SYSTEMATIC TREATMENT OF APHID GENERA
(in alphabetical order)
Two North American oak-feeding species with a short antennal terminal process, indented anal plate and rounded cauda. Apterae have the head and thorax fused and an extensively sclerotised tergum, and alatae have wings strongly resembling those of Patchia, with thickly bordered veins and scale-like imbrication of the wing membrane. Quednau (1999) reviewed and illustrated both species.
Apterae are mid- to dark brown or almost black; BL 1.6-2.0 mm. Alatae are dark reddish brown with dusky
wings, with dark dorsal abdominal cross-bands and wing veins thickly bordered
with fuscous. In small colonies on
twigs and leaf petioles of young Quercus
(nigra, michauxii), attended by ants, or in small numbers on undersides
of leaves, where the alatae look very conspicuous against the green of the
leaf. In south-eastern
(= Patchia winforii Miller)
Apterae are dark brown to black, "almost shiny"; BL 1.7-1.9
mm. Alatae are dark brown to black,
with black abdminal cross-bands and wing veins broadly and heavily bordered
with fuscous. In small colonies,
mainly on leaf petioles and young bark of tender one-year-old shoots, of Quercus rubra (= borealis) and Q. velutina,
and also reported forming dense clusters on twigs and leaf petioles of Q. palustris (Miller 1933b, as Patchia winforii). Closely attended by the ant Crematogaster lineolata, which may
construct shelters over the colonies.
15 palaearctic and one nearctic species
of medium to large long-legged aphids, the alatae usually having pigmented
wings, mostly associated with Fagaceae and attended by ants. It is a taxonomically difficult genus, as
indicated under L. roboris. Accounts are available for central
acutihirsutus Kumar & Burkhardt Apterae have black head and thorax and
brownish black abdomen, with a smooth membranous conical protrusion in the
centre of the dorsal abdomen; BL 2.8-4.4 mm.
Alatae have forewings pigmented except for two large clear triangles
on basal half and a small clear spot between pterostigma and RS.
On twigs of Quercus incana in
allegheniensis McCook (= montanus
Lachnus chosoni Szelegiewicz Apterae are in life very shiny dark brown to black (Kanturski et al. 2018c); BL 3.7-4.1 mm. On bark of older branches of an unidentified Quercus sp. in North Korea (Szelegiewicz 1975a, and see also Binazzi & Remaudičre 2010). Kanturski et al. (2014b) re-examined the type material, and Kanturski et al. (2018c) reported this species from South Korea (also from an unidentified Quercus sp.), and described oviparae and apterous males collected in October-November.
crassicornis Hille Ris Lambers (fig. 93A)
Colour of apterae in life is not recorded;
BL 2.2-3.0 mm. On Quercus ithaburensis (BMNH collection, leg. D. Hille Ris Lambers)
Takahashi Apterae brownish black; BL
c. 4.5 mm. Alatae have blackish brown
pigment at bases of wings, not extending to Cu1b in forewing, and
wing membrane otherwise only slightly dusky around margin. On branches of an unidentified Ficus sp. in
iliciphilus (del Guercio) Apterae are shining blackish brown; BL
2.6-3.7 mm. Alatae have a pattern of forewing pigmentation very similar to
that of L. roboris. On twigs and
branches of Quercus spp. (ilex, suber, coccifera) in
longirostrum David & Ghosh Colour of apterae in life unknown; BL c.
4.4 mm. Alata with hyaline forewings. Described from Salix fragilis in
Lachnus margallaensis Kanturski & Zia Apterae in life are dark green; BL 2.4-2.8 mm. Alatae have forewing pigmentation similar to L. acutihirsutus. On branches and shoots of Quercus oblongata in Pakistan (Kanturski et al. 2017b).
(= Lachnus longirostris (Mordvilko);
Mróz et al. 2015) Apterae are shining grey-brown to dark
reddish or blackish brown, with a clothing of fine pale hairs (see influentialpoints.com/Gallery);
BL 2.8-5.0 mm. Alatae have a pattern
of forewing pigmentation similar to L.
roboris, but the clear patch between the Rs and the media is
larger. On older branches and stems of
Fagus sylvatica, causing feeding
damage by rupture of the cambium, and also on twigs and branches of mostly
evergreen species of Quercus, since
Mróz et al. (2015) have provided
molecular and morphometric data indicating that L. longirostris is a synonym. In
Lachnus pseudonudus Kanturski, Wieczorek & Junkiert Appearance in life unknown; BL of aptera 2.6-3.6 mm. On branches and shoots of Quercus macrolepis (= Q. ithaburensis spp. macrolepis) in Turkey (Kanturski et al. 2014b). Other morphs and life cycle are unknown.
