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Preface vii
I INTRODUCTION 1
The Association Between Aphids and Trees 1
Life Cycles and Polymorphism 3
IV TECHNIQUES 926
Collecting Methods 926
Preservation and Mounting 927
Labelling and Storage 928
V BIBLIOGRAPHY 930
Regionally Classified Works on Tree-dwelling Aphids 930
List of References 930
One of the first responses that we had to the book Aphids on the World's
Crops, soon after its publication in 1984, was from a colleague in forest
entomology, who pointed out that, despite the title of the work, we had
neglected even to mention a crop that he regarded as rather important, namely
timber. Almost ten years later, we hope that we have now gone some way
towards rectifying that omission.
When we compiled Aphids on the World's Crops (1984) we had good
reason to turn a blind eye to the aphids on timber trees, as we knew that keys
to identify tree-dwelling aphids would be a very different task, much more
complicated than the rather simple keys that we put together ten years ago
to identify the aphids most commonly found on agricultural and horticultural
crops. When we actually got down to the job, we found it necessary to adopt
a far more comprehensive approach than in the earlier work, as we could think
of no justifiable reason for including some aphid species and omitting others.
There was also no way of providing keys to identify aphids that had not been
slide-mounted, as many of the key couplets involve relatively minor differ-
ences between closely related species, requiring microscopic examination. We
think, nevertheless, that non-specialist entomologists will be able to use this
book to identify tree-dwelling aphids at least to genus, and it may then be
possible to confirm the identity of a species by taking into account the
biological information provided in Section III.
Another small problem was to decide what is, and what is not, a tree. We
claim no expertise in this matter, and have relied on standard botanical refer-
ence works, especially Bailey (1949), Uphof (1968), Willis (1973), Boutelje
(1980) and Mabberley (1987). If a genus contains plant species classified as
trees by any or all of these authorities then we have included it. If, as is often
the case, the same genus includes species of shrubby habit, then we have also
generally included such shrubs in the host lists because, in choosing a host plant,
aphids have a lot of regard for its genus but very little regard for its growth habit.
Vll
Preface
We are grateful to several colleagues who have given permission for the
results of their (as yet) unpublished work to be included; namely B. Agarwala,
S. Akimoto, S. Aoki, S. Barbagallo, P.A. Brown, C.I. Carter, S. Chakrabarti,
R. Danielsson, J. Holman, N. Moran, J.H. Martin, D. Noordam, W.
Quednau, G. Remaudière, J.T. Sorensen, D. Stern, M.B. Stoetzel and D.J.
Voegtlin.
INTRODUCTION
1
2 Introduction
think, however, that the explanation for this can be found in the evolutionary
history of aphids rather than in their present-day host relations or ecology.
Psyllids have similar ecology and host relations to aphids, yet many tropical
trees with few aphids bear a large psyllid fauna. It seems likely to us that
aphids have failed to diversify in the tropics because of one particular,
primitive feature of aphid biology, their cyclical parthenogenesis.
Cyclical parthenogenesis is a very successful way of exploiting the short-
lived growth flushes of temperate plants, but cannot be readily adapted
to tropical conditions. Aphids moving into the tropics simply lose the sexual
phase of the life cycle, and in doing so they lose the potential to evolve
and diversify that is dependent on the recombination of genes. The tropics
may also have acted in this way as a barrier to aphid colonization of southern
temperate regions, which also have very small indigenous aphid faunas.
The occurrence of Neophyllaphis on Podocarpus, Araucaria and related
conifers throughout the southern continents testifies to the age of aphid-tree
relationships, but very little is known about such evolutionarily ancient asso-
ciations. Most ecological and experimental studies of aphid-tree interactions
have concerned introduced species. In Britain, economic damage to spruce by
sporadic outbreaks of Elatobium abietinum has been documented since 1846.
Damage to the more recently introduced Picea sitchensis is particularly severe,
heavy infestations resulting in complete needle loss. Aphid infestations have
been shown to reduce the accretion of wood (e.g. of sycamore; Dixon, 1971a),
and have deleterious effects on tree root growth (e.g. of Tilia; Dixon, 1971b).
However, none of these trees is native to Britain. There are no native British
Picea, sycamore is an introduction from Central Europe, and the common
British lime tree (or linden) is thought to be a hybrid between a native and
an introduced species.
Planted forests of exotic trees cover enormous areas of the globe. There
are more than 5.5 million hectares of planted forests in Brazil, of which at
least 40% are Eucalyptus spp. (Anon., 1985). Pinus radiata occupies only a
small area in its native California but has been widely planted in New Zealand
and elsewhere. During this century many European, oriental and American
species of Pinus were introduced to various parts of Africa and grew aphid-
free for many years. In recent times three aphids, Eulachnus rileyi from
Europe, Cinara cronartii from North America and Pineus boerneri of uncer-
tain origin, have appeared on pines in Africa and caused far greater damage
than they do in Europe or America. Similarly, Cinara cupressi is much more
damaging to Cupressaceae in Africa than in Europe. These exotic conifers may
be growing under stress, and the aphids are certainly without the complex of
natural enemies associated with them in their countries of origin.
Most aphid damage to trees seems to result directly from feeding, either
by removal of sap or wounding of tissue, or in at least some cases by the toxic
effect of saliva. Aphids are rarely recorded as vectors of viruses infecting trees
(Biddle and Tinsley, 1967). Given the astronomical numbers of aphids in the
air and the length of life of trees, there must be strong selection among trees
for resistance to aphid-transmitted viruses. It would be interesting to know
the mechanism of this resistance, and whether it could be transferred to
Introduction 3
1. The various families and subfamilies of Aphidoidea each have life cycles
with characteristic features, indicating that they have evolved independently.
2. A complete life cycle (that is, a holocycle) typically consists of one genera-
tion of sexual morphs (sexuales) and several generations in which only parthe-
nogenetic females are produced. This phenomenon of cyclical parthenogenesis
is a basic, primitive feature of aphid biology.
3. In the more primitive families, Adelgidae and Phylloxeridae, both sexual and
parthenogenetic females are oviparous, but in the Aphididae parthenogenetic
females always give birth to live young; in Aphididae the parthenogenetic
4 Introduction
Table 1. Some features of the biology, host associations and life cycles of Aphidoidea. The
bracketed figures are total numbers of genera/total numbers of species : numbers of genera
known to live on trees/numbers of species known to live on trees. The classification follows
that used in B & E (1984) rather than that of Heie (1980), which raises most subfamilies to
family level. Table 3 (p. 539) compares the two classifications.
1 ADELGIDAE (2/49:2/49) alternation from galls on Picea to other Pinaceae, holarctic
1.1 Pineini (1/21:1/21) Picea to Pinus,
1.2 Adelgini (1/28:1/28) Picea to Larix, Pseudotsuga, etc.
APHIDIDAE
3 Pemphiginae (48/319:47/204) dwarf arostrate sexuales, oviparae produce only one egg
3.1 Eriosomatini (13/96 :13/58) galls or pseudogalls on Ulmaceae to various
angiosperms, often on roots, with ants; holarctic
3.2 Pemphigini (18/168 :17/108) holarctic
3.2.1 Pemphigina (6/78 :6/49) galls on Populus to roots of various herbs, more
rarely trees or aerial parts of herbs
3.2.2 Prociphilina (12/90:7/59) pseudogalls on various dicot trees to roots of
Coniferae, more rarely other plants
3.3 Fordini (17/55:17/38) alternation from galls on Anacardiaceae
3.3.1 Fordina (12/41:12/26) Pistacia, mostly to grass roots with ants, mostly
Mediterranean and East Asia, 1 American genus
3.3.2 Melaphidina (5/14:5/12) Rhus, mostly to mosses, mostly oriental, 1 American
genus
4 Hormaphidinae (45/183:42/144) host alternating, with small rostrate sexuales
4.1 Cerataphidini (13/91:10/57) galls on Styrax to Gramineae (esp. Bambuseae),
Palmaceae, Zingiberaceae, where often with ants; oriental
4.2 Hormaphidini (4/11:4/11) galls on Hamamelis to Betula, holarctic
4.3 Nipponaphidini (28/81:28/76) galls on Distylium to Lauraceae, Fagaceae, etc., where
often with ants; oriental
5 Phloeomyzinae (1 ?+) Populus, alate sexuales; all viviparae apterous; holarctic
6 Thelaxinae (3/12:3/12) Betulaceae, Fagaceae, Juglandaceae, often with ants; small
apterous sexuales; holarctic
7 Anoeciinae (2/34:2/15)
7.1 Aiceonini (1/14:1/9) Lauraceae; alate males; Asia
7.2 Anoeciini (1/20:1/6) alternation, Cornus to roots of Gramineae; sexuparae producing
small apterous males
8 Mindarinae (1/5:1/5) Pinaceae; Picea and Abies, few generations per year, holarctic
9 Drepanosiphinae (88/530:74/382) free-living on leaves, mostly on trees; fundatrices often
alate, oviparae apterous, many without ants
9.1 Drepanosiphini (7/45:6/37) Aceraceae; all viviparae alate; holarctic
9.2 Phyllaphidini (72/410:68/345) many on Fagaceae and Betulaceae, some on
Lauraceae, Annonaceae, Magnoliaceae, Ulmaceae, Juglandaceae, Combretaceae,
Burseraceae, Rosaceae, Leguminosae, bamboo; often all viviparae alate
Introduction 5
Table 1. (cont.)
9.3 Saltusaphidini (9/75:0/0) Cyperaceae and Juncaceae; apterous viviparae common;
holarctic
10 Chaitophorinae (11 /159:6/138) free-living on leaves and shoots, apterous viviparae
common, often with ants
10.1 Chaitophorini (6/138:6/138) holarctic
10.1.1 Chaitophorina (3/91:3/91) Salicaceae
10.1.2 Periphyllina (3/47:3/47) Aceraceae
10.2 Siphini (5/21:0/0) Gramineae; holarctic
11 Greenideinae (18/154:13/87) dicot trees, esp. Fagaceae, mostly East Asia
11.1 Greenideini (9/132:7/77) oviparae often alate
11.2 Cervaphidini (9/22:6/10) mostly southern hemisphere; 4 genera in Australia, 1 in
South America
12 Aphidinae (244/2517:59/299) fundatrices and oviparae large, males apterous or alate;
worldwide but mostly holarctic
12.1 Pterocommatini (5/49:5/37) Salicaceae, without host alternation; holarctic
12.2 Aphidini (25/665:14/90) alternation in many genera but many species are
monoecious; often with ants
12.2.1 Aphidina (20/585:9/67) mostly Rosidae and Asteridae
12.2.2 Rhopalosiphina (5/80:5/23) alternation, Rosaceae to Gramineae and
Cyperaceae
12.3 Macrosiphini (214/1803:40/222) alternation in many genera but many species are
monoecious on herbs, often without ants
13 Lachninae (20/361:14/299) fundatrices and oviparae large, males alate or apterous
(sometimes small); often with ants
13.1 Lachnini (9/57:8/50) mostly Fagaceae and Rosaceae; holarctic, esp. central and
eastern palaearctic
13.2 Cinarini (6/269:5/248) Coniferae; holarctic, most numerous in nearctic
13.3 Tramini (5/35:1/1) roots, mostly Compositae; palaearctic, esp. central and eastern
females are therefore termed viviparae, and the sexual females are distin-
guished as oviparae.
4. The more complex life cycles involve host alternation; the technical term
is heteroecy. In heteroecious aphids, the sexuales mate and fertilized eggs are
laid on a tree or shrub, the primary host, but a regular migration occurs
at some stage in the life cycle to another, totally unrelated plant, which may
be herbaceous or woody - the secondary host. On the secondary host, only
parthenogenetic generations (exules) occur, and a return migration to the
primary host is needed before the next sexual generation.
5. Because host alternation has evolved several times independently in
Aphidoidea, there are important differences at the family and subfamily levels
in the way in which it is achieved (see Figs 1-4). It may occur as part of a one-
year cycle (this happens in all Aphidinae, Hormaphidinae, Pemphigini,
Eriosomatini), or the complete cycle may take two years (Adelgidae, Fig. 1;
Fordina, Fig. 2).
6 Introduction
6. The great majority of aphids go through both the sexual and partheno-
genetic phases of their life cycle on one host plant, or on a small range of
closely-related plants. The technical term for this is monoecy. Drepanosi-
phinae, Chaitophorinae, Greenideinae and Lachninae do not have host
alternation; all species in these subfamilies are monoecious. Some examples
of monoecious life cycles are depicted in Figs 5, 6 and 7. Monoecious aphids
generally have fewer morphs, and there are smaller differences between
morphs, than in heteroecious aphids, although there may be considerable
seasonal variation.
7. Some aphids have lost the sexual part of the life cycle; that is, they are
anholocyclic. Some species are entirely anholocyclic and have no known sexual
morphs (e.g. Tuberolachnus salignus, Pineus boerneri), while others may be
anholocyclic in warmer climates and holocyclic in cold temperate regions
(e.g. Eulachnus rileyi). Populations of certain species maintain the options of
both sexual and parthenogenetic reproduction in mild climates, by producing
sexuales while at the same time continuing to produce parthenogenetic females
(e.g. many Greenideinae). Anholocyclic populations of heteroecious aphids
lose their link with the primary host and live all year reproducing partheno-
genetically on secondary host plants.
