark brown or reddish curled leaves on Crataegus tomentosa, noted in the
original description of P. corrugatans
(Sirrine 1894), are considered by Smith (1972a) to be due to feeding by
another Prociphilus sp., possibly
the undescribed primary host generations of P. erigeronensis.] Prociphilus
crataegicola Shinji Described from Crataegus sp. in June in Japan; alata has light greenish abdomen,
and antennae with secondary rhinaria distributed III 22, IV 7, V 5 (Shinji
1922a). Other morphs and life cycle
unknown, except that alatae collected from pine roots in Taiwan and Japan by
Tao (1970), as "Prociphilus pini
n. sp." could be the sexuparae of this species. Possibly crataegicola is a synonym of pini
Burmeister; alatae (BL 3.2 mm) collected on "apple" in Japan and
identified by Takahashi as crataegicola
(BMNH collection, leg. T. Tanaka) have secondary rhinaria distributed III
24-29, IV 6-8, V 6-8, and closely resemble European P. pini. [P. sasakii Monzen, described from Malus domestica was synonymised with crataegicola by Shinji (1941), but
this seems doubtful as Monzen's species had siphunculi (unusual in Prociphilus), and less rhinaria on
III.] Prociphilus
(Stagona) crataegistrobi Smith Fundatrices and their progeny in spring
curl leaves of Crataegus sp., which
remain green but may tend to fade.
Their progeny are all alate, BL 2.2-2.5 mm, migrating in June to found
colonies on roots of Pinus strobus. The apterous exules live in indvidual
wax-lined cells associated with small roots and mycorrhizae (Smith
1969). Sexuparae return to trunks of Crataegus in October-December, but
pine root-feeding populations also persist through winter. In USA; only known from North Carolina. Prociphilus (Stagona) dilonicerae Zhang Apterae (fundatrices) are undescribed; BL
of alata (fundatrigenia) is c.3.6 mm. On Lonicera
sp. in Tibet. Secondary host is unknown. Prociphilus
emeiensis Zhang
Described from two alatae (BL c. 4.0 mm) collected in April on Fraxinus chinensis in Sichuan, China
(G. Zhang & Qiao 1997c). Appearance
in life and biology are unknown. Prociphilus erigeronensis (Thomas) Plate 2h Apterae (exules) are white to pale yellow
with darker head, antennae, legs, and end of abdomen, secreting white wax; BL
1.7-2.3 mm. In ant-attended colonies on roots of many plants, especially
Compositae (Ambrosia, Aster, Cichorium, Erigeron, Lactuca, Solidago, Tragopogon),
Poaceae (Agrostis, Muhlenbergia, Poa, Triticum) and
Leguminosae (Arachis, Phaseolus, Trifolium). Widely distributed in USA, and also recorded from
Ontario, Canada, and from Puerto Rico. Predominantly anholocyclic; Smith
(1974) redescribed this species, including alate sexuparae, and discussed a
possible partial holocycle with Crataegus
(or Amelanchier) as primary
host. The name is possibly being applied to more than one species. Prociphilus
formosanus Takahashi
Dense spring colonies of this species distort the leaves of Fraxinus sp. ("insularis")
in Taiwan (Takahashi 1935). Alatae
(collected in May) have a dark green abdomen, and BL c. 4 mm; antennae with
secondary rhinaria distributed III 32-39, IV 7-11, V 7-9, VI 0. Immatures yellow with cottony wax on
abdomen. Other morphs are
unknown. Alatae were also collected on
Picea morrisonicola, which is
therefore thought to be the secondary host, but confirmation of this seems
necessary. Prociphilus
fraxini (Fabricius)
Host-alternating between Fraxinus
excelsior and roots of Abies,
and probably also Picea (see
below). Fundatrices feed at bases of
new shoots of ash, giving rise to large colonies of immature alatae which
develop to maturity in dense leaf-nests, often high up in trees (Heie
1980b). Alatae (BL 3.3-5.1 mm) fly in
late June-July to found wax-covered colonies on Abies roots, the apterous exules being almost indistinguishable
from those of P. bumeliae. Sexuparae return to Fraxinus in October-November.
The male genitalia were described and
illustrated by Wieczorek et al.
(2012). [Immature apterous exules identified as P. fraxini have now been observed attended by
ants (Lasius brunneus and Prenolepis nitens) at bases of trunks of Picea abies, and these may have overwintered parthenogenetically deep underground on
spruce roots in symbiotic association with specialised ants such as P. nitens (Purkart et al. 2019).] P.
fraxini is widespread in Europe, and also recorded from Iran; specimens
from Turkey on Fraxinus oxycarpa
(BMNH collection, leg. H. Canakçioglu) are possibly an undescribed,
closely-related species. Also recorded from China (G. Zhang et al. 1993c). Prociphilus
(Meliarhizophagus) fraxinifolii (Riley) Living all-year-round on American Fraxinus spp. Yellow-green to pale green fundatrices and
their progeny (both apterae and alatae) live in curled and clumped leaves at
tips of twigs throughout the growing season (see influentialpoints.com/Gallery).
Alatae (BL 1.4-2.8 mm, with yellow-green abdomen) are produced in April-October. Ant-attended colonies also occur
all-year-round on Fraxinus
roots. Alate sexuparae occur in
October-November (Smith 1974a). Common
and widely distributed in USA; also in Canada (British Columbia, Manitoba)
and Mexico, and introduced to Chile (Carrillo 1977), South Africa (on planted
F. pennsylvanica and F. velutina; Müller & Schöll 1958,
and BMNH collection); also to Europe, where a colony was found on a planted F. pennsylvanica in Hungary
(Remaudière & Ripka 2003), and it has subsequently spread to Serbia
(Petrović-Obradović et al.