Lachnus quercihabitans (Takahashi) Described from some alatae said to be
collected (with immatures) on Quercus
Apterae are shining blackish brown (see
BL 2.5-5.5 mm. Alatae have forewing membrane pigmented except for clear
patches on either side of Rs, a clear band running obliquely across base
of media and distal part of Cu1a, and a clear triangle from Cu1b
to the wing-base. On twigs and small
branches of Quercus spp. and also
sometimes in large numbers on Castanea
sativa in southern
salicis Chakrabarti & Raha Colour in life is not recorded, probably
brownish; BL of aptera 3.5-5.4 mm.
Alatae have clear wings. In
large dense colonies on stems of Salix
babylonica and S. tetrasperma,
attended by ants. Sexual morphs are
not recorded; apterous viviparae and immatures may overwinter in bark
crevices (Chakrabarti & Raha 1988).
Lachnus shiicola Sorin Apterae are shining brownish black; BL 3.7-4.0 mm. Forewing pigmentation of alata is similar to that of L. tropicalis. On Castanopsis sp. (?cuspidata) in Japan, feeding in scars of the bark of the trunk near its base, and on exposed roots of old trees, covered by ant shelters. Sexuales and life cycle are unknown. Closely related to L. tropicalis, but rather longer-haired and with shorter antennae and HT II (Sorin 1980).
sorini Binazzi & Remaudičre Apterae are shining blackish, with a large
prominent rounded dorsal abdominal tubercle ; BL 3.2-5.0 mm. Alatae have
forewing pigmentation similar to that of L.
roboris. On Quercus spp. in
swirskii Hille Ris Lambers Apterae are reddish brown, broadly
pear-shaped; BL 3.1-4.4 mm. Alatae
have forewing pigmentation closely resembling that of L. roboris. On branches of
young Quercus spp. (aegilops, infectoria, ithaburensis), recorded from
Apterae are brownish black, not shiny; BL 3.8-5.1 mm. Forewings of alata mainly infuscated, but
with several clear patches of irregular shape. On Quercus
tatakaensis Takahashi Apterae are brownish black, with many
large black circular spots in rows on dorsal abdomen. Antennae black, legs
reddish brown and black; BL 5.5-5.7 mm (Takahashi 1937b). Alatae have not been described. On trunk and branches of an unidentified Salix sp. at high altitude in
tropicalis (van der Goot)
(= japonicus Matsumura) Apterae are shining brownish black; BL
3.8-5.3 mm. Alatae have forewing
wholly infuscated, except for a clear patch between pterostigma and Rs, and a
clear band from base of media to distal part of Cu1a. On twigs and stems of Quercus spp. Lithocarpus
spp. and Castanea spp., in east and south-east
Apterae are grey with black hind tibiae; BL 2.1-2.8 mm. Alatae have
forewing pigmentation similar to that of L.
roboris. On Quercus persica in
wichmanni Hille Ris Lambers Apterae are dull blackish with greyish
waxy markings; Bl 3.3-4.6 mm. Sides of
head and sections of the femora and tibiae are orange-brown. Alatae have the forewing membrane
infuscated distally, and very thick bands of fuscous along Rs, media and
Cu1b. On trunk and branches
of Hippophae rhamnoides, attended
by ants. Recorded from
yunlongensis Zhang Known only from apterous viviparae (BL c.
4.6 mm) collected on Salix sp. in
One narrow-bodied species differing from Atheroides by the position of the siphunculi (on abdominal tergite 6, as opposed to abdominal tergite 5 in Atheroides) and by lack of a dorsal sclerotic carapace. Wieczorek (2010) provided a full account with redescriptions of all morphs.