Introduction 7
Fig. 1. Life cycle of Adelges laricis, an example of the 2-year heteroecious cycle of Adelgidae (partly
based on Carter, 1971). Stippled sector is time spent on primary host. Fundatrices hatch in autumn and
the young larvae (1) overwinter on spruce twigs. They feed at the bud scale bases in early spring (2),
inducing a gall (3) in late May-June, which opens in June-July. The alate gallicolae (4) fly to larch and
lay eggs on the needles (5). The larvae hatching from these eggs, the 'neosistens' stage (6),
overwinters on larch twigs, becoming the adult sistens (plural sistentes) in mid-April (7). The
progrediens larvae (8) which hatch from the eggs of the sistens stage develop on new larch shoots,
becoming either sexuparae (9) or apterous progredientes (9a). The sexuparae migrate to spruce in
May-June and lay eggs which develop as sexual males and females (10). The fertilized eggs laid by
the sexual females develop into fundatrix larvae, completing the holocycle. The apterous progredientes
give rise to one or more summer generations on larch, producing copious wax and honeydew.
8 Introduction
Fig. 2. Life cycle of Baizongia pistaciae, an example of the 2-year heteroecious cycle of Fordina in the
Mediterranean area and southwest Asia (based mainly on Wertheim, 1954). Stippled sector is time
spent on primary host. Fundatrix larvae (1) hatch from eggs on Pistacia in late March-April and feed on
young shoots causing bending (2). They later move to a leaflet which becomes folded and eventually
develops into a large pod-like gall (3). Two generations of apterae occur in the gall before alatae start
to be produced. The gall splits open near its apex in late October-November, releasing large numbers of
emigrant alatae (4), which found colonies of apterous viviparae on roots of grasses (5). Alate sexuparae
(6) are produced by these colonies in March-April and return to Pistacia, where they give birth to very
small sexual females (oviparae) and males (7) on the bark. The oviparae each mature a single egg
which is not laid, the ovipara dying with the egg still inside her. The egg (9) does not hatch until the
following spring.
9
Fig. 3. Life cycle of Thecabius affinis, an example of the life cycle of a heteroecious member of the
Pemphiginae (based on Mordvilko, 1935; Roberti, 1938; Danielsson, 1976). Stippled sector is time
spent on the primary host. In late spring, the fundatrix (1) inhabits a small gall formed by folding one
edge of a poplar (Populus nigra) leaf. Her progeny leave this gall and feed along the mid-rib of another
leaf, causing it to fold in half longitudinally (2), become roughened and blistered on the outer surface
and eventually turn red. The progeny of the fundatrix developing inside this gall all become emigrant
alatae (3), which emerge in June-July and give rise to waxy colonies of apterous exules (4) at stem
bases and on runners of Ranunculus repens. In September, alate sexuparae (5) are produced which fly
to trunks of poplar and give birth to the very small, arostrate males and oviparae (6). (Colonies may
also persist on the secondary host and alate viviparae may be produced that found new colonies on
Ranunculus.) The sexuales develop rapidly to adult and mate on the poplar bark (7). The oviparae each
lay a single egg (8) which does not hatch until the following spring.
10 Introduction
Fig. 4. Life cycle of Aphis fabae, as an example of a heteroecious member of the Aphidinae (after
Blackman, 1974). Stippled sector is time spent on the primary host. The fundatrix (1) develops from an
overwintering egg and founds a colony on new growth of Euonymus europaeus in spring, which gives
rise to emigrant alatae (2) in May-June. Successive generations occur on various herbaceous secondary
hosts through spring and summer (3); alate viviparae are produced as the colonies become larger and
more crowded, and these fly to found fresh colonies on other plants. In autumn, in response to
decreasing daylength, the colonies on the secondary hosts produce alate gynoparae (4) and alate males
(5), which migrate back to Euonymus. The progeny of the alate gynoparae are apterous sexual females,
the oviparae (6), which mature on the spindle leaves just before they fall, and mate with males arriving
independently from secondary hosts. The oviparae lay overwintering eggs (7) in the bud axils.
Introduction 11
Fig. 5. Life cycle of Periphyllus testudinaceus, holocyclic and monoecious on Acer spp., e.g. A.
campestre (based mainly on Essig and Abernathy, 1952). The first instar fundatrix (1) hatches early in
February and develops on swelling buds and young growth of Acer, becoming adult (2) in March. She
has numerous progeny, both apterous and alate. Some of the alatae (3) fly to other maple trees and
produce further generations of apterous (4) and alate (5) viviparae. Other progeny of the fundatrix, both
apterous (6) and alate (7), remain on the same tree; they produce specialized aestivating first instar
nymphs ('dimorphs') with foliate hairs, which spend the summer scattered on both upper and lower
sides of leaves (8). By mid-June, the only individuals to be found are the dimorphs. Eventually, in
September, these start to develop into adult sexuparae, which have a distinct cruciform dark dorsal
patch (9). In late October-November their progeny, small alate males and apterous sexual females
(oviparae), reach maturity and mate (10). Overwintering eggs (11) are laid on branches in bark crevices
and on twigs near winter buds.
12 Introduction
Fig. 7. Life cycle of Cinara schwarzii on Pinus ponderosa in western USA, as an example of the
monoecious, holocyclic annual life cycle of Cinarini (after Palmer, 1926). First instar fundatrices hatch
in late March (1) and feed on the twigs, developing to adult (2) in April. Large, ant-attended colonies of
viviparae, including both apterae (3) and alatae (4), develop on the bark of twigs and small branches.
In some species of Cinara alatae occur only in June-July, but in C. schwarzii they are found from May
through to September. Alate males (5) and apterous oviparae (6) occur in September-October. (Males of
some species of Cinara are apterous, and oviparae often have a conspicuous perianal ring of white
wax.) The shiny black eggs (wax-covered in some species) are laid in rows along the needles (7).
THE TREES AND THEIR APHIDS
14
The Trees and Their Aphids 15
3. The tree genera arranged in alphabetical order with host lists of the aphids
recorded from each tree species, followed by or including keys to the aphids.
The arrangement varies slightly according to the number of aphid species
and their degree of specificity. If numerous aphid species in one genus are
involved - Cinara on pines or Eriosoma on elms for example - then we have
taken advantage of any specificity shown by the aphids to limit the number
of species that have to be discriminated in any one key. The very large aphid
faunas of some tree genera - Quercus and Salix for example - are divided up
in a preliminary 'master' key, to avoid long and cumbersome keys with 100
or more couplets. Species which are keyed on the basis of published descrip-
tions and have not been seen by the authors are indicated by an asterisk (*).
4. This section ends with a key to the 23 species of polyphagous aphids that
may each be found on various tree genera. Many tree genera, particularly
those in the tropics and southern hemisphere, have no aphids specific to them,
but are sometimes colonized by such polyphagous aphids, particularly when
there is vigorous new growth. Most of these aphid species are also found on
herbaceous plants, and some of them are important pests of agricultural crops.
A few of the species included are not strictly polyphagous, but tend to limit
their feeding to both woody and herbaceous members of one plant family; for
example Aphis craccivora on many genera of Leguminosae.
Fig. 8. Morphology of a generalized adult female alate viviparous aphid, with parts named in accordance
with terminology used in keys. In this example the media of the forewing is twice-branched.
Most keys will only work for the particular morph or morphs for which
they were intended. Wherever possible the morph keyed is the adult apterous
vivipara (aptera for short), but there are plenty of exceptions. Many
The Trees and Their Aphids 17
Fig. 9. Measurements frequently required in the keys: A, length of last antennal segment (base and
processus terminalis); B, length of last segment of rostrum (R IV+V); C, length of second segment of
hind tarsus (HT II); D, length of CAUDA.
the identification process. They may give the user a possible name for the
specimens, but no reliance should be placed on this alone. They are based on
existing host plant records, and it is quite possible that aphids will be collected
that are not in the key because they have not previously been recorded from
the plant in question. This is particularly likely in places where the aphid fauna
is less well known. More confidence in the identification may be gained if the
information given in Section III under that name also fits the specimens, but
even then one should proceed with caution; perhaps take the further step of
comparing your aphids with museum specimens identified by a recognized
authority, or consult a more detailed published description. When correct
identification is essential - in biological control work for example - it is
always advisable to consult a taxonomie specialist, who has learnt by experi-
ence many of the pitfalls involved!
Table 2. Tree genera with aphids on them, arranged by families [The nomenclature follows
Brummitt (1992), with some additional subfamily divisions from Willis (1973) and Mabberley
(1987).]___________________________________________
Aceraceae: Acer
Anacardiaceae
Anacardieae: Anacardium, Buchanania, Mangifera
Pistaciaceae (Rhoideae): Harpephyllum, Lannea, Loxostylis, Ozoroa, Pistacia, Rhus, Schinus
Spondieae: Sclerocarya, Spondias
Annonaceae: Annona, Xylopia
Apocynaceae: Apocynum, Carissa, Conopharyngia, Rauwolfia, Tabernaemontana
Aquifoliaceae: Ilex
Araliaceae: Aralia, Cussonia, Eleutherococcus (= Acanthopanax), Kalopanax, Panax, Polyscias,
Pseudopanax, Schefflera
Araucariaceae: Agathis, Araucaria
Asclepiadaceae: Calotropis, Margaretta
Caprifoliaceae: Sambucus
Caryocaraceae: Caryocar
Casuarinaceae: Casuarina
Cecropiaceae: see Moraceae
Celastraceae: Celastrus, Euonymus, Maytenus
Cercidiphyllaceae: Cercidiphyllum
Chrysobalanaceae: Chrysobalanus, Parinaria
Combretaceae: Anogeissus, Combretum, Terminalia
Compositae: Piptocarpha, Vernonia
Cornaceae: Cornus
The Trees and Their Aphids 19
Table 2. (cont.)
Corylaceae: Carpinus, Corylus, Ostrya
Cupressaceae: Callitris, Calocedrus, Chamaecyparis, X Cupressocyparis, Cupressus, Juniperus,
Pilgerodendron, Tetraclinis, Thuja, Widdringtonia
Cyatheaceae: Cyathea (= Alsophila)
Cycadaceae: Cycas
Cyrillaceae: Cyrilla
Labiatae: Hyptis
Lauraceae: Actinodaphne, Cinnamomum, Laurus, Lindera, Litsea, Machilus, Nectandra,
Neolitsea, Ocotea, Persea, Phoebe, Sassafras, Syncarpia, Umbellularia
Lecythidaceae: Barringtonia
Leguminosae
Caesalpinioideae: Brachystegia, Caesalpinia, Cassia, Ceratonia, Cercis, Cynometra,
Dimorphandra, Gleditsia, Julbemardia, Peltophorum, Schotia, Tamarindus
Mimosoideae: Acacia, Albizia, Inga, Leucaena, Mimosa, Pithecellobium, Prosopis
Papilionideae: Aganope (= Ostryoderris), Andira, Castanospermum, Dalbergia, Laburnum,
Lonchocarpus, Maackia, Millettia, Mundulea, Pterocarpus, Robinia, Sesbania,
Sophora, Virgilia
Loganiaceae: Strychnos
Lythraceae: Duabanga, Lagerstroemia
20 The Trees and Their Aphids
Table 2. (cont.)
Magnoliaceae: Liriodendron, Magnolia, Michelia
Malpighiaceae: Banisteriopsis, Malpighia
Malvaceae: Hibiscus, Plagianthus, Thespesia
Meliaceae: Cedrela, Dysoxylum, Khaya, Melia, Toona
Melianthaceae: Bersama
Memecylaceae: Memecylon
Moraceae: Artocarpus, Brosimum, Broussonetia, Cecropia, Cudronia, Ficus, Morus, Streblus
Moringaceae: Moringa
Myricaceae: Myrica
Mysinaceae: Rapanea
Myrtaceae
Leptospermoideae: Eucalyptus, Leptospermum, Melaleuca, Metrosideros, Syncarpia, Tristania
Myrtoideae: Eugenia, Myrtus, Psidium, Syzigium
Sabiaceae: Meliosoma
Salicaceae: Chosenia, Populus, Salix
Sapindaceae: Dodonaea, Koelreuteria, Pappea, Sapindus
Sapotaceae: Malacantha, Mimusops, Pouteria
Scrophulariaceae: Paulownia
The Trees and Their Aphids 21
Table 2. (cont.)
Simaroubaceae: Aiianthus
Sonneratiaceae: Duabanga (see Lythraceae)
Staphyleaceae: Euscaphis, Staphylea
Sterculiaceae: Brachychiton, Dombeya, Guazuma, (Theobroma)
Stilaginaceae: Antidesma
Styracaceae: Sinojackia, Styrax
Symplocaceae: Symplocos
Tamaricaceae: Tamarix
Taxaceae: Taxus
Taxodiaceae: Cryptomeria, Metasequoia, Sequoia, Sequoiadendron, Taxodium
Theaceae: Camellia, Schima, Stewartia
Tiliaceae: Corchorus, Grewia, Microcos, Schoutenia, Tilia
Trochodendraceae: Euptelia, Trochodendron
Xanthophyllaceae: Xanthophyllum
Zamiaceae: Encephalartos
Zygophyllaceae: Guaiacum
About 25 Cinara species are described from Abies, but many of the North
American species are unrecorded since their original descriptions or little
known. Abies is the secondary host for all species of Adelges subgenus
Dreyfusia, which are undoubtedly the most economically important group,
migrating from cone-like galls on their primary host Picea, or reproducing
parthenogenetically on Abies throughout the year.