2007), Bulgaria (Trenchev & Trencheva 2009), Spain (Pérez Hidalgo &
Mier Durante 2012), UK (Baker & Martin 2011), Poland (Halaj et al. 2016), Iran (Tajmiri et al. 2016) and Kazakhstan
(Kadyrbekov 2017a). In 2015 it was found for the first time on the native
European ash F. excelsior at four
localities in Poland (Halaj & Osiadacz 2017). It has also recently been
reported from Beijing, China (Yu et al.
2015). Carver (1980) described an
aphelinid parasitoid on P. fraxinifolii
in Iowa, and Brudrett & Kendrick (1987) studied symbiosis between
root-feeding populations and a bolete fungus.
2n=20 (Robinson & Chen 1969) or 2n=22 (Blackman & Eastop
1994). Prociphilus
gambosae Zhang & Zhang
Apterae are greyish green, secreting long wax filaments; BL c. 2.4 mm.
On Syzygium caryophyllus (= aromaticum)
in Hebei, China (G. Zhang et al.
1993c). Other morphs, biology and life cycle are unknown. Prociphilus
(?Neoparacletus) ghanii Hille Ris Lambers Apterae are broadly oval, appearance in
life unknown; BL 1.3-1.7 mm. Collected
on Salix wallichiana in Pakistan
(Hille Ris Lambers 1973b). Other
morphs, biology and life cycle are unknown. Prociphilus
himalayaensis Chakrabarti Described from sexuparae (BL 2.6-3.4 mm,
wax-covered) collected on roots of Pinus
excelsa (=griffithii) in
Himachal Pradesh, India (Chakrabarti 1976a).
This aphid was originally placed in subgenus Stagona, although the pterostigma is not elongated, perhaps
because the apterous exules described as paratypes (from an unidentified
host) have a tibial chaetotaxy like that of P. (S.) crataegistrobi, but there is no real evidence that these
are the same species as the sexuparae from pine roots. Banerjee &
Chakrabarti (1993) confirmed experimentally the suspected (Chakrabarti et al. 1985) migration from
rolled-leaf galls on Lonicera
quinquelocularis. The spring forms on Lonicera
had earlier been recorded as P. caryae
fitchii by A.K. Ghosh (1984b). The apterae (fundatrices) are pale brown; BL
2.2-3.3 mm. All progeny are alate (BL 2.3-3.0 mm), migrating in May-June to
roots of P. excelsa. Alate
sexuparae (BL 2.6-3.4 mm) return to Lonicera
in November-December (Banerjee & Chakrabarti 1993). Samples
have been seen from the Punjab, Himachal Pradesh, Kashmir and Pakistan. Rather
similar spring migrants but with more rhinaria were collected from Eriobotrya petiolata in West Bengal (BMNH collection, leg. A.N.
Basu), so there remains the possibility that two species with different
primary hosts are being confused. There are apterae of other undescribed
species on roots of Pinus griffithii
in the Himalayas, and the group merits further investigation. P. trinus Zhang,
collected from L. japonica in
China, is probably a synonym on the basis of the published description. The distinction
between P. himalayensis and the Central Asian species P. umarovi is also unclear. Prociphilus
(Stagona) konoi Hori
Host-alternating between Lonicera
spp. and Picea. Fundatrices are yellowish green, BL 2.5-3.0 mm,
curling terminal leaves of Lonicera
longitudinally. All their progeny are
alate (BL 2.5-2.8 mm, with greenish brown abdomen) and migrate in June-July
to found colonies on roots of Picea
glehnii (Hori 1938a; requires experimental confirmation). Sexuparae (with secondary rhinaria distributed
ANT III 15-16, IV 8-9, V 7-8, VI 3-4) return to Lonicera in October-November.
In Japan and east Siberia (Pashchenko 1988a). 2n=18. Prociphilus
kuwanai Monzen (=
orientalis Mordvilko 1929) Feeding by the fundatrix in spring on Pyrus spp. (and occasionally Malus) causes leaf to curl downwards
forming a bag-like gall, coloured yellowish green tinged with red. Alatae (BL 2.6-2.9 mm, with a yellowish
green abdomen) fly in late June-early July.
The secondary host is uncertain; roots of Larix according to Shinji (1932; possibly due to confusion with
another species), or more probably of Rumex
spp. (Mordvilko 1935, Higuchi & Miyazaki 1969). Hori (1938a) also noted that the secondary
host "may probably be Rumex". Sexuparae occur in October-November, but
have not been described. In Japan,
Korea and east Siberia. Prociphilus
(?Stagona) laricis Shinji Only alate sexuparae are described, BL c.
1.6 mm, found swarming out of the ground along the roots and trunk of Larix kaempferi in Japan in late
October-early November (Shinii 1941).
The primary host is unknown. Prociphilus
ligustrifoliae (Tseng & Tao) Spring populations occur in curled leaves
of Ligustrum japonicum in China
(Tseng & Tao 1938), and are also recorded from L. lucidum in Japan (Wieczorek et al. 2012). The globular
yellowish white fundatrices, covered in white cottony wax, develop on the
buds in March-April and cause economic damage to trees used in rearing the
coccid Ericerus pela for commercial
wax production in Sichuan (Zhang 1981).