Laingia psammae Theobald Apterae are dirty straw-coloured to
greyish green (see influential
points.com/Gallery); BL 1.6-2.8 mm. Alatae have dark transverse bars on
the dorsal abdomen. Typically found in inflorescences of Ammophila
arenaria in sand dunes, but also on Calamagrostis
epigeios at more inland locations, and also recorded in Sweden from Elymus, Calamagrositis arundinacea and
Deschampsia caespitosa (Heie 1982),
and in Iberian peninsula on Carex
acutiformis. Sometimes attended
by ants. Widely distributed in
A genus for one species on Populus, differing from Chaitophorus in the very short ANT PT, sparse short needle-like dorsal hairs, crater-like siphunculi without reticulate sculpturing, and semicircular cauda. The alatae have thickly fuscous-bordered wing veins like the North American Chaitophorus populicola (which also has a rounded cauda).
Apterae are blackish brown (Qiao et
al. 2003a); BL c. 1.5-1.7 mm. On
young shoots of Populus pruinosa
and P. diversifolia (= P. euphratica) in central
Landisaphis Knowlton & Ma
A genus for one little-known North American species with flattened, mostly spatulate dorsal hairs, a reticulate dorsal cuticle and clavate siphunculi. The sclerites of the spinal hairs on abdominal tergites 6-8 are developed into rugose conical processes (less developed in alatae).
Landisaphis davisi Knowlton &
Ma (Fig.37a) Colour of apterae in life is unrecorded,
BL 0.9-1.4 mm. Described from apterae collected on undersides of leaves of Chenopodium album, although this
perhaps requires confirmation as subsequent records have been from
Brassicaceae (Lepidium perfoliatum,
Descurainia sophia; BMNH
collection), and it has been reared on Capsella
bursa-pastoris (A. Jensen, pers. comm).
Only known from the state of
or two species on Alnus in
orizabaensis Remaudičre (fig. 27F,G) Apterae are flattened dorsoventrally, pale
yellow with black antennae; BL 1.6-2.2 mm.
Alatae have black head, thorax and appendages and black dorsal
abdominal markings. On undersides of
leaves, especially on young branches, of Alnus
acuminata ssp. arguta in
One species in east
papillicaudus Gredina Apterae are yellowish green; BL c.1.2 mm.
Alatae are unknown. On Elsholtzia
pseudocristata (= E. ciliata) in east
Lepidaphis Kadyrbekov, Renxin & Shao
Two eastern palaearctic species on Lepidium in desert regions, related to Brevicoryne and Brachycolus but with numerous hairs on the front of the head and abdominal tergite 8, and a short antennal terminal process (Kadyrbekov et al. 2002). Life cycles are unknown.
Apterae are greenish, with slight wax film; BL 1.7-2.2 mm. The single
known alata has 12-14 secondary rhinaria on ANT III. In leaf galls on Lepidium spp. in
terricola Kadyrbekov, Renxin &
Shao Apterae are greenish, with
slight grey wax film; BL 1.9-2.1 mm. On roots of Lepidium spp. in south-east
lini Zhang Apterae are grass-green; BL c. 1.8 mm. On
Linum usitatissimum in
Three palaearctic and one
nearctic species resembling Illinoia
in the tendency for siphunculi to be swollen proximal to the subapical,
reticulated zone, but the degree of swelling varies considerably within
species. Heie (1994) provided an account of two species in
asperulophagum Holman Apterae are green or pinkish with shiny
brown-black dorsum, head and basal antennal joints, and dark siphunculi; BL
1.7-2.3 mm. Alatae are as yet undescribed; a specimen in the BMNH collection
(leg. H.L.G. Stroyan) has ANT III with 5-6 small rhinaria, and dorsal abdomen
with incomplete dark cross bars and large marginal and postsiphuncular
sclerites. On undersides of leaves and
upper parts of stems of Asperula
galii Mamontova Plate 26a Apterae are shiny green or brown with
variably developed black pigmentation of dorsum, and having dusky brown
siphunculi with dark apices; BL 1.7-2.0 mm. Alatae have incomplete dark
dorsal abdominal cross bands and large marginal and postsiphuncular
sclerites. On undersides of leaves of Galium spp. and Asperula odorata. This species has been confused with L. asperulophagum but has a shorter R
IV+V and longer HT II, the ratio between these two providing a reliable
galiophagum (Wimshurst) Apterae are shiny green, their
siphunculi pale with dark apices; BL 1.7-2.5 mm. Alatae have dark marginal
abdominal and intersegmental pleural sclerites but no dark cross bands. On Galium spp., living on undersides of
leaves along veins, and on young shoots.