22 Abies
Elatobium momii
Mindarus abietinus, japonicus
A. forrestii Adelges piceae
A. fraseri Adelges piceae
(Southern Balsam Fir) Mindarus abietinus
A. grandis Adelges nordmannianae, pectinatae,
(Grand Fir) piceae
Cinara confinis, curvipes, gentneri,
occidentalis, pacifica, sonata
Mindarus abietinus, victoria
A. holophylla Cinara longipennis, smaragdina
(Needle Fir) Mindarus japonicus
A. homolepis Cinara todocola
(Nikko Fir) Mindarus abietinus
A. koreana Adeiges pectinatae, piceae
A. kosteri Elatobium abietinum
A. lasiocarpa Adelges pectinatae, piceae
(Subalpine Fir) Cinara chinookiana, confinis,
curvipes, minuta, occidentalis,
osborni
Mindarus abietinus
A. magnifica (incl. var. shastensis) Cinara curvipes, setulosa, sonata
A. mariesii Cinara hattorii
A. mayriana (see A. sachalinensis var. mayriana)
A. nebrodensis Adelges nebrodensis, piceae
Mindarus abietinus
A. nephrolepis Adelges pectinatae
[Prociphilus bumeliaeformis
Mordvilko ex Shaposhnikov,
1955]
Cinara smaragdina
A. nobilis Adelges nordmannianae, piceae
Cinara confinis, curvipes
A. nordmanniana Adelges merken, nordmannianae,
pectinatae, piceae, prelli
Cinara confinis
Elatobium abietinum, sp. nr
momii
Mindarus abietinus
Prociphilus fraxini
24 Abies
Fig. 10. Dorsal (right) and ventral (left) views of adult sistens of A, Pineus abietinus and B, Adeiges
nordmannianae.
1 Adult apterae with very short antennae, less than 0.1 of body length, with
at most 3 segments. Small insects (BL less than 1.2mm) producing white
wax wool. Always oviparous, adults having a distinct chitinous ovipositor
(Adelgidae) .................................................. 2
- Adult apterae with 5- or 6-segmented antennae, at least 0.2 of body length.
BL 1.3-7.8 mm. With or without wax wool. Viviparous in spring and
summer (Aphididae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Body of adult aptera broadly pear-shaped to almost spherical. Cephalo-
prothoracic shield entire, uniformly pigmented. Abdomen with 4 pairs of
evident spiracles. Wax pores discontiguous (Fig. 10A) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus abietinus
- Body of adult aptera pear-shaped or oval. Head and prothorax not
uniformly pigmented. Abdomen with 5 pairs of evident spiracles. Wax
pores contiguous (Fig. 10B) . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges spp.
(see key to resting-stage first instar larvae of Adelges spp. at end of main
key)
3 SIPH tubular. ANT PT/BASE equal to or greater than 1.0 . . . . . . . 4
- SIPH broadly conical, pore-like or absent. ANT PT/BASE less than 1.0
............................................................. 5
4 ANT PT/BASE 1.6 or less . . . . . . . . . . . . . . . . . . . Elatobium abietinum
- ANT PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . Elatobium momii
26 Abies
11 SIPH cones with hairs of two sizes, the longest hairs at least twice as long,
and clearly thicker than, the shortest hairs (Fig. 11B) . . . . . . . . . . . . 12
- SIPH cones with hairs varying in length by less than a factor of 2
(Fig.11A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
12 SIPH cones pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara setulosa
- SIPH cones dark ............................................ 13
13 Pigmented bases of SIPH cones very extensive, with no clear limits,
extending anteriorly and posteriorly to join pigmented lateral areas on
other abdominal segments (Fig. 13A) . . . . . . . . . . . . . . . . Cinara osborni
- Pigmented bases of SIPH cones less extensive, delimited, restricted to
ABD TERG 6, or 5 + 6, any pigmented lateral areas on other segments
being separate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 Length of sclerotized part of stylet groove less than 1.8 mm. ANT V about
equal in length to, or shorter than, ANT VI (including PT) . . . . . . 15
- Length of sclerotized part of stylet groove (Fig. 74, p. 292) greater than
1.8mm. ANT V clearly longer than ANT VI including PT . . . . . . . 17
15 BL greater than 4 mm. Hind tibiae conspicuously bicoloured, with yellow
region on basal half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara alacra*
28 Abies
Fig. 13. SIPH and surrounding sclerites of aptera of A, Cinara osborni and B, Cinara sonata.
- ANT III about equal to or a little shorter than ANT IV and V together,
and less than 3 x ANT VI (incl. PT) . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 R IV over 3 times longer than R V . . . . . . . . . . Cinara abietihabitans*
- R IV less than 3 times longer than R V . . . . . . . . . . . . . . . . . . . . . . . . 22
22 BL 3.8-7.8 mm. ANT PT/BASE less than 0.4 . . . . . . . Cinara confinis
- BL 3.2-4.4 mm. ANT PT/BASE equal to or greater than 0.4. (Alatae with
forewing membrane between Cu1a and Culb fuscous) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara hattorii
23 All tibiae black or very dark brown, although sometimes with a slightly
less dark dusky brown section on basal half . . . . . . . . . . . . . . . . . . . . 24
- Fore and mid-tibiae pale yellow to amber, dark at apices if at all, and hind
tibiae if mainly dark then at least with paler section at base . . . . . . 26
24 BL greater than 4.8mm. Hairs on hind tibiae clearly longer than width
of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara grande*
- BL less than 4.8 mm. Hairs on hind tibiae longer or shorter than width
of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 BL less than 3.2mm. ANT III with up to 5 rhinaria. Hairs on hind
tibiae mainly longer than width of tibia at its midpoint . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara schuhi*
- BL greater than 3.2 mm. ANT III with or without rhinaria. Hairs on hind
tibiae shorter than width of tibia at its midpoint . . . . . . Cinara kiusa*
26 Coxae dark brown to black . . . . . . . . . . . . . . . . . . . . . . . Cinara curvipes
- Coxae yellow to amber . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
30 Abies
27 BL less than 4 mm. Hairs on SIPH cones sparse, totalling about 20, with
basal part of cone nearly devoid of hairs . . . . . . . . Cinara curtihirsuta
- BL more than 5 mm. Hairs on SIPH cones very numerous all over
pigmented area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara gentneri
28 Abdomen without wax pore plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
- Abdomen with segmental wax pore plates, at least marginally .... 30
29 Body oval. Hairs on ANT III longer than basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pacifica*
- Body elongate, about 3 times longer than its greatest width in dorsal view.
Hairs on ANT III shorter than basal diameter of segment . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Essigella pergandei*
30 Antennae of aptera shorter than rostrum. Hairs on antennae and legs
long. (On roots; cannot be keyed to species) . . . . . . . Prociphilus spp.
- Antennae of aptera longer than rostrum. Hairs on antennae and legs
short. (On needles) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31 ANT III of alata with 6-12 squarish to almost circular rhinaria (Fig. 15A)
............................................... Mindarus victoria
- ANT III of alata with 12-27 narrow, tranversely elongate rhinaria ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 ANT III of alata with rhinaria in a single row, mostly extending across
complete width of segment (Fig. 15B) . . . . . . . . . . . Mindarus abietinus
- ANT III of alata with rhinaria not aligned in a single row, many of those
on basal half not extending across full width of segment (Fig. 15C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mindarus japonicus
Fig. 15. ANT III of alata of A, Mindarus victoria, B, M. abietinus and C, M. japonicus.
Fig. 16. Adelges on Abies (neosistens stage except where otherwise indicated). A, dorsal sclerites of
A. pectinatae; B, dorsal sclerites of Adelges (Dreyfusia) sp., showing also the five central areas referred
to in couplets 3 and 4; C, spinal sclerites on meso- and metathorax of A. nordmannianae; D, same for
A. piceae (after Pschorn-Walcher and Zwölfer, 1958); E, second instar sistens of A. prelli (after
Francke-Grosmann, 1937a); F, wax pore plate on spinal sclerite of mesonotum of A. knucheli (after
Schneider-Orelli and Schneider, 1954); 6, thoracic sclerites and wax pores of A. joshii; H, the same for
A. todomatsui (after Inouye, 1953).
the spinal wax pore plates of the meso- and metathorax plus ABD TERG
1-3 (i.e. a total of 10 central areas) is 57-104 .....................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges nordmannianae
- Wax pore plates on inner margins of meso- and metathoracic spinal
sclerites divided into 3-4 angular or rounded areas (Fig. 16D), the most
Abies 33
central area containing 2-8 relatively large pits of irregular shape, rounded
or polygonal. Total number of pits in central areas of spinal wax pore
plates of meso- and metathorax plus ABD TERG 1-3 (i.e. 10 central areas)
is 18-63 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Central areas of wax pore plates on meso- and metathoracic spinal
sclerites usually more-or-less rounded, each with 3-8 pits. Total number
of pits on the 10 central areas (as defined in couplet 3) is in range 26-63
(usually more than 40) . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges merkeri
- Central areas of wax pore plates on meso- and metathoracic spinal
sclerites often more-or-less triangular, each with 3-6 pits. Total number
of pits on the 10 central areas is usually less than 40 (range 18-59)
............................................................. 5
5 Neosistens strongly sclerotized. 2nd instar sistens with a series of dark
conical humps on the head, thorax and 1st to 3rd (or 4th) abdominal
segments, diminishing in size posteriorly (Fig. 16E) . . . . . . Adelges prelli
- Neosistens variably sclerotized, but usually with rather weak sclerotization
(at least in European populations). 2nd instar sistens without pronounced
sclerotic humps on head, thorax and anterior abdominal segments . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges piceae
6 Dorsal wax pore plates of mesothorax, metathorax and abdomen com-
prising groups of 15-25 small rounded facets . . . . . . . Adelges glandulae
- Dorsal wax pore plates either much larger and taking up most of the area
of the sclerite, or much smaller and composed of small groups of no more
than 7 facets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Spinal, pleural and marginal sclerites strongly sclerotized, those of the
mesothorax, metathorax and abdomen each almost fully occupied by
a large wax pore plate composed of 2-5 segments separated by grooves,
each segment having a reticulate structure (Fig. 16F) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges knucheli
- Spinal, pleural and marginal sclerites weakly sclerotized, with wax pore
plates on mesothorax, metathorax and abdomen much smaller, comprising
only 2-7 grouped or loosely-connected facets, and without reticulate
structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Marginal sclerites of meso- and metathorax each with a single wax pore
plate comprising a compact group of 4-7 facets (Fig. 16G) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges joshii
- Marginal sclerites of meso- and metathorax each containing 2 distinct wax
pore plates, each usually with 4 facets (Fig. 16H) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges todomatsui
34 Acacia
ACACIA Leguminosae
ACANTHOPANAX = ELEUTHEROCOCCUS
There are several aphid genera which are virtually specific to Acer, and within
these genera there is a high degree of monophagy, although a few species are
able to colonize several Acer species.
The first key takes adult aphid specimens colonizing any Acer to species,
with the exception of the two largest genera, Periphyllus (35 spp.) and
Drepanaphis (20 spp.). The species of Acer are then listed in alphabetical
order, with a list of the aphids recorded from them followed by keys, where
necessary, to the species of Periphyllus and/or Drepanaphis.
Finally, a key is given to Periphyllus spp. described from unidentified
species of Acer, together with the seven most polyphagous (oligophagous)
species of this genus (acericola, bulgaricus, californiensis, kuwanaii, lyro-
pictus, testudinaceus and villosi).
To identify an Acer-feeding aphid, use the main key first, then if you know
the species of Acer refer to the alphabetical list to see whether your answer is
appropriate, using the subsidiary keys to Periphyllus and Drepanaphis if
necessary. If you have a species of Periphyllus from an unidentified Acer, or
one that does not appear in the alphabetical list, use the final key. The final
key may also help if your Periphyllus does not seem to be one of those
previously recorded from your species of Acer.
Except where otherwise stated, all the keys can be applied to both apterous
and alate viviparous females, but are unlikely to work for fundatrices,
oviparae and males.
Acer 35
Fig. 18. A, ANT III of spring migrant alata of Neoprociphilus aceris; antenna of spring migrant alata of
B, Mimeuria ulmiphila and C, Paraprociphilus tessellatus.
Fig. 19. A, CAUDA of Megalophyllaphis sp.; B, aestivating nymph of Periphyllus americanus; C, SIPH of
aptera of Drepanosiphoniella aceris.
8 ANT VI of spring migrant alata with 4-8 secondary rhinaria (Fig. 18C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraprociphilus tessellatus
- ANT VI of spring migrant alata with 0-1 secondary rhinaria . . . . . . . .