The alatae produced in the leaf-nests, have BL c. 3.75 mm,
dark-bordered wing veins, a pale green abdomen, and secondary rhinaria
distributed III 32-36, IV 13-16, V 12-16, VI (0-)1-7. They presumably fly to an unknown secondary
host. This species is closely related
to, and possibly synonymous with, P.
osmanthae. Prociphilus longianus Smith
The appearance of the apterae (exules) in life is unrecorded,
presumably secreting white wax, with conspicuous dark sclerotic tubular
extension of anal plate around cauda; BL 2.7-3.3 mm. The host is uncertain;
it was recorded from a “gall on Conophilus
americanus” (presumably Conopholis americana), also from base
of stem of a “garden geranium”, roots of an unidentified grass, and from
ants’ nests. Prociphilus (Stagona?) lonicerae Shinji Apterae (fundatrices) are green; BL not
recorded. Rolling young leaves of Lonicera japonica in Japan. All
progeny are alate (BL c.2.1 mm), migrating in early June to an unknown
secondary host. Prociphilus
(Paraprociphilus) mexicanus Remaudière & Munoz
Viveros Apterae are dark-coloured,
entirely covered with a dense waxy secretion, with long wax filaments
projecting posteriorly; BL 2.9-3.6 mm.
On undersides of leaves of Alnus
firmifolia in Mexico (Remaudière & Munoz Viveros 1986). Sexuparae are produced in autumn, and there
is at least a partial migration to Acer
negundo, from which fundatrices and emigrant alatae were described by
Remaudière & Munoz Viveros (1992b). Prociphilus
micheliae Hille Ris Lambers Only spring migrant alatae are described;
appearance in life unknown, BL 3.3-4.0 mm, secondary rhinaria distributed III
28-35, IV 8-13, V 0, VI 0. In curled
leaves of Michelia (= Magnolia) champaca in northern India (Kashmir, West Bengal, Uttar
Pradesh), becoming mature in April-June (BMNH collection). Other morphs amd
life cycle are unknown. 2n=14 (Kar et al. 1990). Prociphilus
oleae Leach ex Risso
Fundatrices (BL c.3.2 mm) and their progeny form compact spring
colonies covered in white wax wool on shoots near base of trunk of Olea europea. It is also recorded from Phillyrea latifolia Unusually for a Prociphilus, no leaf deformation is caused. Progeny of the fundatrix are all alate, BL
3.7-4.0 mm with smoky yellowish abdomen, migrating in May. The life cycle is unknown, but apterae
described from the roots of Vitis
vinifera in Sicily (Barbagallo & Stroyan 1982) could be the secondary
host form of this species. In
Mediterranean area (France, Italy, Greece, Turkey). Roberti & Monaco (1987) gave an account
of this aphid in Italy. Prociphilus
oriens Mordvilko
Fundatrices and their progeny feed on young growth of primary hosts in
spring causing curling and clustering of leaves into leaf-nest galls (see http://www.agr.hokudai.ac.jp/systent/akimoto/gall/P.%20oriens.jpg). Typical primary hosts are Fraxinus spp., but there are records
of spring populations on a remarkable range of other plants, not only in
Oleaceae (Ligustrum, Lonicera, Syringa) but including Rosaceae (Malus, Prunus, Sorbus) as well as Salix, Tilia and Ulmus (Kono
1940). The progeny of the fundatrix
are all alatae, BL 3.5-5.5 mm, with yellowish brown to dull reddish green
abdomen covered with white wax; migrating in June-early July to roots of Abies sachalinensis (incl. var. mayriana). Records from Picea roots need confirmation.
Return migration of sexuparae occurs in September-October (Hori
1938a). Recorded from India, China, east
Siberia, Korea and Japan (type locality).
Yamaguchi (1985) and Li & Akimoto (2017) studied sex ratio
variation in this species, and Akimoto (2006) investigated the effects of
in-breeding. Prociphilus
osmanthae Essig & Kuwana Apterae in curled-leaf pseudogalls on Osmanthus spp. in spring are greyish
green, wax-covered (see influentialpoints.com/Gallery);
BL c. 1.8 mm. Emigrant alatae have BL 3.3-4.0 mm, wax-covered olive-green
abdomen, and dark-bordered forewing veins (Essig & Kuwana 1918, Moritsu
1983). Secondary rhinaria of paratypes
are distributed III 27-31, IV 9-12, V 8-12, VI 0. They migrate in late May to an unknown
secondary host. In Japan, and reported
to occur in Meghalaya, India (A.K. Ghosh & Raychaudhuri 1968c, on an
unidentified host, and Khuda-Bukhsh & Kar 1990, on Croton sp.). It has recently been introduced to Hungary (Ripka et al. 2020). A.K. Ghosh (1984b)
included various Japanese specimens collected by Takahashi in his
redescription of osmanthae;
emigrant alatae from Ligustrum,
apterous exules from Michelia, and
alate sexuparae (with an anocaudal process) from Osmanthus. It is not
certain that all these forms are the same species. P.
ligustrifoliae also has dark-bordered wing veins and may be a synonym,
although alatae from galls have more rhinaria, including some on ANT VI. 2n=18 (for aphids collected on Croton; Khuda-Bukhsh & Kar 1990). Prociphilus
pergandei Smith
Fundatrices curl leaflets of Fraxinus
americana in eastern USA. Progeny
are all alate, BL 2.7-3.2 mm, covered with long flocculent wax secretion,
flying in June to an unknown secondary host (Smith 1974a). Prociphilus
(Stagona) piceaerubensis Smith Apterae are white to pale yellowish with a
tuft of wax on abdomen; BL 0.9-1.2 mm.