& Frison) Apterae have pearly
white to yellowish head and posterior part of abdomen including siphunculi
and cauda, the thorax and anterior abdomen being bright shining ruby-red
(presumably due to sap of host); BL c. 1.6 mm. On undersides of leaves,
feeding singly near larger veins, of Sanguinaria canadensis. In
north-eastern and north-central
Four species resembling Cavariella but without a supracaudal process, and typically associated with Berberis.
atra Hille Ris Lambers Apterae are dark purplish brown, dirty
greenish in centre of dorsal abdomen,
with pale legs and antennae, siphunculi dark on distal half; BL c.1.3-1.6 mm.
Alatae have a brown dorsal abdominal patch (L.K. Ghosh & Pramanik 1976).
On Berberis spp. in central and
east Asia (Iran, Kazakhstan,
(= Liosomaphis turanica Narzikulov) Plate 16a (Fig.17e,f) Apterae are yellow to yellow-green, or
pinkish to orange-red (two colour forms), slightly wax-powdered (see influentialpoints.com/Gallery); BL 1.1-2.3 mm. Alatae have a dark head, thorax
and antennae, but little or no dark dorsal abdominal pigmentation. On
undersides of leaves of Berberis and
Basu Apterae are shiny pale
yellowish, with green to yellowish brown markings on thorax and abdomen; BL
1.7-2.5 mm. Alatae have the dorsal abdomen with variable brownish
sclerotisation (A.K. Ghosh 1969). On
undersides of leaves of Berberis spp.
A genus for one species superfically resembling Lipaphis, but perhaps more closely related to Myzus.
About 12 mostly western
palaearctic species associated with Brassicaceae, related to Brevicoryne and characterised by
weakly developed antennal tubercles, apterae with rather short antennae which
are almost always without secondary rhinaria, a sclerotic but often weakly
pigmented tergum, and weakly swollen siphunculi. Subgenus Lipaphidiella is distinguished by the presence of a conical,
rather scabrous, supracaudal process. Accounts are available by
alliariae Müller Apterae are dark blue-green to almost
black (see influentialpoints.com/Gallery); BL 1.6-2.1 mm. Alatae have secondary rhinaria
distributed III c. 22, IV 5-6. On Alliaria
cochleariae Jacob Apterae are dull olive green with
variably-developed brown patches; BL 1.2-2.1 mm. Alatae have secondary
rhinaria distributed III 19-30, IV 5-11, V 0-4. On Cochlearia officinalis in the intertidal zone, on rosettes of
young plants, or in flower-heads. Only known from
Lipaphis erysimi (Kaltenbach) Apterae are yellowish green, dirty green or brownish; BL 1.5-2.3 mm. Alatae have secondary rhinaria distributed III 9-32, IV 2-10, V 0-3. On various Brassicaceae (Arabis, Capsella, Coronopus, Erysimum, Isatis, Lepidium, Matthiola, Sinapis, Sisymbrium, Thlaspi, etc.), but not usually on field Brassica crops. In northern Europe, and probably in western Siberia and Central Asia, but its distribution needs clarification due to past confusion with L. pseudobrassicae. Monoecious holocyclic with apterous males. The name has been commonly applied to the world-wide crucifer pest, L. pseudobrassicae (q.v.), but they differ in karyotype, and Ronquist & Ĺhman (1990) showed in laboratory trials that L. erysimi performed poorly in comparison with L. pseudobrassicae on Brassica oilseed crop plants. 2n=10.
fritzmuelleri Börner Apterae are dark green; BL 1.3-1.8 mm.
Alatae have secondary rhinaria distributed III 11-16, IV 0. On Sisybrium spp., occurring on flowers
in spring and on lower leaves in summer. In
Lipaphis (Lipaphidiella) jungarica Kadyrbekov, Renxin & Shao Apterae are greenish; BL c. 1.7 mm.
Apterae have 3-5 secondary rhinaria on ANT III (based on 1 specimen), alatae
have them distributed III 7-12, IV 0-3. On upper sides of leaves of Syrenia
siliculosa (= Erysimum siliculosum) in western
Lipaphis (Lipaphidiella) lepidii (Nevsky) (Fig.37b) Apterae are pale green; BL 1.2-1.6 mm.