....................................... Paraprociphilus mexicanus
9 SIPH generally stump-shaped, those of aptera with an apical zone of
polygonal reticulation, and those of alata generally with more extensive
reticulation. Antennal hairs usually long and conspicuous, mostly much
longer than diameter of segment from which they arise (except in one
North American species on A. glabrum and one East Asian species on
A. rufinerve). Alatae never with dark markings on wings . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Periphyllus spp. (see separate keys)
- SIPH of variable shape, usually without polygonal reticulation, or if with
some reticulation near apices, then adults are all alatae with dark markings
on wings. Antennal hairs generally small and inconspicuous, usually
shorter than segment from which they arise; if there are long hairs then
they are confined to ANT III (with occasionally one also at the base of
ANT IV) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 CAUDA with an elongate knob, the knob more than twice as long as wide
(Fig. 19A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megalophyllaphis sp(p).
- If CAUDA knobbed then the knob is about as long as wide ..... 11
11 Hairs on front of head and sides of abdomen modified as flattened leaf-
like lamellae (Fig. 19B). (Very small insects, BL less than 1 mm) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . aestivating nymphs of Periphyllus spp.
- Hairs not thus modified . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Front of head and at least the sides of the abdomen with long hairs, those
on frons exceeding length of ANT I . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
38 Acer
Fig. 20. Marginal hairs of mesothorax and metathorax of aptera of A, Trichaitophorus aceris (after
Takahashi, 1937a), B, T. japonicus (after Sorin, 1979b) and C, T. koyaensis.
Fig. 22. A, dorsal abdomen of Drepanosiphum dixoni; B, dorsal abdomen of D. iranicum; C, fore femur
of Drepanosiphum oregonensis; D, SIPH of Shenawheum minutum; E, SIPH of Drepanaphis acerifoliae; F,
forewing of Yamatocallis takagii; 6, forewing of Y. hirayamae; H, forewing of Y. tokyoensis; I, SIPH of
Yamatocallis takagii, J, SIPH of Y. hirayamae.
42 Acer
5 Length of the shorter of the 2 hairs on base of ANT VI less than basal
diameter of segment. Apterae with long hairs on antennae, legs and dor-
sum all blunt or slightly expanded at their apices, and with SIPH at least
1.5 times as long as their basal widths (Fig. 25A) . . . . . . . . . hirticornis
Length of the shorter of the 2 hairs on base of ANT VI greater than basal
diameter of segment. Apterae with long hairs on antennae, legs and
Acer 45
dorsum all fine-pointed, and with SIPH about as long as or a little longer
than their basal widths (Fig. 25B) . . . . . . . . . . . . . . . . . . . . . . . venetianus
6 SIPH pale or only slightly dusky, 1.5-2.0 times longer than their basal
widths ................................................... villosi
SIPH dark and only about as long as their basal widths ... obscurus
A. knowltoni B. idahoensis
C. utahensis D. granovskyi
E. carolinensis F. acerifoliae
G. saccharini H. keshenae
I. sabrinae J. simpsoni
K. parva L. kanzensis
Acer 49
Fig. 26. Drepanaphis: anteriolateral view of spinal tubercles on ABD TERG 1-4 of 12 species.
50 Acer
Aptera with long spinal hairs similarly pigmented on all ABD TERG.
Longer of 2 hairs on ANT BASE VI 0.3-0.8 x length of BASE VI, and
shorter hair 0.12-0.27 x BASE VI . . . . . . . . . . . . . . . . . . . . . . bulgaricus
SIPH of aptera with at least distal half dark. Abdomen of alata with only
faint dorsal markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . coracinus
Paraprociphilus tessellatus
Periphyllus americanus, lyropictus,
testudinaceus
Dorsal body hairs mostly shorter than, or as short as, basal diameter of
ANT III, and with blunt or only very slightly expanded apices. ANT III
with 7-16 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tissoti
ACTINODAPHNE Lauraceae
ACTINOPHLOEUS Palmae
ACTINOSTROBUS Cupressaceae
ADANSONIA Bombacaceae
ADENIUM Apocynaceae
Few aphids are recorded from Aesculus, and only three are possibly specific
to members of this genus.
AGANOPE Leguminosae
Aphis craccivora
ALBIZIA Leguminosae
ALCHORNEA Euphorbiaceae
ALEURITES Euphorbiaceae
Euceraphis ontakensis
Paraprociphilus baicalensis
A. glutinosa Betacallis alnicolens
(European Alder) Clethrobius comes
[Euceraphis betulijaponicae]
Glyphina betulae, schrankiana
Pterocallis alni, maculata
Stomaphis quercus
A. hirsuta Betacallis alnicolens
[Euceraphis betulae, caerulescens]
Hannabura alnicola
Paraprociphilus baicalensis
Pterocallis (Recticallis)
alnijaponicae, nigrostriata
Stomaphis alni
Symydobius alniarius
Tinocallis (Sappocallis) ulmicola
A. incana (N.B. incana of American Betulaphis quadrituberculata
authors = rugosa) Clethrobius comes
(Speckled Alder) Glyphina betulae, schrankiana
Pterocallis albidus, alni
Pterocallis (Recticallis)
alnijaponicae
A. japonica Betacallis alnicolens
Glyphina schrankiana
Hannabura alnicola
Mesocallis alnicola
Pterocallis (Recticallis)
alnijaponicae, nigrostriata
Symydobius alniarius, minutus
[Tinocallis zelkovae]
A. kamtschatica Boernerina alni insularia
Paraprociphilus baicalensis
A. mandshurica Boernerina alni insularia
A. matsumurae Betacallis alnicolens
Boernerina alni
Hannabura alnicola
Mesocallis alnicola
A. maximowiczi Boernerina alni, alni insularia
Euceraphis ontakensis
Paraprociphilus baicalensis
Pterocallis (Recticallis)
alnijaponicae
64 Alnus
Fig. 27. A, proximal part of hind tibia of aptera of Mollitrichosiphon montanum with stridulatory (?)
ridges; B, front of head of alata of Crypturaphis grassii; C, front of head (right side) of aptera of
C. grassii; 0, ornamentation of abdominal tergites of Glyphina schrankiana; E, the same for G. betulae;
f, head of aptera of Latgerina orizabaensis; G, head of alata of L. orizabaensis.
- Body of adult aptera less than 2.5 mm long. Wax pore plates absent ... 12
10 Antennae, legs, anal plate and subgenital plate of aptera all lightly
pigmented, and wax pore plates pale ..... Paraprociphilus baicalensis
- Antennae, legs, anal plate and subgenital plate of aptera brown to black,
and wax pore plates pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 R IV+V of aptera clearly longer than HT II, R IV bearing 4-10 accessory
hairs. Alate sexupara with narrow, transverse secondary rhinaria on
antenna, including 1-5 on base of ANT VI . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraprociphilus tessellatus
- R IV+V of aptera a little shorter than HT II, R IV bearing 2-3 accessory
hairs. Alate sexupara with secondary rhinaria on antennae more oval,
absent from base of ANT VI . . . . . . . . . . . . Paraprociphilus mexicanus
12 Eyes of aptera multifaceted. Dorsum of aptera pale with long capitate
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis albidus
Alnus 67
Fig. 28. A, median dorsal process of ABD TERG 2 of Pterocallis alnifoliae; B, dorsal abdomen of
Recticallis nigrostriata; C, tip of forewing of R. nigrostriata.
- ANT III with 4-9 secondary rhinaria extending over dusky apical half of
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis rhombifoliae
23 SIPH distally black, pale only at their bases . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis (Recticallis) pseudoalni
- SIPH entirely pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Forewings with dark markings at tips between wing veins (Fig. 28C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis (Recticallis) nigrostriata
- Forewings without dark markings between wing veins . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis (Recticallis) alnijaponicae
25 Antennae longer than body . . . . . . . . . . . . . . . . . . . . . Oestlundiella flava
- Antennae shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Antennal flagellum (ANT III-IV) wholly dark except for sensoriated basal
part of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . Boernerina occidentalis
- At least the basal parts of ANT IV and V pale . . . . . . . . . . . . . . . . . 23
27 R IV+V longer than HT II. Front of head usually pigmented. Anal vein
of forewing dark-bordered . . . . . . . . . . . . . . . . . . . . . . Mesocallis pteleae
- R IV+V shorter than HT II. Front of head pale. Anal vein of forewing
pale or dark-bordered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28 ANT PT/BASE about equal to or more than 1.0. Anal vein of forewing
pale. R IV+V 0.5-0.7 x HT II . . . . . . . . . . . . . Mesocallis obtusirostris
Alnus 69
Fig. 29. A, end of abdomen of aptera of Boernerina alni; B, front of head of same; C, groups of
marginal tubercles on ABD TERG 4 and 5 of Symydobius kabae; D, dorsal abdomen of Calaphis alni.
- Aptera with at least some of hairs on ANT III longer than basal diameter
of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
38 Anterior abdominal segments of aptera each with only one capitate
marginal hair on each side . . . . . . . . . . . . . . . . . . . . Mesocallis alnicola*
- Anterior abdominal segments of aptera each with 2 capitate marginal
hairs, arising from a well-developed marginal tubercle . . . . . . . . . . . 39
39 ANT III of alata with 2-8 rhinaria confined to middle part, which is
slightly swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis alnifoliae
- ANT III of alata with 4-9 rhinaria extending over dusky apical half
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis rhombifoliae
Alnus 71
ALSOPHILA = CYATHEA
2 SIPH present as raised pores with rims partly sclerotized, ringed by hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma americanum
SIPH absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus caryae
3 CAUDA triangular, helmet-shaped or rounded, shorter than or about as
long as its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
CAUDA at least 1.5 times longer than its basal width . . . . . . . . . . . . 6
4 SIPH 2-3 times longer than CAUDA . . . . . . . . . . . Dysaphis parasorbi
SIPH about as long as, or a little longer than, CAUDA . . . . . . . . . . 5
5 Dorsum with extensive dark pigmentation. Basal part of hind tibia slightly
swollen, bearing a group of circular 'pseudosensoria' (scent glands)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis sensoriata
Dorsum with pigmented areas restricted to posterior abdominal segments.
Hind tibiae of viviparous morphs without scent glands . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis whiteshellensis
6 SIPH and CAUDA both black. Antennal tubercles weakly developed
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
SIPH pale or dark, but if dark then the CAUDA is paler. Antennal
tubercles well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 ANT III in aptera with about 40 rhinaria, distributed over entire length
of segment. SIPH with an apical zone of polygonal reticulation
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum amelanchiericolens
- ANT III in aptera with 1-2 rhinaria near base . . . . . . . . . . . . . . . . . . . 8
8 SIPH of aptera dark and very long, nearly half as long as body . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon macrosiphum
SIPH of aptera about one quarter of body length, pale except sometimes
at their apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Aptera with antennal tubercles diverging in dorsal view (Fig. 42K). SIPH
wholly pale. Alata with a solid dark dorsal abdominal patch . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbriaphis gentneri
Aptera with antennal tubercles parallel in dorsal view (Fig. 42J). SIPH
often dark-tipped. Alata with dark transverse bars on dorsal abdomen,
sometimes fused across segments in midline only . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
ANACARDIUM Anacardiaceae
ANDIRA Leguminosae
- ANT PT/BASE more than 3.5. Stridulatory apparatus present (Fig. 122)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
6 Lateral (marginal) tubercles present on most abdominal segments ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis sassceri
- Lateral tubercles usually only on ABD TERG 1 and 7 . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
ANOGEISSUS Combretaceae
ANTIDESMA Euphorbiaceae
APHANANTHE Ulmaceae
APOCYNUM Apocynaceae
ARALIA Araliaceae
ARBUTUS Ericaceae
ARECA Palmae
ARENGA Palmae
ARISTOTELIA Elaeocarpaceae
Key to species
SIPH with numerous long hairs. Stridulatory apparatus absent . . . . . .
............................................. Greenidea artocarpi
SIPH without hairs. Stridulatory apparatus present (Fig. 122) . . . . . . . .
.............................................. Toxoptera aurantii
Arundinaria is here used in its older, broad sense, including Pleioblastus and
some other groups now distinguished as separate genera, e.g. Yushania. It
seems to have a more restricted aphid fauna than Bambusa and some species
are specific to it, so a separate, shorter key is provided. However, it is probable
that other bamboo-feeding aphids, not so far recorded from Arundinaria, will
feed on this genus and it may be worthwhile to work through the Bambusa
key if your aphid is not one of those listed and keyed here.