On roots of Picea rubens,
inhabiting individual wax-lined cells associated with mycorrhizae, especially
in humus or where roots have penetrated decaying logs. Sexuparae occur in
September-October. Described from
North Carolina, USA (Smith 1969); sexuparae which may be this species have
also been trapped in Germany and England (BMNH collection), and it is perhaps
the form described as xylostei
"forma minima" by
Tüllgren (1909). Prociphilus
(Stagona) pini (Burmeister) Plate 6a, b Fundatrices and their progeny curl and
yellow the leaves of Crataegus spp.
in spring. Progeny are all alate, BL 1.7-3.1
mm, with light green to greyish green abdomen, flying in June-August to found
colonies of small, wax-covered apterae on fine roots of Pinus sylvestris and its close relatives; also recorded on P. contorta in UK (Carter &
Danielsson 1993) and from P. sibirica
in Kazakhstan (Kadyrbekov 2017a).
Sexuparae (BL 1.6-2.3 mm; (see influentialpoints.com/Gallery)
return to Crataegus in
October-November, but populations also persist throughout the year on Pinus roots. Tambs-Lyche & Heie (1994) recorded
(vagrant?) P. pini from Malus domestica. In north-west and central Europe, and
Central Asia (Kazakhstan; Kadyrbekov 2017a).
Smith (1969) gave distinguishing characters from the closely-related
North American species crataegistrobi. 2n=16. Prociphilus
piniradicivorus Smith Apparently living all-year-round on Pinus spp. (Smith 1969). Apterae (presumably wax-covered in life, BL
c. 2 mm, with protruding dark cauda and anal plate) live on roots of Pinus spp., attended by ants and often
in association with the very similar but host-alternating species P. carolinensis. Sexuparae (BL 2.9-3.3 mm) are produced in
October-November and fly only a short distance, depositing sexual morphs on
trunks of nearby pine trees. Colonies
were experimentally reared from fundatrices that hatched from eggs
transferred to seedling P. taeda
(Smith 1969). In USA (North Carolina,
Maryland, Indiana). Prociphilus
(Pulvius) probosceus (Sanborn) Living all-year-round on Fraxinus. Fundatrices and their developing alate
progeny curl and distort leaves and terminal growth of Fraxinus spp. in spring.
More commonly found are the very large apterae (BL 4.5-5.3 mm), in ant
shelters at bases of water shoots and trunks of young trees, at or below soil
level (Smith & Stroyan 1972).
Sexuparae are produced in October-November, but underground colonies
persist throughout the year. In
eastern North America. Prociphilus
sasakii Monzen
Described from Malus domestica
in Japan (Monzen 1927, 1929). Large,
globular fundatrices (BL c. 3.3 mm) and their developing alate progeny,
covered with white woolly wax, cause apple leaves to become curled, bloated
and yellowed in spring. Alatae, BL c.
3.1 mm with pale green abdomen, and secondary rhinaria distributed ANT III
15-18, IV 5-8, V 5-7, VI 0, migrate to an unknown secondary host. (See also discussion of this species under P. crataegicola.) Prociphilus
taxus Ghosh, Chakrabarti, Chowdhuri &
Raychaudhuri Only known from the
alate sexupara, collected on roots of Taxus
baccata in Himachal Pradesh, India (A.K. Ghosh et al. 1969); BL 2.9-3.5
mm, secondary rhinaria distributed ANT III 18-22, IV 8-10, V 6-10,
VI2-5). Sexuparae have also been
trapped in Pakistan (Naumann-Etienne & Remaudière 1995). The anal plate
projects as a collar around the cauda; a similar anocaudal process is known
in sexuparae of certain other Prociphilus
(keyed by A.K. Ghosh 1984, p.287). The
life cycle is unknown. Prociphilus
(Paraprociphilus) tessellatus (Fitch) (fig.18C) Brownish aphids living beneath a copious
secretion of white woolly wax.
Fundatrices and their alate progeny develop on undersides of leaves of
Acer saccharinum in May-July,
causing slight leaf-curl. Migration
occurs to Alnus, where colonies of
large brown wax-covered apterae (BL 3.6-4.4 mm) are formed along the branches
(see influentialpoints.com/Gallery). Alate sexuparae return to Acer in September, but anholocyclic
overwintering also occurs under fallen leaves or in loose earth at base of
alder trees (Patch 1911, Mordvilko 1935).
Widely distributed in North America.
A syrphid (Chrysopa slossonae)
lays its eggs specifically in relation to colonies of P. tessellatus (Bristow 1988).