Alatae have secondary rhinaria distributed III 34-48, IV (0-)1-6, V 0-1. On
undersides of leaves, stems and flower-stalks of Lepidium spp. in
pseudobrassicae (Davis) Mustard
Aphid, Turnip Aphid, False Cabbage
Aphid Plate 12d Apterae are yellowish green, grey-green
or olive-green, with a white wax bloom, which in humid conditions may become
a dense mealy coat (see aphids
of Karnataka website); BL 1.4-2.4 mm. Alatae have secondary rhinaria
distributed III 15-30, IV 3-13, V 0-3. On many
genera and species of Brassicaceae, inclunding Barbarea, Brassica, Capsella, Iberis, Raphanus and Rorippa. An important world-wide pest
of brassica crops (Blackman & Eastop 2000). Monoecious holocyclic in
rossi Börner Apterae are dark grey green with a slight
waxy bloom, with a dark grey brown head, broad dark grey brown bars across
the dorsum, and large marginal sclerites; BL 1.2-1.6 mm. Alatae have
secondary rhinaria distributed III 27-53, IV 10-26, V 2-12. Monoecious holocyclic on Arabis hirsuta in UK, with oviparae
and apterous males in October (Prior 1971). Infested plants have stunted
flower stems and deformed inflorescences. Also recorded from
Lipaphis (Lipaphidiella) ruderalis Börner ( = Lipaphis berteroaella Mamontova) (Fig.37c) Apterae are greyish green; BL 1.8-2.2 mm.
Alatae have secondary rhinaria distributed III 35-40, IV 5-7. On undersides
of leaves, stems and flower-stalks of Lepidium
spp. in eastern Europe, and also in
Lipaphis sisymbrii Bozhko Apterae are yellow-green; BL c. 1.7-1.8 mm. Alatae have secondary rhinaria distributed III 26-30, IV 8. On Sisymbrium polymorphum in Ukraine, and also reported on the same host as well as on S. loeselii in Kazakhstan (Kadyrbekov 2009a, 2017a).
turritella (Wahlgren) Plate 12c Apterae are greenish yellow to yellowish
brown, dusted with white wax; BL 1.5-2.3 mm. Alatae have secondary rhinaria
distributed III 40-53, IV 10-24, V 2-12. On Arabis (=Turritis) glabra, causing deformation of
inflorescences. Also recorded from Arabis
pendula (Kadyrbekov 2017a) and Erysimum
cheiranthoides (Ivanoskaya 1977). In
unguibrevis Zhang Colour of apterae in life is unrecorded;
BL c.1.9 mm. Alatae have secondary rhinaria distributed III 5-8, IV 0-2. On Brassica sp. at 3,800 m in
Two little-known east Asian species on Fagaceae, tentatively placed together because they have heavily sclerotised apterae with fused head, thorax and abdominal tergites 1-7, and siphuncular pores. Qiao et al. (2005a) reviewed the genus.
Apterae are black, rather densely covered with white secretion, almost
circular, and dorsally flattened; BL c. 0.75 mm. On undersides of leaves of Lithocarpus sp. in
Apterae are black, shining, almost circular, and strongly convex
dorsally; BL 1.2-1.5 mm. Alatae have
secondary rhinaria distributed ANT III 21-23, IV 10-11, V 8-11. Apterae are found on branches of Castanopsis cuspidata in
A South American genus with primitive features and affinities to the African genus Paoliella, but with no clear host relationships. Four species are known to be tree-dwelling, two on Combretaceae and three on Lauraceae, but several other species are only known from trapped alatae. Quednau (2010) reviewed the genus and provided keys to apterae and alatae and illustrations of most of the known morphs.
Apterae are dark brown, with blue-grey wax especially around bases of
dorsal processes; BL 1.3-1.8 mm. On undersides of leaves of Nectandra reticularis in
(Paralizerius) brasiliensis Quednau Apterae are yellow-green, with long
marginal processes and eyes reduced to triommatidia; BL 1.2-2.1 mm. Alatae have dark head, pterothorax,
antennae and legs. On young shoots and
suckers of Terminalia australis in
(Paralizerius) cermelii Quednau Plate 6b Apterae
are whitish with green head and darker antennae, legs and posterior abdominal
processes; BL 1.8-2.4 mm. Described from a large colony of mainly alatae on Bougainvillea sp. in Brazil. Alatae have
been trapped in
Lizerius jorgei Cunha & Sousa-Silva Apterae are bright yellow with small greenish stripes on thorax and anterior abdomen, and eyes reduced to triommatidia; BL 1.2-2.1 mm. Alatae are also bright yellow with a dark brown thorax. On Terminalia brasiliensis in Săo Paulo, Brazil (Cunha & Sousa-Silva 2019); also found on Persea americana – possibly a less-favoured host. An alate male was collected (on Persea) in January.
ocoteae Blanchard Plate 7c, d Apterae are dark olive to brownish black,
thickly coated with white wax; BL 1.6-2.0 mm.