The following aphids are recorded from Arundinaria (and related genera)
Astegopteryx minuta, pallida
Cerataphis bambusifoliae
Ceratovacuna sylvestrii
Chaitoregma tattakana
Cranaphis formosana, [indica]
Melanaphis arundinariae,
bambusae, meghalayensis ssp.
bengalensis, ?pahanensis
Arundinaria 79
Neocranaphis arundinariae
Paracolopha morrisoni
Pseudoregma koshuensis
Rhopalosiphum arundinariae
Sitobion papillatum ssp.
subnudum
Takecallis affinis, arundicolens,
arundinariae, sasae, taiwanus
Fig. 30. A, dorsal (right) and ventral (left) views of aptera of Cerataphis bambusifoliae; B, marginal
wax pore plates on right side of abdomen in Astegopteryx spp., from left to right minuta, pallida,
bambusae, bambucifoliae; C, front of head of aptera of Pseudoregma koshuensis; D, front of head of
aptera of Ceratovacuna sylvestrii; E, marginal tubercles of Chucallis bambusicola; F, wax gland on ABD
TERG 8 of Pseudoregma pendleburyi; G, the same in Ceratovacuna japonica.
wax pore plates. Frontal horns conical, often pointed in adults and always
so in young immatures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 SIPH pores placed on shallow cones encircled with fine hairs. Aptera with
wax pore plates (if present) marginal except on posterior abdominal
segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
82 Bambusa
- SIPH cones small and sclerotic, not encircled by hairs. Aptera with spinal
as well as marginal wax pore plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18 Apices of frontal horns of adult aptera rounded. Marginal wax pore plates
on anterior abdominal segments either small, oval and discontiguous
(Fig. 3OB), or absent . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx minuta
- Apices of frontal horns of aptera pointed. Marginal wax pore plates on
anterior abdominal segments large, mainly oblong, closely opposed to one
another, in rows of 3-7 per segment (Fig. 3OB) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx pallida
19 Dorsal cuticle of fused head and prothorax with denticular ornamenta-
tion. Frontal horns short, curved outwards and narrowed apically, about
as long as ANT I (Fig. 30C) . . . . . . . . . . . . . . Pseudoregma koshuensis
- Dorsal cuticle of fused head and prothorax smooth or slightly wrinkled,
without denticular ornamentation. Frontal horns longer, erect, usually
exceeding length of ANT I (Fig. 3OD) . . . . . . . . Ceratovacuna sylvestrii
Astegopteryx bambusae,
bambucifoliae, basalis, flava,
glandulosa, [insularis],
liukueinensis, [malaccensis],
minuta, [neelagiriensis], pallida,
[salatigensis], [similis],
singaporensis, [striata],
unimaculata, [vandermeermohri]
Cerataphis bambusifoliae
Ceratoglyphina bambusae,
bengalensis, [styracicola]
Ceratovacuna [brevicornis],
floccifera, hoffmanni, indica,
japonica, keduensis, [longifila],
sylvestrii
Chaitoregma [aderuensis], tattakana
Glyphinaphis bambusae
[Indoregma bambusae]
[Pseudoastegopteryx himalayensis],
Pseudoregma albostriata,
bambusicola, [cantonensis],
gombakana, koshuensis,
montana, pendleburyi, [uscare]
Pemphiginae (Fordini) Forda hirsuta
Smynthurodes betae
BANISTERIOPSIS Malpighiaceae
BARRINGTONIA Lecythidaceae
BAUHINIA Leguminosae
BERCHEMIA Rhamnaceae
BERSAMA Melianthaceae
Glyphina betulae
Hamamelistes betulinus
Monaphis antennata
Symydobius kabae, oblongus
B. maximowicziana Betacallis odaiensis
Betulaphis japonica
Calaphis flavabg
Clethrobius comes
Hamamelistes betulinus ssp.
makabiae, cristafoliae
Monaphis antennata
Symydobius kabae
B. middendorfii Betulaphis quadrituberculatabg
Calaphis flavabg
Euceraphis caerulescens
Glyphina betulaebg
Monaphis antennatabg
B. nana Betulaphis pelei
(European Dwarf Birch) Calaphis arctica, betulicola, ?flava
Euceraphis sp. near betulae
Symydobius sp.
B. nigra Calaphis betulella, neobetulella
(River Birch) Callipterinella tuberculata
Hamamelistes spinosus
Hormaphis hamamelidis
Pemphigus ephemeratus
B. occidentalis (=fontinalis) Betulaphis quadrituberculata
(Western Black Birch) Calaphis betulaecolens,
betulaefoliae, coloradoensis,
flava
Euceraphis sp. near betulae
Symydobius intermedius
B. papyrifera Calaphis betulaecolens,
(Paper-bark Birch) betulaefoliae, flava, viridipallida
Callipterinella callipterus,
minutissima
Euceraphis sp. near betulae,
punctipennisbg
Glyphina setosa
Hamamelistes spinosus
Hormaphis hamamelidis
Monaphis antennata
Symydobius americana
92 Betula
4 Antenna of aptera 3-segmented, ANT III thin, at its base less than half
the width of ANT I, and tapering gradually from base towards apex
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamamelistes betulinus
- Antenna of aptera usually 4-segmented, sometimes 5-segmented, with
ANT III at its base at least half the width of ANT I . . . . . . . . . . . . . 5
5 Aptera (from gall) with pigmented siphuncular pores . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamamelistes gibberi
- Aptera (from gall) without siphuncular pores . . . . . . . . . . . . . . . . . . . . 6
6 Galls comprising reddish to crimson corrugations of leaf lamina between
veins. (Apterae in life dark brownish to purple with white wax) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamamelistes spinosus
- Galls comprising cockscomb-like bags protruding on upperside of leaf.
(Apterae reddish-yellow in life) . . . . . . . . . . . Hamamelistes cristafoliae
7 ANT PT/BASE less than or about equal to 1.0 . . . . . . . . . . . . . . . . . . 8
- ANT PT/BASE clearly more than 1.0, and usually more than 1.5 (the
single exception is one species with very small apterae, Callipterinella
minutissima, which has PT/BASE in range 1.1-1.8) . . . . . . . . . . . . . 37
8 Rostrum longer than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Rostrum shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
9 HT II shorter than maximum diameter of SIPH cone . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis radicicola
- HT II longer than maximum diameter of SIPH cone . . . . . . . . . . . . 10
10 HT II more than 2 x longer than 2nd segments of fore and mid-tarsi
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis takahashii
- HT II less than 1.5 x longer than 2nd segments of fore and mid-tarsi
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis quercus
11 ANT PT peg-like, less than 2 x its basal width. Eyes of aptera reduced
to 3 facets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- PT of antenna finger-like, more than 2 x its basal width. If apterae are
present then their eyes are multifaceted . . . . . . . . . . . . . . . . . . . . . . . . 15
12 Dorsum pale. Body hairs inconspicuous. SIPH absent . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus ephemeratus
- Dorsum pigmented, hairs conspicuous. SIPH present as pores on small
cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Dorsal and lateral body hairs of aptera long and fine . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glyphina setosa
- Dorsal and lateral body hairs of aptera thick and spine-like . . . . . . 14
Betula 95
Fig. 31. Hairs on ABD TERG 2 and 3 of apterae of A, Betulaphis quadrituberculata (longer-haired form,
B, B. brevipilosa, C, B. pelei, D, B. quadrituberculata (shorter-haired form).
24 Aptera with spinal and pleural hairs on ABD TERG 7 as short and
inconspicuous as those on more anterior tergites. Marginal hairs on thorax
and ABD TERG 1-4 also very short. Dorsal abdominal hairs only long
and capitate on ABD TERG 8, and lateral hairs only long and capitate
on ABD TERG 5-8. Dorsal cuticle very wrinkled (Fig. 3IB) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Betulaphis brevipilosa
- Aptera with at least some of spinal and pleural hairs on ABD TERG 5-7
long and capitate, in addition to marginal hairs and those on ABD TERG
8. Dorsal cuticle smooth or somewhat wrinkled . . . . . . . . . . . . . . . . . 25
25 R IV+V only about 0.06mm long, about as long as its basal width, less
than 0.6 x ANT VI BASE . . . . . . . . . . . . . . . . . . . . Betulaphis japonica
- R IV+V more than 0.07 mm long, more than 0.7 x ANT VI BASE
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Aptera with marginal hairs on ABD TERG 1-3 shorter than or about as
long as basal diameter of ANT III (Fig. 31C) . . . . . . . Betulaphis pelei
- Aptera with marginal hairs on ABD TERG 1-3 mostly much longer than
basal diameter of ANT III (Fig. 31D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . Betulaphis quadrituberculata (short-haired form)
Betula 97
41 Abdomen with separate dark dorsal transverse bars on all tergites ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callipterinella calliptera
- Abdomen with a large dark quadrate patch on ABD TERG 4 to 6
....................................... Callipterinella tuberculata
42 Head with a black transverse band across the venter between the eyes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
- Head without a ventral transverse band . . . . . . . . . . . . . . . . . . . . . . . . 46
43 Marginal tubercles on ABD TERG 2-5 or 4-5, and whole of SIPH,
pigmented. R IV+V less than 1.2 x HT II . . . . . . . . . . . . . . . . . . . . . . 44
- Marginal tubercles on ABD TERG 2-4 pale, those on 5 either pale or
pigmented. R IV+V at least 1.25 x HT II . . . . . . . . . . . . . . . . . . . . . . 45
44 Head and prothorax with a dark dorsal longitudinal median stripe.
Forewing veins evenly pigmented . . . . . . . . . . . . . Betacallis querciphaga
- Head and prothorax without a dorsal longitudinal median stripe. Media
of forewing pale, but Cu1b and distal part of Cu la thickly bordered with
fuscous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Betacallis odaiensis
45 ANT III with 17-25 secondary rhinaria, on basal 0.5-0.7 of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Betacallis sikkimensis
- ANT III with 37-45 secondary rhinaria, extending more than 0.75 of
length of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . Betacallis luminiferus
46 Alata with a distinctive pattern of dark markings dorsally; longitudinal
stripes on head and thorax, and transverse bars or patches on abdomen
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
- Alata without a distinctive pattern of dark dorsal markings . . . . . . 48
47 Dorsal abdomen with separate double black bars on ABD TERG 4
and 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis betulella
- Dorsal abdomen with a large black quadrate patch on ABD TERG 4
and 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis neobetulella
48 Knob of CAUDA elongate, longer than wide . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis betulaefoliae
- Knob of CAUDA globular, no longer than its maximum width ... 49
49 Apterous morph . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
- Alate morph ................................................ 55
50 Most proximal secondary rhinarium on ANT III located near base of
segment, no further from base of segment than the length of ANT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
100 Betula
60 R IV+V 0.09-0.105 mm, less than 0.8 x HT II, and bearing only 2-4
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis viridipallida
- R IV+V at least 0.11 mm long, more than 0.8 x HT II, and bearing 4-8
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis flava
BISCHOFIA Euphorbiaceae
BIXA Bixaceae
BOMBAX Bombacaceae
BRACHYCHITON Sterculiaceae
BRACHYSTEGIA Leguminosae
BRIDELIA Euphorbiaceae
BROSIMUM Moraceae
BROUSSONETIA Moraceae
BUCHANIA Anacardiaceae
BUDDLEJA Buddlejaceae
BURSERA Burseraceae
BUTEA Leguminosae
BUXUS Buxaceae
CAESALPINIA Leguminosae
Aphis spiraecola
Myzus persicae
(Use key to polyphagous aphids, p. 532)
CALAMUS Palmae
Cerataphis ?variabilis
(For palm aphids see B & E, 1984)
CALLIGONUM Polygonaceae
CALLITRIS Cupressaceae
CALOCEDRUS Cupressaceae
CALODENDRUM Rutaceae
CALOPHYLLUM Guttiferae
CALOTROPIS Asclepiadaceae
CAMELLIA Theaceae
CANARIUM Burseraceae
CARISSA Apocynaceae
The aphids of pecan (Carya illinoensis) were keyed in Blackman and Eastop
(1984). The present key is extended to cover other Carya species, but there
are certain difficulties. The hickory phylloxerids (Phylloxera spp.), with more
than 30 available names, are impossible to key in the present state of know-
ledge. Pergande (1904) provided a classification and keys based on the galls.
Many species of the three North American genera Monellia, Monelliopsis and
Protopterocallis are morphologically similar, yet most are recorded from
several Carya species, so a single key is necessary with some rather difficult
and possibly unreliable discriminants within these genera.
thorax, and mid- and hind femora, all very dark, as are the well-developed
spinal and marginal abdominal tubercles ..... Melanocallis caryaefoliae
- SIPH pale and often inconspicuous, small, tuberculate. Head thorax and
all femora mainly pale or dusky, abdominal tubercles variably developed
and/or pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Forewing vein Cu1b ('anal' vein) darker than vein Cu1a, lightly bordered
with fuscous along most of its length. Rhinaria-bearing part of ANT III
only slightly thickened, rarely up to 1.5 x diameter of basal articulation
of segment (Fig. 33B). Embryos (seen through adult cuticle) with spinal
hairs on ABD TERG 5 displaced laterally, and spinal hairs on ABD TERG
6 displaced medially (Figs 34A, B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Forewing vein Cu lb as pale as Cu1a (except sometimes at ends). Rhinaria-
bearing part of ANT III conspicuously swollen, usually more than
1.5 x diameter of basal articulation of segment (Fig. 33C). Embryos with
spinal hairs on ABD TERG 1-6 in two longitudinal rows without any
lateral or medial displacement (Figs 34C-E) . . . . . . . . . . . . . . . . . . . . . 14
6 ANT PT/BASE 0.6-1.1. Small, delicate aphids, BL rarely more than
2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- ANT PT/BASE 0.3-0.6. More robust aphids, BL usually more than 2 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
7 ABD TERG 1-6 each with only one pair of spinal hairs, sometimes arising
from tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- ABD TERG 1-6 each with at least 4 spinopleural hairs, either as separate
spinal and pleural pairs, or with spinal hairs duplicated on flat tubercles
(bosses) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 Pronotum, metanotum and ABD TERG 1-3 each with a pair of well-
developed tubercles, much longer than the flattened, pigmented bosses on
ABD TERG 4-6 (Fig. 33D). Embryos (seen through adult cuticle) with
spinal hairs on ABD TERG 2-4 long enough to overlap when directed
towards one another (Fig. 34A) . . . . . . . . . . . . Monelliopsis tuberculata
- Pronotum and metanotum without tubercles, and ABD TERG 1-3 with
flattened hair-bearing bosses like those on ABD TERG 4-6 (Fig. 33E).