2n=6 (Blackman & Eastop 1994). Prociphilus umarovi Narzikulov Apterae
(fundatrices) are green; BL 2.0-2.3 mm. Curling leaves of Lonicera spp. in spring. All progeny
are alate (BL 2.2-3.6 mm), migrating in June to conifer roots (Abies sibirica, Picea schrenkiana; Kadyrbekov 2017a). In Central Asia (Kazakhstan, Tajikistan). It is
possible that P. himalayensis (q.v.) is a synonym. Gabrid (1989)
described a much larger aphid from Lonicera spp. in Kyrgyzstan (BL of
fundatrices 3.8-4.0 mm, BL of alatae 3.8-4.3 mm), with 14-20 rhinaria on ANT
III, as Prociphilus sp. aff. alatavicus Juchnevitch. P.
alatavicus is currently regarded as a synonym of P. umarovi.] Prociphilus
ushikoroshi Shinji Fundatrices and their developing all-alate
progeny feed on leaves of Photinia
(=Pourthiaca) villosa in spring, causing edges of leaves to curl towards
underside and become thickened and yellowish green (Monzen 1929; as P. pourthiacae). Alatae (BL c. 2.1 mm, with yellowish green
abdomen covered in white woolly wax, and secondary rhinaria distributed ANT
III 10-14, IV 4-7, V 1-4, VI 0-1) migrate in June. The secondary host is
uncertain, possibly roots of Larix kaempferi (Shinji 1932, Moritsu
1983). Sexuparae return to deposit
sexual morphs on trunk of Photinia
in October (Monzen). In Japan and
Korea. [In India, alatae with many more secondary rhinaria (III 35-36, IV
9-10, V 10-11, VI 0-2) have been collected on Photinia notaniana (A.K. Ghosh et al. 1970; Prociphilus
"sp. A").] Prociphilus
(Stagona) xylostei (De Geer) (Fig.38a) Fundatrices (BL 3.0-4.6 mm) and their developing
alate progeny live in copious wax on shrubby species of Lonicera in spring, causing leaves to curl and become spotted
with yellow (Mordvilko 1935, Heie 1980b, and see bladmineerders.nl). Alatae (BL 2.3-3.7 mm, with pale green
abdomen and white woolly wax) complete their development on the twigs in late
June-July and migrate to found colonies of small, wax-covered apterae (BL
0.8-1.2 mm) on fine roots of conifers. Picea
abies is the recorded secondary host in Europe, and P. obovata in Central Asia. In North America it is reported from Abies grandis, but there are some
discrepancies between the redescription of apterae from Abies roots by Pike et al.
(2012) and the descriptions of European material from roots of Picea.
Sexuparae return to Lonicera
in October-November. Widely
distributed in Europe, and in Central Asia (Kazakhstan; Kadyrbekov 2017a),
India (Uttar Pradesh, Kashmir), China (G. Zhang et al. 1993c; recorded from Abelia
biflora), and introduced to North America. Differences from P. piceaerubensis were discussed by Smith (1969). Populations in east Siberia are regarded as
a subspecies, P. xylostei ussuricus Pashchenko 1988a (= P. xylostei ssp. orientalis Mordvilko 1935; nec orientalis Mordvilko 1929).
2n=10 (samples
from Europe and India).
About 50 nominal species
closely related to Aphis, but with
short appendages and other morphological features adapted to feeding at stem
bases and on roots of plants in close association with ants. Most species
feed on Compositae/Asteraceae. It is a taxonomically difficult group, many
very similar species having been described, and there is a danger that
morphological resemblance could be due to convergence associated with feeding
position and ant attendance. Most nominal species are in Europe and Central
Asia, but three were described from Africa, and six are North American. The
North American and Eurasian/African species show some differences, but there
is now molecular evidence that they may be sister groups clustering
separately from Aphis (Lagos et al. 2014), and therefore justifying
the treatment of Protaphis as a
distinct genus. However this still needs to be confirmed by more
comprehensive DNA studies of Palaearctic species. Kadyrbekov (2001b) reviewed and keyed the
central Asian species, and Nieto Nafría et al. (2005a)
reviewed those on the Iberian peninsula. Protaphis afghanica (Narzikulov &
Umarov) Apterae are green with
darker dorsal markings; BL c. 1.6 mm. On leaves and shoot apices of Artemisia
turanica in Afghanistan. Protaphis alhagii Juchnevitch Apterae are dark green, wax dusted; BL
c.1.7 mm. In ant-attended colonies at stem bases and on roots of Alhagi spp. in southern Kazakhstan. Protaphis ancathiae Kadyrbekov Apterae are greenish marked with dark
brown, siphunculi black; BL 1.46-1.7 mm. On roots of Ancathia igniaria in Kazakhstan (Kadyrbekov 2001b); subsequently
recorded from Cirsium incanum (=
arvense) (Kadyrbekov
& Aoitzhanova 2005). Protaphis anthemiae Ivanoskaya Apterae are dark greenish, with slight grey
wax film, siphunculi black and cauda green; BL 1.7-1.8 mm. On roots of an Anthemis sp. attended by ants.
Described from Azerbaijan, and subsequently recorded from Kazakhstan
(Kadyrbekov 2017a). Protaphis anthemidis
(Börner) Colour of apterae in life unrecorded; BL
1.4-2.0 mm. On roots of Anthemis
tinctoria in central and eastern Europe. It is also recorded from Iran
(Rezwani et al. 1994) and
Kazakhstan (on Achillea millefolium;
Kadyrbekov 2011b). Monoecious holocyclic with apterous males (Tashev 1964a).