Alatae are brownish black with a sparse covering of wax meal. In dense colonies on young stems and leaves
of Ocotea acutifolia in
Adult apterae in life unknown; BL of alata 1.3-1.9 mm. On Nectandra picurim and Nectandra sp.
(alatae and immatures) in
pustulatus Quednau Colour in life unknown; BL of aptera c.
1.4-1.5 mm. On Ocotea sp. near glomerata and Nectandra sp. in
tuberculatus (Blanchard) Apterae are brownish black; BL 1.2-1.5
mm. Alatae are also brownish black,
and weakly pruinose. On young growth
of Nectandra sp. (Blanchard 1939)
Longicaudinus Hille Ris Lambers
corydalisicola (Tao) Apterae are pinkish grey with pale
appendages, a pale spinal stripe and glassy white wax secretion on sides and
end of abdomen; BL 1.7-2.3 mm. Immatures are pale greenish yellow, and alatae
have a dark dorsal abdominal central patch. On Corydalis spp. in
Longicaudus van der Goot
A rather distinctive genus of
about 7 species in
cornutus Chakrabarti & Banerjee Colour of apterae in life is unrecorded;
BL c.2.5 mm. On Thalictrum sp. in
dunlopi Hille Ris Lambers Apterae are creamy white; BL 1.7-2.4 mm
Longicaudus himalayensis Hille Ris Lambers Described from two alatae collected on ?Quercus, which according to Chakrabarti & Banerjee (1991a) were gynoparae that originated from Thalictrum sp. An aptera from Rosa described as himalayensis (David et al. 1971a) is included in the Rosa key on the basis of David et al.’s description, although it seems to be a different species with shorter siphunculi and higher ratio of R IV+V to HT II (see Remaudičre 1993). L. himalayensis is also included in the Thalictrum key on the basis of information in the key by Chakrabarti & Banerjee (1991a), who reared one aptera from Thalictrum. There is however no proper description of the aptera of himalayensis, either from Rosa or Thalictrum.
kumauni Chakrabarti & Banerjee Colour of apterae in life is unrecorded;
BL c.1.2 mm. On Thalictrum sp. in
naumanni Remaudičre (Fig.11c) Apterae are very pale, presumably whitish
green, BL 1.6-2.7 mm. Heteroecious holocyclic between
netuba (Zhang, Chen, Zhong & Li
) Apterae (fundatrices) are yellowish
green; BL (fundatrices) 1.6-1.7 mm (only fundatrices and immatures are
described). On young shoots of
Longisiphoniella Chakrabarti, Saha & Mandal
One species in
Saha & Mandal Appearance of
apterae in life is not recorded, BL 2.0-2.5 mm. Collected from roots of
various plants (Artemisia sp., Cynoglossum glochidiatum, Conyza
bonariensis, Rumex sp.) in west
species have been described in this genus, one in
Apterae are pale brownish-grey, slightly pruinose, with conspicuous
rows of dark dorsal spots, and dark siphunculi (see
BL 5.1-7.8 mm. Alatae have similarly
conspicuous black markings. On the bark of numerous tree species in
liquidambarus (Takahashi) Apterae black or brownish black, lightly
dusted with wax powder; BL c.7.0-7.7 mm.
On Liquidambar formosana in
xizangensis Zhang Apterae of BL 6.5-7.1 mm, colour in life
unknown. Recorded from several genera
of trees (Salix, Phoebe, Populus, Prunus, Quercus) in
One little-known species
overwintering on Lonicera in
paradoxa Narzikulov Apterae (fundatrices) are whitish green;
BL 3.4-3.9 mm. In spring colonies on undersides of leaves and growing points
of Lonicera spp., attended by ants.