Embryos with spinal hairs on ABD TERG 2-4 too short to overlap when
directed towards one another (Fig. 34B) . . . . . . . . . Monelliopsis caryae
Fig. 33. A, SIPH of Melanocallis caryaefoliae; B, ANT III of Monelliopsis caryae; C, ANT III of Monellia
caryella; D, tubercular bases of spinal hairs on ABD TERG 1-6 of Monelliopsis tuberculata; E, the same
for M. caryae; F, position of dark spot on hind femur of Monelliopsis bisetosa; G, the same for
M. nigropunctata; H, tubercular bases of spinal hairs on ABD TERG 2-4 of Monelliopsis nigropunctata;
I, the same for M. pecanis; J, hair-bearing tubercles on prothorax of Protopterocallis quadrata; K, the
same for P. pergandei; L, the same for P. gigantea; M, hair-bearing tubercles on ABD TERG 1 and 2 of
P. gigantea.
114 Carya
Carya 115
9 ABD TERG 1-6 without pleural hairs, but with 2 spinal hairs on each
spinal boss. Hind femur with a black spot dorsally near apex (Fig. 33F)
............................................ Monelliopsis bisetosa
- ABD TERG 1-6 with separate spinal and pleural hairs on separate bosses.
Hind femur with a black spot on ventral side near apex (Fig. 33G) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Spinal tubercles (bosses) on ABD TERG 2-4 clearly larger than pleural
bosses (Fig. 33H) . . . . . . . . . . . . . . . . . . . . . . Monelliopsis nigropunctata
- Spinal bosses on ABD TERG 2-4 of similar size to pleural bosses
(Fig. 33I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monelliopsis pecanis
11 Pronotum with 2 posterior tubercles (Fig. 33J), and ABD TERG 1-4 each
with 4 tubercles . . . . . . . . . . . . . . . . . . . . . . . . . Protopterocallis quadrata
- Pronotum usually with at least 4 posterior tubercles (Figs 33K, L), and
ABD TERG 1-4 each usually with more than 4 tubercles . . . . . . . . 12
12 Pronotum with 6-7 posterior tubercles, in groups of 2-4 (Fig. 33K)
....................................... Protopterocallis pergandei
- Pronotum with 4-5 posterior tubercles (Fig. 33L) . . . . . . . . . . . . . . . . 13
13 Some of spinal hairs on ABD TERG 1-3 neotenic; as long as or longer
than the tubercles from which they arise, 45-80 µm long, expanded at tips
and very stout-based (Fig. 33M) . . . . . . . . . . . Protopterocallis gigantea
- Spinal hairs on ABD TERG 1-3 all shorter than the tubercles from which
they arise, less than 40 µm long, similar in size to hairs on pronotum
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Protopterocallis fumipenella
14 ABD TERG 1-6 each with 3-8 fine, often very short hairs, with small
round pigmented spots at their bases, which are never tuberculate ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monellia microsetosa
- ABD TERG 1-6 each with only 2 spinal hairs, often on low tubercles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Spinal hairs on ABD TERG 1-4 all short and fine, less than 40 µm
long, less than 1.5 x diameter of most swollen part of ANT III. ABD
TERG 1-6 of embryos (seen through adult cuticle) with very short spinal
hairs and long overlapping marginal hairs (Fig. 34C) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monellia caryella
- At least some of spinal hairs on ABD TERG 1-4 long, 40-80 µm,
1.5-3.2 x diameter of most swollen part of ANT III. If ABD TERG 1-6
of embryos have very short spinal hairs (Monellia hispida) then the
marginal hairs do not overlap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Embryos (and 1st instars) with marginal hairs on abdominal segments
1-6 short, not overlapping between segments, and spinal hairs minute
(Fig. 34D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monellia hispida
116 Caryocar
- Embryos (and 1st instars) with marginal and spinal hairs all long, the
marginal hairs all overlapping between segments (Fig. 34E) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monellia medina
CARYOCAR Caryocaraceae
CASEARIA Flacourtiaceae
CASSIA Leguminosae
2 Many dorsal tubercles present on head, thorax and ABD TERG 1-6,
decreasing in size and number posteriorly . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moritziella castaneivora
- Only 4 dorsal tubercles present, confined to head . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera castaneae
3 SIPH tubular, several times longer than w i d e . . . . . . . . . . . . . . . . . . . . . 4
- SIPH truncate or broad, hairy cones . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
4 Body of aptera with very long, branched, hair-bearing lateral processes
(Fig. 92A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cervaphis quercus
- Body of aptera without hair-bearing processes . . . . . . . . . . . . . . . . . . . 5
5 SIPH without hairs. Antennal tubercles well developed, approx. parallel-
sided in dorsal view (Fig. 42J) . . . . . . . . . . . . . Aulacorthum magnoliae
- SIPH with numerous long fine hairs. Antennal tubercles undeveloped
............................................................. 6
6 Abdomen of aptera broadly rounded, uniformly brown pigmented. SIPH
of aptera with a heavily sclerotized, reticulate zone at the bases, and
CAUDA with a distinct median conical papilla (as in Fig. 92B) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea (Trichosiphum) kuwanai
- Abdomen of aptera oval, less pigmented, at least with a pale anterio-
median area. SIPH of aptera without basal reticulation, and CAUDA
without a median papilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 SIPH of aptera more than 0.7 x BL. Hind tibiae with transverse
stridulatory ridges (as in Fig. 27A) .... Mollitrichosiphon tenuicorpus
- SIPH of aptera less than 0.7 x BL. Hind tibiae without stridulatory ridges
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum tattakanum
8 ANT PT/BASE about 1.0 or more. Anal plate divided by a median cleft
into 2 lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- ANT PT/BASE 0.5 or less. Anal plate not bilobed . . . . . . . . . . . . . . 21
9 CAUDA triangular, apically pointed, without a constriction (Fig. 35A).
Body and appendages almost entirely brown-black . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patchia virginiana
- CAUDA with a constriction and a terminal knob. Body and appendages
mainly pale, sometimes with dark markings . . . . . . . . . . . . . . . . . . . . . 10
10 Head and pronotum with a longitudinal dark spinal stripe, and ABD
TERG 1-7 each with paired dark dorsal spinal as well as marginal sclerites
(Fig. 35E). (Wings not maculate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Head and pronotum without a spinal longitudinal stripe. If ABD TERG
1-7 have a complete set of paired dark dorsal sclerites then wings are
strongly maculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Castanea 119
Fig. 35. A, CAUDA oi Patchia virginiana; B, front of head and ANT I-II of Castaneomyzocallis
castaneae; C, the same for C. nanae; D, the same for C. castaneoides; E, dorsal body markings of
Myzocallis castanicola; F, marginal tubercles of ABD TERG 3-6 and SIPH of Castanocallis cereus;
6, spinal tubercles on ABD TERG 2 of Castanocallis castanocallis; H, the same for Myzocallis kuricola;
I, the same for Tuberculatus capitatus.
11 R IV with 4-10 accessory hairs. First tarsal segments with 5-6 ventral
hairs. Embryos with long hairs (55-70 µm) on ABD TERG 2 . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis castanicola castanicola
- R IV with 11-18 accessory hairs. First tarsal segments with 6-8 (usually
7) ventral hairs. Embryos with hairs on ABD TERG 2 shorter (27-41 µm)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis castanicola leclanti
12 Tibiae mainly brown to black, or at least brown on basal half. ANT
PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Tibiae mainly pale, sometimes dark apically. ANT PT/BASE less than 2.0
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
120 Castanea
13 Anterior frontal hairs of head long and finely pointed, 2 or more X basal
diameter of ANT III (Fig. 35B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Anterior frontal hairs less than 2 x basal diameter of ANT III, and
sometimes with capitate apices (Figs 35C, D) . . . . . . . . . . . . . . . . . . . . 15
14 Tibiae and ANT III-VI uniformly black . . . . . . . Myzocallis castaneae
- Tibiae dark at base but with paler middle region. ANT IV, V and BASE
VI each pale at base and dark at apex . . . . . . . . . . . . Myzocallis tissoti
15 Anterior frontal hairs 7-15 µm long, shorter than basal diameter of ANT
III, and capitate (Fig. 35C). ANT III-VI pale basally and dark apically,
and tibiae usually with paler middle region . . . . . . . . Myzocallis nanae
- Anterior frontal hairs 30-38 µm, longer than basal diameter of ANT III,
pointed or slightly capitate (Fig. 35D). ANT III-VI and tibiae often
completely dark . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis castaneoides
16 Forewings with pigment extending between veins, or with veins at least
heavily bordered with pigment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Forewings not pigmented between veins . . . . . . . . . . . . . . . . . . . . . . . . 20
17 ANT III with 4-6 hairs, maximally about 1.5 x basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castanocallis margituberculatus
- ANT III with 7 or more hairs, including anteriorly directed long ones more
than 2 x basal diameter of the segment . . . . . . . . . . . . . . . . . . . . . . . . 18
18 ANT III with more than 20 hairs. Marginal tubercles each with 4-6
transparent noduli (Fig. 35F) . . . . . . . . . . . . . . . . . . Castanocallis cereus
- ANT III with less than 15 hairs. Marginal tubercles without any
transparent noduli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 ABD TERG 2 and 3 each with a pair of quadrate spinal tubercles, as long
as or longer than their basal widths (Fig. 35G). ANT III longer than head
width across (and including) eyes . . . . . . . . . Castanocallis Castanocallis
- Spinal tubercles on ABD TERG 2 and 3 if developed then rounded, and
shorter than their basal widths (Fig. 35H). ANT III shorter than head
width across eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis kuricola
20 Spinal tubercles on ABD TERG 1-3 all similarly developed, finger-like,
longer than their basal widths (Fig. 35I). Anteriorly directed hairs on front
of head and ANT I-III long and capitate. SIPH pale . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus capitatus
- Spinal tubercles only ever well developed on ABD TERG 3, and then
conical, shorter than their basal widths. Hairs on front of head and ANT
I-III short or very short, and pointed. SIPH black except at bases
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus annulatus
21 BL less than 2 mm. Antennae 5-segmented. CAUDA knobbed . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelaxes suberi
Castanopsis 121
CASTANOPSIS Fagaceae
E
C
Fig. 36. A, margin of prosoma of aptera of Reticulaphis shiiae; B, dorsal view of aptera of
Neonipponaphis shiiae, with detail of cuticular ornamentation of prosoma; C, the same for
Metanipponaphis sp.; D, cuticular ornamentation of prosoma of aptera of Lithoaphis shiiae; E, the same
for Nipponaphis brevilosa.
CASTANOSPERMUM Leguminosae
CASUARINA Casuarinaceae
CATALPA Bignoniaceae
CECROPIA Cecropiaceae
CEDRELA Meliaceae
Fig. 37.A, dorsal hair of aptera of Cinara (Cedrobium) laportei; B, HT I of Cinara cedri.
CELTIS Ulmaceae
Fig. 38. A, posterior part of abdomen (right side) of Sumatraphis celti; B, CAUDA and anal plate of
Shivaphis catalpinari; C, the same for Sh. celti; D, SIPH of Sh. hangzhouensis; E, SIPH of Sh. celticola;
f, SIPH of Sh. szelegiewiczi (after Quednau, 1979).
- (Alata from leaf-nest.) BL more than 4mm. Hind wing with 2 oblique
veins arising from a common base. Secondary rhinaria on ANT III-V (not
VI), narrowly transverse but not extending more than half-way around
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
4 BL more than 4 mm. Rostrum longer than body ... Stomaphis yanonis
- BL less than 3 mm. Rostrum shorter than body . . . . . . . . . . . . . . . . . . 5
5 Antenna 4-segmented in aptera, 5-segmented in alata. Posterior abdomen
with lateral processes. SIPH tubular, swollen, constricted apically, and
with a subapical ring of 4 hairs (Fig. 38A) . . . . . . . . Sumatraphis celti
- Antenna 6-segmented. Abdomen without lateral processes. SIPH are
pores, very small cones, or truncate, without hairs . . . . . . . . . . . . . . . 6
130 Celtis
CERATONIA Leguminosae
CERCIDIPHYLLUM Cercidiphyllaceae
CERCIS Leguminosae
CHAMAECYPARIS Cupressaceae
3 BL often more than 2.4mm. Aptera with dark sclerites on thorax and
ABD TERG 8. Apices of tibiae dark. ANT VI BASE with 8-14 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara (Cupressobiurri) tujafilina
BL less than 2.4 mm. Aptera without dark dorsal sclerites. Apices of tibiae
pale or dusky. ANT VI BASE with 5-6 hairs . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara (Cupressobium) louisianensis
N.B. Other aphids whose more normal hosts are Cedrus and Cupressus may
occasionally colonize Chamaecyparis, so it may be advisable to look also at
the keys to aphids on those plant genera.
CHORISIA Bombacaceae
CHOSENIA =SALIX
CHRYSOBALANUS Chrysobalanaceae
CINCHONA Rubiaceae
Fig. 39. A, dorsal view of body of a Euthoracaphis; B, prosomal hairs of Eu. heterotricha; C, prosomal
hairs of Eu. umbellulariae.