Very similar to P. carlinae and P. dudichi; experimental work is
needed to determine whether or not these are all distinct species. Protaphis anuraphoides (Nevsky) Plate 7d Apterae are green to dark green, coated
with mealy wax; BL 1.5-2.1 mm. Alatae
have secondary rhinaria distributed III 5-10, IV 0-2. On upper sides of leaves,
stems and flowerheads, often in large numbers. On plants in several composite
genera of tribe Cardueae (e.g. Carthamus,
Carduus, Cirsium, Cousinia,
Onopordum). Records from genera in other subfamilies of
Compositae/Asteraceae are likely to be misidentifications. In Ukraine,
southern Russia and south-west and Central Asia. Kadyrbekov (2001b)
redescribed this species; it is very similar
to P. carthami. Protaphis aralensis Kadyrbekov Apterae are greenish, with dark brown
head, dorsal markings, siphunculi and cauda, covered with a slight grey wax
film; BL 1.7-2.0 mm. On Scorzonera
parviflora, at base of stem, attended by ants. In southern Kazakhstan
(Kadyrbekov 2001b). Subsequently recorded from Tragopogon dubius (Kadyrbekov
& Aoitzhanova 2005). Protaphis astrakhanica Stekolshchikov Apterae are greyish green, lightly
wax-powdered; BL 1.9-2.4 mm. Alatae have secondary rhinaria distributed III
4-8, IV 0-1, V 0. At stem bases near root collars of Lactuca tatarica (= Mulgedium
tataricum) in southern Russia (Astrakhan province), covered by soil and
attended by ants (Stekolshchikov 2005). The life cycle is unknown, but
probably monoecious holocyclic. Protaphis betpakdalensis Kadyrbekov
Apterae are dark greenish variably marked dorsally with dark brown,
and a slight grey film of wax; BL 1.9-2.2 mm. On roots of Taraxacum monochlamydeum, attended by
ants (Kadyrbekov 2001b). In Kazakhstan. Protaphis
bimacula (Narzikulov & Umarov) Apterae are dark green, wax-dusted, with
dark brown head, antennae, apices of femora and tibiae, and a pair of black
patches on the dorsal abdomen between the short, dark siphunculi; BL c. 1.45
mm. Colonising apices of stems of Campanula sp. in Afghanistan. Protaphis carlinae (Börner)
Apterae are dark blue-green; BL 1.5-2.1 mm. On rosette leaves of Carlina spp. in central and eastern
Europe, and now recorded from southern England (see influentialpoints.com/Gallery).
Monoecious holocyclic. Tashev (1964a) redescribed this species,
including oviparae; it is very similar to, and possibly synonymous with, P. dudichi. Protaphis
carthami (Das) Apterae are dark green or reddish brown
coated with greyish white wax, and often there are a pair of dark spots on
dorsal abdomen just anterior to the siphunculi; BL 1.6-2.3 mm. Apterae have
secondary rhinaria distributed III 0-6, IV 0-2, and alatae have them
distributed III 5-7, IV 2-3. On stems and flowers of Carthamus tinctorius and
other Carthamus spp., attended by
ants. In north Africa, Lebanon, Iran, Kazakhstan, Pakistan and north-west
India (Naumann-Etienne & Remaudière 1995). Kadyrbekov (2005b) records it also from several
other composite genera. Specimens in the BMNH collection from Atractylis flava and Hedypnois rhagadioloides in Egypt and
from Cirsium lanceolatum in North
Yemen may also be this species, although these are intermediate in some
respects between carthami and anuraphoides.
North African records of anuraphoides and
pseudocardui from Carthamus are possibly referable to carthami. P. echinopis on Echinops is
very similar and possibly a synonym. Protaphis chondrillae
(Mordvilko in Tarbinsky and Plaviltchikov) Apterae are dark green with slight grey
wax film, with black siphunculi and dark brown dorsal markings; BL 1.8-1.9
mm. On roots or subterranean stems of Chondrilla
sp., ant-attended. Southern Kazakhstan (Kadyrbekov 2001b). Protaphis cousiniae Kadyrbekov
Apterae are dark greenish with slight grey film; BL 1.6-2.3 mm. On
roots of Cousinia spp., attended by
ants (Kadyrbekov 2001b, 2005d, 2017a). In Central Asia (south and east
Kazakhstan). Very similar to P. terricola. Protaphis
debilicornis (Gillette
& Palmer) Apterae are dark olive
buff; BL 1.6-2.1 mm. In curled leaves and on stems of Helianthus spp., widely distributed in USA. Monoecious
holocyclic, with oviparae and alate males in September (in Colorado; Palmer
1952). Lagos et al. (2014)
transferred this species to Protaphis. Protaphis
delottoi Eastop Colour of apterae in life unknown,
probably dark green or brown dusted with wax; BL 1.5-1.8 mm. Apterae have 0-4
secondary rhinaria on ANT III, and alatae have them distributed III 9-13, IV
0-4. On Pluchea dioscoridis,
probably at base of plant with ants. In Eritrea, and possibly Nigeria (Eastop
1961a). Protaphis dudichi (Börner) Colour of apterae in life unrecorded; BL
1.5-2.0 mm. On roots of Matricaria and Tripleurospermum spp. in central and eastern Europe, and
also in western Siberia (Stekolshchikov et
al. 2007a). Börner’s material was redescribed by Tashev (1964a), who gave
some possible differences from the very similar P. anthemidis, but these could be host-related, and it has been
treated in catalogues as a synonym. However, it resembles P. carlinae even more closely, and
further work is needed to establish whether one, two or three species are
involved. Protaphis
echinaceae (Lagos &
Voegtlin) Colour of apterae in life
is unrecorded; BL 1.5-2.0 mm. On underside of flowerhead and leaves of Echinacea angustifolia, attended by
ants, which often build shelters over leaf-feeding colonies (Lagos &
Voegtlin 2009). Only known from Minnesota, USA. Alatae have secondary
rhinaria distributed ANT III 2-15, IV 3-10, V 1-5. Sexual morphs and life
cycle are unknown. Lagos et al.
(2014) transferred this species to Protaphis. Protaphis echinopis (Hille
Ris Lambers) Colour of apterae in
life is unrecorded, probably dark green with some wax; BL 1.2-2.2 mm. Apterae
without secondary rhinaria, alatae with them distributed III 6-10, IV 0-1. On
stems and leaves of Echinops
spinosus, and possibly other Echinops
spp., in north-east Africa (Egypt, Eritrea, Ethiopia) and Middle East
(Israel, Yemen). Records of P.
anuraphoides and P. pseudocardui on
Echinops in this region should
perhaps all be referred to this species. However, the distinction from P. carthami on Carthamus is unclear, and P. echinopsicola
(Kadyrbekov), described from Echinops
albicaulis in Central Asia, seems to be intermediate in morphology
between A. echinopis and A.
terricola. Protaphis elatior (Nevsky)
Apterae are dark green, with a fine blue pruinose covering, black eyes
(cf. elongata) and black transverse
bands on posterior abdomen; BL 1.5-2.0 mm. Described from roots of Artemisia
annua, and subsequently recorded from other Artemisia and Seriphidium spp.
Records from various other Compositae/Asteraceae (Ivanoskaya 1960; Narzikulov
et al., 1971) are perhaps referable
to other Protaphis spp.
Ant-attended. In Russia, Iran (Rezwani et
al. 1994) and Central Asia (Kadyrbekov 2001b). Protaphis elongata (Nevsky) Apterae are glaucous green with blue-grey
pruinose covering, dark head and black markings on thorax. Eyes are red (cf. elatior). BL 1.4-1.7 mm. On roots and
subterranean stems of Artemisia and Seriphidium spp., attended by ants. There are also a few records
from other Compositae/Asteraceae, e.g. Xanthium
strumarium. In Uzbekistan, south-east Kazakhstan, Ukraine, and Iran (BMNH
collection, leg. R. van den Bosch), and also reported from Poland, Rumania
and Moldova. Some records of this species including those from western
Siberia and Denmark are possibly ascribable to P. miranda (Kadyrbekov 2001b). Protaphis evansi (Eastop) (Fig.57a) Apterae are dull, mottled greenish or
brownish red; BL 1.2-1.9 mm. On Vernonia
glabra, at base of stem with ants. Also recorded from Leucospermum cordifolium and Hypochaeris radiata. In southern
Africa (Botswana, Tanzania, Zimbabwe, South Africa). Kadyrbekov (2001c)
placed this species in a new genus (Eastopiella),
but some of the characters that he described (shape of cauda, presence of
median ocellus in aptera, number of accessory hairs on R IV+V) do not agree
with type material in the BMNH collection, and in most respects P. evansi seems to be a typical Protaphis. Protaphis filaginea (del Guercio) Apterae are
dark green to brownish; BL not recorded. Described from Filago germanica (= F.
vulgaris), and also possibly on Filago (=Logfia) gallica, if it is the same species as Aphis
filaginis Lichtenstein, which is a nomen nudum. In southern France and
Portugal. Protaphis formosana (Takahashi) Apterae are dark green, with black head,
antennae, siphunculi and cauda; BL c.1.8 mm. On Blumea spp. in Taiwan.
Apterae have 1-5 secondary rhinaria on ANT III, alatae have 6-8 on III, and
0-1 on IV. Protaphis
hirsuta (Nevsky) Apterae (immature?) are dark green to
greenish brown, wax-coated; BL c.1.2 mm. On undersides of leaves and stems of
an Artemisia sp. (?vulgaris) in Uzbekistan. Described in Cryptosiphum, and subsequently placed
in Xerobion, but apparently the
type specimens are an immature Protaphis
(Kadyrbekov 2014b). Protaphis iliensis Kadyrbekov
Apterae are dark greenish with grey wax film, with dark dorsal
sclerotisation, siphunculi and cauda; BL 1..6-2.3 mm. On stems and flowers of
Acroptilon australe (= Rhaponticum repens), attended by ants; subsequently recorded
from Karelinia caspia (Kadyrbekov & Aoitzhanova 2005). Sedighi et al. (2020b) also record it from Artemisia (= Seriphidium) sieberi and
Gundelia sp., and provide a
redescription. In Kazakhstan and Iran.. Protaphis kareliniae Kadyrbekov, Renxin & Shao Apterae are yellow-green, with slight
grey wax film and dark brown head, siphunculi and dorsal markings; BL 2.1-2.1
mm. Secondary rhinaria in alata (1 specimen) distributed III 6-8, IV 1. On
roots of Karelinia caspia, attended
by ants. In Xinjiang-Uygur region, western China (Kadyrbekov et al. 2002). Protaphis
knowltoni (Hottes & Frison) Apterae are greenish blue to yellowish
green, with black dorsal markings and greyish pruinose reticulation; BL
1.8-2.3 mm. Apterae have secondary rhinaria distributed III 0-13, IV 1-8, V
0-3, VI BASE 0-1; alatae have them distributed III 20-32, IV 7-15, V 3-8. In
ant-attended colonies on roots of Taraxacum officinale and also recorded
from Achillea millefolium (Palmer
1952) and Dahlia sp. (BNHM
collection, leg. A.G. Robinson). Palmer also records it from Zea mays, but this needs confirmation
as there is possible confusion with P.