CITHAREXYLUM Verbenaceae
CITRUS Rutaceae
COCOS Palmae
COMBRETUM Combretaceae
Apart from a few polyphagous species, the aphid fauna of Combretaceae and
Burseraceae consists entirely of members of the genus Paoliella. Species
of Paoliella are specific at least at the level of host genera (Combretum,
Commiphora, Terminalia), with eight known from Combretum. Three of
these are only known as alate morphs, and one only from a single aptera, so
apterae and alatae are keyed separately.
Fig. 40. Typical spinal processes of Paoliella spp.: A, papillata (aptera), B, papillata (alata),
C, chiangae (alata), D, longirostris (alata), E, delottoi (aptera or alata), F, commiphorae (alata),
G, ufuasi (aptera), H, hystrix (aptera), I, echinata (aptera), J, ayari (aptera), K, mesothoracic process
of ayari (alata).
COMMIPHORA Burseraceae
This genus has a similar aphid fauna to Combretum, q.v., but with different
species of Paoliella.
1-3 finger-like, the longest at least 2 x their basal widths. Radial sector
and media not bordered distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Mesothoracic dorsal processes dark and heavily spiculose (Fig. 40K).
Forewing vein Cu1b, bordered with fuscous . . . . . . . . . . . Paoliella ayari
Mesothoracic dorsal processes pale and no more spiculose than those on
other tergites. Forewing vein Culb not bordered ..... Paoliella ufuasi
CONOPHARYNGIA = TABERNAEMONTANA
CORCHORUS Tiliaceae
CORDIA Boraginaceae
Cornus is the primary host of the genus Anoecia, and the primary or only host
of seven species of Aphis. In both these genera some species are difficult to
separate, and identifications made with the following key need to be treated
Cornus 141
with caution. The North American Cornus-feeding species of the genus Aphis
were reviewed by Robinson and Chen (1969). For Anoecia only the alatae (i.e.
the spring migrant fundatrigeniae leaving Cornus in spring, and the sexuparae
returning to it in autumn) are keyed, as in several species the only apterous
morphs on Cornus are the fundatrix in spring and the sexuales in autumn.
Fig. 41. SIPH and marginal tubercles on ABD TERG 5-7 of apterae of A, Anoecia corni alate sexupara,
B, Pseudosiphonaphis corni aptera and C, Aphis caliginosa aptera; SIPH of D, Aphis cornifoliae,
E, A. neogillettei and F, A. salicariae.
Aulacorthum solani
Macrosiphum euphorbiae,
hamiltoni, manitobensis
Myzus persicae
Cornus sp. Prociphilus cornifoliae
12 Hairs on ANT III more than 2 x longer than basal diameter of segment.
SIPH clearly shorter than CAUDA. Closely-spaced spinulose ridges
present on ABD sternites 5 and 6 . . . . . . . . . . . . . . . . Toxoptera odinae
- Hairs on ANT III less than 2 x longer than basal diameter of segment.
SIPH usually as long as or longer than CAUDA. No evident spinulose
ridges on ABD sternites 5 and 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Hairs on ANT III short, maximally 35 µm, shorter than or about equal
to middle diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Hairs on ANT III long, maximally 35-75 µm, all or most clearly exceeding
middle diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
14 Marginal tubercles present on prothorax and on ABD TERG 2-5 as well
as 1 and 7; those on prothorax, 1 and 7 being exceptionally large and
broad-based (Fig. 41C) . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis caliginosa
- Marginal tubercles usually present only on prothorax and ABD TERG 1
and 7, and these not unusually large . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 CAUDA finger-like, with a slight constriction basad of midpoint, the
distal part elongate oval (Fig. 121T) . . . . . . . . . . . . . . . Aphis spiraecola
- CAUDA tongue-shaped or bluntly triangular, without elongate oval distal
part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Aptera with ANT III usually more than 1.3 x PT (except in very small
specimens). Alata with 15-36 secondary rhinaria on ANT III, and usually
none (sometimes 1-2) on ANT IV . . . . . . . . . . . . . . . . . . Aphis helianthi
- Aptera with ANT III usually less than 1.3 x PT. Alata with 3-16
secondary rhinaria on ANT III, and 0-5 on ANT IV . . . . . . . . . . . . 17
17 Hind femora, SIPH and CAUDA all uniformly dark. SIPH cylindrical,
only a little narrower at apex than at base, and often curved outwards
(Fig. 41D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis cornifoliae
- Hind femora pale (aptera) or pale on basal third (alata); SIPH dark,
CAUDA pale or dusky. SIPH tapering, much narrower at apex than at
base (especially in aptera), and usually rather straight . Aphis gossypii
18 Tibiae of cleared specimens uniformly brown-black .................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis nigratibialis
- Tibiae of cleared specimens pale, or dark only apically . . . . . . . . . . 19
19 SIPH long, about 0.25 of BL. Aptera with 15-22 secondary rhinaria on
ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis maculatae
- SIPH much less than 0.25 of BL. ANT III of aptera usually without
secondary rhinaria (rarely 1-4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 CAUDA with 11-27 hairs. Dorsal hairs on ABD TERG 1-6 usually less
than 40 µm long. SIPH usually rather straight . . . . . . . . . . Aphis fabae
Cornus 145
- CAUDA with 6-11 hairs. Dorsal hairs on ABD TERG 1-6 usually long,
more than 40µm. SIPH often curved outwards distally (Figs 41E, F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 ABD TERG 8 with 3-6 hairs. SIPH of aptera about 5 x longer than their
width at base (Fig. 41E). Alata without dark dorsal markings on ABD
TERG 1-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis neogillettei
- ABD TERG 8 with 6-12 hairs. SIPH of aptera only 2-3 x longer than
their width at base (Fig. 41F). Alata with dark bars or sclerotic patches
on ABD TERG 1-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis salicariae
22 Antennal tubercles with inner faces divergent in dorsal view (Fig. 42K).
SIPH with a subapical zone of polygonal reticulation . . . . . . . . . . . . 23
- Antennal tubercles with inner faces parallel or convergent in dorsal view
(Figs 42I, J). SIPH without subapical polygonal reticulation . . . . . . . 26
23 CAUDA long and finger-like, 0.12-0.20 of BL. Hairs on ANT III more
than 0.5 of middle diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . 24
- CAUDA short, less than 0.11 of BL. Hairs on ANT III less than 0.5 of
middle diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
24 SIPH 1.5-2.1 x CAUDA and reticulated over distal 13-25%. R IV+V
0.9-1.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
- SIPH 2.3-2.5 x CAUDA and reticulated over distal 6-11%. R IV+V
1.05-1.20 x HT II . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum hamiltoni
25 ANT BASE VI less than 0.2 mm, much shorter than ANT I + II measured
together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum manitobensis
- ANT BASE VI about 0.3 mm, as long as ANT I + II measured together
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion cornifoliae
26 Antennal tubercles with inner faces approximately parallel in dorsal view
(Fig. 42J). ANT III of aptera usually with a single rhinarium near base.
SIPH straight, tapering from base to flange (Fig. 121K) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
- Antennal tubercles with inner faces convergent in dorsal view (Fig. 42I).
ANT III of aptera never with any rhinaria. SIPH tapering with a slight
S-curve, or slightly swollen on distal half . . . . . . . . . . . . . . . . . . . . . . . 27
27 ANT PT/BASE less than 2.5. Aptera with a dorsal intersegmental pattern
of dark ornamentation (Fig. 121J). SIPH coarsely imbricated, with a
slight S-curve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus ornatus
- ANT PT/BASE more than 3.0. Aptera without any clear dorsal markings.
SIPH slightly swollen on distal half (Fig. 121H) . . . . . . Myzus persicae
146 Corylopsis
CORYLOPSIS Hamamelidaceae
8 ANT III dark, with 7-14 rhinaria. Abdominal marginal tubercles each
bearing a single small hair . . . . . . . . . . . . . . . . . . . . . . Mesocallis pteleae
- ANT III pale or only dark at apex, with 2-11 rhinaria. Abdominal
marginal tubercles mostly with 2-3 hairs . . . . . . . . . . . . . . . . . . . . . . . . 9
9 ANT PT/BASE 0.5-0.9. ABD TERG 1-7 with 6-10 spinopleural hairs,
the spinal hairs often grouped on paired tubercles . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis affinis
- ANT PT/BASE 0.9-1.2. ABD TERG 1-7 each with a single pair of spinal
hairs only, those on 3, 5 and 7 being displaced laterally . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis (Paratinocallis) corylicola
10 Dorsal hairs of aptera long, thick and slightly capitate, arising from large
tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corylobium avellanae
- Dorsal hairs of aptera shorter, not capitate, not borne on tubercles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Dorsal cuticle strongly sclerotized and very wrinkled (although quite pale)
. . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobium) vandenboschi
- Dorsal cuticle not sclerotic, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Hind tibiae and SIPH of aptera wholly black, CAUDA dusky to dark
. . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobiuni) corylicola
- Hind tibiae of aptera with paler middle section; SIPH wholly black or pale
at bases, CAUDA pale to dusky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Aptera with abdominal cuticle usually pigmented marginally, and also
dorsally across ABD TERG 5-7. SIPH wholly black, ABD TERG 8
and CAUDA contrastingly pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobium) coryli
- Aptera with abdomen wholly pale. SIPH usually paler at base, CAUDA
pale or dusky . . . . . . . . . . Macrosiphum (Neocorylobium) pseudocoryli
C. mexicana = C. pubescens
C. mollis Rhopalosiphum insertum
C. monogyna Aphis pomi
Dysaphis angelicae, apiifolia ssp.
petroselini, crataegi, lauberti,
ranunculi
Fimbriaphis gentneri
Nearctaphis bakeri
Ovatus crataegarius
Prociphilus (Stagona) pini
Rhopalosiphum insertum
C. orientalis Dysaphis crataegi ssp. heraclei
C. oxyacantha Aphis fabae, pomi
Aulacorthum solani
Dysaphis crataegi
Fimbriaphis gentneri
Macrosiphum euphorbiae
Nearctaphis crataegifoliae
Ovatus crataegarius, insitus
Prociphilus (Stagona) pini
Rhopalosiphum insertum
C. phaenopyrum Rhopalosiphum insertum
C. pinnatifolia Ovatus crataegarius
C. praeformosa Hyalomyzus tissoti
C. x prunifolia Rhopalosiphum insertum
C. pubescens Muscaphis mexicana
Rhopalosiphum insertum
C. punctata Eriosoma lanigerum
Muscaphis stroyani
Nearctaphis crataegifoliae
C. rivularis Eriosoma lanigerum
Nearctaphis bakeri
C. songorica Dysaphis crataegi ssp. pallida
C. succulenta Nearctaphis crataegifoliae
C. tomentosa Eriosoma crataegi
Prociphilus (Neoparacletus)
corrugatans
C. uniflora Eriosoma lanigerum group
Hyalomyzus eriobotryae
Crataegus 151
Schizoneurata tissoti
Utamphorophora crataegi
C. vicana Hyalomyzus tissoti
Crataegus sp. Dysaphis laserpitii
[Dysaphis ramani Das and
Raychaudhuri, 1983]
Muscaphis canadensis
Rhopalosiphum nigrum, rufulum
Fig. 42. A, dorsal cuticle of fundatrix of Muscaphis stroyani; B, CAUDA of Muscaphis canadensis
(fundatrix); C, CAUDA of M. stroyani (fundatrix); D, CAUDA of Nearctaphis bakeri (aptera); E, CAUDA of
Dysaphis crateaegi; F, dorsal abdominal patch of Dysaphis crataegi kunzei; G, the same for D. crataegi;
H, comparison of positions of marginal tubercle on TERG 7 of Rhopalosiphum (above) and Aphis (below);
I, inner faces of antennal tubercles convergent; J, inner faces of antennal tubercles parallel; K, inner
faces of antennal tubercles diverging; L, SIPH of Rhopalosiphum rufulum spring migrant alata; M, SIPH
of Nearctaphis bakeri aptera; N, SIPH of Hyalomyzus eriobotryae aptera; 0, SIPH of H. tissoti aptera;
P, SIPH of Utamphorophora crataegi aptera; Q, SIPH of Fimbriaphis gentneri aptera.