middletonii. In USA and Canada (Manitoba). Monoecious holocyclic, with
oviparae in September (males undescribed). 2n=8. Protaphis kvavadzei Bardjadze & Özdemir Colour of apterae in life unknown; BL
1.5-1.9 mm. Apterae have secondary rhinaria
distributed III 0-6, IV 0-3, V 0; alatae (2 specimens) have
them distributed III 12-19, IV 2-3, V 0. On Eryngium
campestre in central Turkey (Barjadze et
al. 2014). Protaphis lactucicola Kadyrbekov Apterae are dark greenish with slight
grey wax film, brown head, usually a brown central dorsal transverse mark
just anterior to siphunculi, and dark brown siphunculi; BL 1.6-1.9 mm. In
ant-attended colonies on roots of Lactuca and Crepidiastrum in south-east Kazakhstan. Very similar to P. pseudocardui (q.v.). Protaphis
middletonii (Thomas) (= Aphis
armoraciae Cowen, = Aphis
maidiradicis Forbes, = Aphis
menthaeradicis Cowen) Plate 7e Apterae are pale green, bluish green,
grey-green or olive-green, often dusted with greyish wax, and with
variably-developed dark dorsal markings or cross-bands, more apparent in apterae than in alatae (see
influentialpoints.com/Gallery); BL 1.5-2.5 mm. Apterae have secondary rhinaria
distributed III 0-17, IV 0-8, V 0-5*, and alatae have them distributed III
4-22, IV 0-9, V 0-6*. Ant-attended colonies occur commonly in western and
central USA on roots of plants in numerous families including
Compositae/Asteraceae, Brassicaceae, Poacaeae, Lamiaceae and Apiaceae. Aphids
of this group are also recorded from Compositae/Asteraceae in Brazil, and
from Opuntia roots in Australia and
South Africa. Records from Centaurea
in Turkey (Akyürek
et al. 2011) and from Taraxacum in Algeria (Laamari et al. 2013) need additional
confirmation. Eastop & Blackman (2005) concluded on the basis of multiple
discriminant analysis that a single name should be applied to this group, the
oldest available being middletonii
Thomas. Monoecious holocyclic, but anholocyclic overwintering is probably
also common. In the North American literature, populations identified as armoraciae are stated to produce
alate males, whereas maidiradicis and
menthaeradicis have apterous males,
and oviparae of menthaeradicis are
said to have very few scent glands on their hind tibiae (Palmer 1952).
However, the criteria for naming these populations are unclear, and
experimental work is needed to show whether these differences are of any
taxonomic significance. Two other North American taxa described by Pack &
Knowlton (1929), Aphis
crypta and A. gutierrezis (as Anuraphis gutierrezis), are closely related and probably part
of this group. 2n=8. *the
numbers of rhinaria given in Blackman & Eastop (2006) were erroneous Protaphis miranda Kadyrbekov
Apterae are dark greenish with slight grey film, dark reddish eyes,
black siphunculi and green cauda: BL 1.4-1.8 mm. In ant-attended colonies on
roots of Artemisia and Seriphidium spp. in Kazakhstan
(original description), west Siberia (Ivanoskaya, 1977, as alexandrae) and Europe (Denmark,
Poland - Heie, 1986, as elongata;
Czech Republic - BMNH collection, leg. J. Holman), and also recorded from two
Tanacetum spp. (Kadyrbekov 2017a).
Probably monoecious holocyclic; an ovipara from A. campestris in Poland (BMNH collection, leg. L. Olesinski) may
be this species. Protaphis
pseudocardui Theobald (Fig.31c) Apterae are dark green or brown with
covering of grey mealy wax, short black siphunculi, and often there are a
pair of dark spots on dorsal abdomen just anterior to siphunculi; BL 1.5-2.0
mm. In ant-attended, often dense colonies on stems, upper sides of leaves,
flowerheads, or on root collars. Described from “thistles (Carduus sp.)” in South Africa; however
this could have been Sonchus, which is widely known as “thistle” in
Africa. Since recorded from other
parts of Africa, Mediterranean region, the Middle East and Georgia (Barjadze
2008), and from various other Compositae (Arctotheca,
Berkheya, Carduus, Carthamus, Cichorium, Hypochaeris, Lactuca, Pluchea,
Pulicaria, Vernonia), although some of these records may be referable to
other closely-related species, including the terricola group. 2n=8. Protaphis
rutae (Nevsky) Apterae are dark green, with a light wax
covering, siphunculi black and cauda green; BL 1.4-2.0 mm. On Ruta sieversii (= Haplophyllum acutifolium?), forming large colonies on upper and
undersides of leaves. In Central Asia (Uzbekistan). This species was redescribed and
transferred to Protaphis by
Kadyrbekov (2001b). Protaphis scorzonerae (Mordvilko) Apterae are brownish green, covered with “rather bright azure-blue velvet-like powder” (Ghilarov 1937, as Xerophilaphis scorzonerae); BL 1.7-2.3 mm. In ant-attended colonies on roots (where they may feed in lesions produced by the ants; Ghilarov1937) and at bases of leaves, of Compositae/Asteraceae (Taraxacum, Hieracium, Scorzonera, |