154 Crataegus
13 Alata with a solid black patch on ABD TERG 3-5, partially fused with
bar on 6 . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis californica nigrescens
- Alata without a solid black dorsal patch, only with transverse bars or
dashes on ABD TERG 5-8 . . . . . . . . . . . . . . . . . . . Nearctaphis sclerosa
14 R IV+V 0.14-0.18 mm long, usually with 4 accessory hairs. Alata without
a solid black dorsal abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- R IV+V 0.10-0.13 mm long, usually with 2 accessory hairs. Alata with a
solid black patch covering ABD TERG 3-5, or 3-6 . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis bakeri
15 Head of aptera blackish-brown, and SIPH darker than CAUDA. Alata
with transverse dark bars on ABD TERG 6-8 or 7-8 . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis crataegifoliae occidentalis
- Head of aptera pale, and SIPH paler than CAUDA. Alata with a complete
transverse dark bar only on ABD TERG 8 . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis crataegifoliae crataegifoliae
16 Spinal tubercles usually present and well developed on head, prothorax
and all abdominal segments. SIPH of aptera pale or dusky, less than
0.14 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis sorbiarum
- Spinal tubercles only on head and ABD TERG 7-8, rarely on other
segments. SIPH dark, more than 0.14 mm long . . . . . . . . . . . . . . . . . 17
17 Spring migrant alata (henceforth 's.m.a.') with marginal tubercles on ABD
TERG 7 in most specimens, frequently on both sides. Hairs on ABD
TERG 8 usually 10-25 µm long . . . . . . . . Dysaphis apiifolia petroselini
- S.m.a. without marginal tubercles on ABD TERG 7 in most specimens,
and very rarely on both sides. Hairs on ABD TERG 8 30-70 µm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 S.m.a. with total number of secondary rhinaria on both ANT V combined
7-35, or if 5-6 then a transverse dark bar is present on ABD TERG 2
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
- S.m.a. with usually only 0-5 secondary rhinaria on both ANT V
combined, rarely up to 6 or 7. ABD TERG 2 without a dark bar ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
19 ABD TERG 1 and 2 of s.m.a. without dark bars, at most with small
scattered circular sclerites . . . . . . . . . . . . . . . . . . . . . . Dysaphis angelicae
- ABD TERG 2, or both 1 and 2, of s.m.a. with a transverse dark bar,
sometimes partly broken into irregular sclerites . . . . . . . . . . . . . . . . . . 20
20 ANT III of s.m.a. with 44-75 secondary rhinaria, ANT IV with 15-35 and
ANT V with 3-19. R IV+V with 4-7 accessory hairs . . . . . . . . . . . . . . .
............................................. Dysaphis ranunculi
Crataegus 155
- ANT III of s.m.a. with 37-47 secondary rhinaria, ANT IV with 10-19 and
ANT V with 2-6. R IV+ V with 2-3 accessory hairs . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis virgata
21 ANT III of s.m.a. usually with 74-125 secondary rhinaria per mm of
length ...................................................... 22
- ANT III of s.m.a. usually with 130-192 secondary rhinaria per mm of
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22 R IV+V of s.m.a. 1.01-1.25 X HT II. Hairs on ANT III blunt or weakly
capitate, 9-21 µm long, those on ABD TERG 3 12-23 µm long . . . . . .
.............................................. Dysaphis incognita
- R IV+V of s.m.a. 0.83-0.98 x HT II. Hairs on ANT III mostly pointed,
20-29 µm long, those on ABD TERG 3 25-35 µm long . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis crataegi pallida
23 S.m.a. with hairs on ABD TERG 2 and 3 pointed, the longest 20-57 µm
long, but rarely less than 25 µm . . . . . . . . . . . . . . . . . . Dysaphis lauberti
- S.m.a. with hairs on ABD TERG 2 and 3 blunt, the longest 9-29 µm, but
rarely more than 25 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 S.m.a. with SIPH 2.9-3.5 x their middle diameter . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis crataegi heraclei
- S.m.a. with SIPH 3.6-5.7 x their middle diameter . . . . . . . . . . . . . . 25
25 S.m.a. with dorsal abdominal patch on ABD TERG 3-6 interrupted by
narrow gaps or lacunae between 3-4 and 4-5, as well as 5-6 (Fig. 42F).
Anterior part of subgenital plate with 4-10 hairs . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis crataegi kunzei
- S.m.a. with dorsal abdominal patch on ABD TERG 3-6 almost entire,
except for a narrow central gap between 5 and 6 (Fig. 42G). Anterior part
of subgenital plate with 2-7 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 (Requires large samples) Most s.m.a. with at least 1 antenna bearing 1 (or
more) secondary rhinaria on ANT V [mean number per antenna 0.53
(UK)-1.1 (former USSR)] . . . . . . . . . . . . . . . . Dysaphis crataegi crataegi
- Most s.m.a. without any secondary rhinaria on ANT V of either antenna
[mean number 0.32 per antenna (UK)] ... Dysaphis crataegi aethusae
27 Antennal tubercles undeveloped or weakly developed. SIPH of aptera
dark or, if only dark at apex, then short (less than 0.25 mm). CAUDA
and anal plate dusky to dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
- Antennal tubercles moderately to well developed. SIPH of aptera pale or
dusky, or dark only at apices and then over 0.3 mm long . . . . . . . . 34
28 SIPH slightly swollen or cylindrical on at least basal 0.7 of length, narrow-
ing only towards apex, and with a well-developed apical flange (e.g.
156 Crataegus
37 R IV+V less than 1.1 x HTII. CAUDA with 6-11 hairs (usually 7). Alata
with a black dorsal abdominal patch . . . . . . . . . . . . . . . Myzus persicae
- R IV+V more than 1.1 x HT II. CAUDA with 4-6 hairs. Alata without
a black dorsal abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38 SIPH of aptera moderately swollen, and only moderately imbricated
(Fig. 42N). ANT PT less than 1.1 x ANT III. Alata with 35 or more
secondary rhinaria on ANT III . . . . . . . . . . . . Hyalomyzus eriobotryae
- SIPH of aptera only slightly swollen, and coarsely imbricated (Fig. 42O).
ANT PT more than 1.2 x ANT III. Alata with 11-25 secondary rhinaria
on ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalomyzus tissoti
39 Spring migrant alata with 60-83 secondary rhinaria on ANT III, 36-52
on IV and 13-22 on V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus insitus
- Spring migrant alata with 22-49 rhinaria on ANT III, 5-20 on IV and 0-10
on V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus crategarius
40 SIPH with a subapical zone of polygonal reticulation (Fig. 121C). Hairs
on ANT III more than 0.5 x basal diameter of segment . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
- SIPH without any polygonal reticulation. Hairs on ANT III less than
0.5 x basal diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41 SIPH swollen on about distal 0.5 of length (Fig. 42P). Alata without dark
dorsal abdominal markings . . . . . . . . . . . . . . . Utamphorophora crataegi
- SIPH tapering, cylindrical or only slightly swollen subapically (e.g.
Fig. 42Q). Alata with dark dorsal abdominal markings . . . . . . . . . . . 42
42 SIPH more than 2 x CAUDA, and in aptera often dark-tipped (Fig. 121K).
Dorsal abdomen of alata with transverse dark bars . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
- SIPH less than 2 x CAUDA, not dark-tipped in aptera (Fig. 42Q). Dorsal
abdomen of alata with an almost solid dark patch across ABD TERG 4-5
or 3-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbriaphis gentneri
CRATERISPERMUM Rubiaceae
CRATOXYLUM Guttiferae
CROSSOPTERYX Rubiaceae
CROTON Euphorbiaceae
CRYPTOMERIA Taxodiaceae
CUDRANIA=MACLURA
160 X Cupressocyparis
X CUPRESSOCYPARIS Cupressaceae
Cupressus has a small but diverse aphid fauna, with no evidence of host
specificity at the species level. The only possible confusion is between the
Cinara species, which are all in the subgenus Cupressobium.
Fig. 44. A, SIPH of Siphonatrophia cupressi; B, ANT VI of Cinara cupressi; C, ANT VI of C. fresai.
7 Aptera with ANT VI BASE bearing 4-7 hairs confined to basal half
(Fig. 44B). R IV 0.12-0.17 mm long, bearing 2-4 accessory hairs; 1.4-1.9 x
HT I. Alata with 1-6 secondary rhinaria restricted to distal half of ANT
III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara cupressi
- Aptera with ANT VI BASE bearing 7-12 hairs, some on distal half
(Fig. 44C). R IV 0.16-0.24 mm long, bearing 5-7 accessory hairs; 1.8-
2.2 x HT I. Alata with 6-11 secondary rhinaria extending over most of
ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara fresai
CUSSONIA Araliaceae
CYCAS Cycadaceae
Aphis gossypii
Dalbergia 163
CYNOMETRA Leguminosae
CYRILLA Cyrillaceae
DAEMONOROPS Palmae
DALBERGIA Leguminosae
DENDROCALAMUS Gramineae
DIMORPHANDRA Leguminosae
Aulacorthum (Neomyzus)
circumflexum
Myzus persicae
(Use key to polyphagous aphids, p. 532)
DIPTEROCARPUS Dipterecarpaceae
DISTYLIUM Hamamelidaceae
Distylium spp. are primary hosts for many members of the Nipponaphidini,
a tribe in the subfamily Hormaphinae. Takahashi (1962a) and Sorin (1987b)
reviewed the species of this group that form galls on Distylium in Japan; Sorin
illustrated the galls of five species, and the embryos from emigrant alatae
of 12 species, but there are probably numerous other species in East and
Southeast Asia still undescribed. Determinations using the partial key below
should therefore be treated very cautiously.
Quadratus yoshinomiyai
Sinonipponaphis monzeni
D. stellare Distylaphis foliorum
Greenidea flacourtiae
Neohormaphis calva
Reticulaphis distylii
Schizoneuraphis gallarum,
longisetosa
Toxoptera aurantii
Fig. 45. A, antenna of Quadratus yushinomiyai (alata from gall); B, same for Dinipponaphis autumna;
C, same for Nipponaphis distychii; D, abdominal spiracle of Schizoneuraphis longisetosa; E, abdominal
spiracle of Nipponaphis monzeni; F, SIPH of N. monzeni; G, ANT I and II of Distylaphis foliorum alata
from gall; H, same for Nipponaphis distyliicola; I, same for Neothoracaphis yanonis; J, same for
Reticulaphis distylii.
- Vein Cu1b in forewing clearly thicker than vein Cula, especially at base
............................................................. 8
8 Wing membrane heavily infuscated, at least between vein Culb and base
of forewing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Wing membrane between veins almost hyaline, or slightly but evenly
pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
9 Antenna more than 2 x as long as head width across (and including) eyes,
ANT III being as long as or a little longer than fore tibia (Fig. 45C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis distychii
- Antenna less than 2 x as long as head width across eyes. ANT III clearly
shorter than fore tibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Distylium 169
Fig. 46. Forewings of alate gallicolae of A, Neohormaphis calva (after Noordam, 1991), B, Monzenia
globuli, C, Distylaphis foliorum, D, Schizoneuraphis longisetosa, E, Nipponaphis distylicola,
F, Schizoneuraphis gallarum, G, Neothoracaphis yanonis, H, Reticulaphis distylii.
DODONAEA Sapindaceae
DOMBEYA Sterculiaceae
DOVYALIS Flacourtiaceae
DUABANGA Lythraceae
DYSOXYLUM Meliaceae
EHRETIA Boraginaceae
ELAEAGNUS Elaeagnaceae
Elaeagnaceae are the primary hosts of the eight to nine species of Capito-
phorus known to have host alternation. One species (shepherdiae) seems to
have a specific primary host association with Shepherdia argentea (which is
not Elaeagnus argentea), but the other species are apparently able to utilize
either Elaeagnus or Hippophae, presumably depending on host availability.
- ABD TERG 1-4 usually with submarginal hairs, so that there are 4
more-or-less complete longitudinal rows of hairs on each side; these hairs
have mushroom- or balloon-shaped apices, and are shorter than their
maximum widths. Marginal hairs usually duplicated. SIPH more than
2.7 x CAUDA. Dorsal cuticle with variolate or reticulate sculpturing
............................................................. 5
5 ABD TERG 1-4 with pleural hairs usually single. BL probably more
than 1.4mm, and ANT PT/BASE probably more than 2.5 except in
fundatrices. (The spring generations of this species on its primary host
are not yet recorded) . . . . . . . . . . . . . . . . . . . . . . . . Capitophorus xanthii
- ABD TERG 1-4 with pleural hairs mostly duplicate. BL less than 1.4 mm,
ANT PT/BASE less than 2.0 . . . . . . . . . . . . Capitophorus shepherdiae
6 ABD TERG 1-4 each with 6-8 hairs (usually only one pair each of spinal,
pleural and marginal hairs) . . . . . . . . . . . . . . . . . . Capitophorus elaeagni
- ABD TERG 1-4 each with 10-26 hairs, due to duplication or multiplica-
tion of spinal, pleural and marginal hairs on each tergite . . . . . . . . . 7
7 SIPH short, tapering continuously from base to apex, less than 11 x
longer than width at midpoint . . . . . . . . . . . Capitophorus himalayensis
- SIPH attenuated, almost cylindrical except at base, or with slight sub-
apical swelling; more than 12 x width at midpoint . . . . . . . . . . . . . . . 8
8 Hairs on ANT III all short, thin and inconspicuous, 0.3-0.5 x basal
diameter of segment. R IV+V 2.5-3.1 x HT II . . . . . . . . . . . . . . . . . . . .
.................................... Capitophorus meghalayensis*
- ANT III with 1-5 forwardly-directed thick clavate hairs like those on ANT
II or a little shorter, 0.5-1.2 x longer than base of segment. R IV+V
1.5-2.6 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Thick capitate hairs on ANT III all less than 0.8 x basal diameter of
segment. (R IV+V 1.5-2.1 x HT II) . . . . . . . . . . . Capitophorus similis
- Longest capitate hairs on ANT III more than 0.8 x basal diameter of
segment. (R IV+V 1.6-2.6 x HT II) . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 ABD TERG 1-4 each with 18-26 capitate hairs. R IV+V 2.1-2.6 x HT
II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capitophorus pakansus
- ABD TERG 1-4 each with 12-19 capitate hairs. R IV+V 1.6-2.1 x HT
II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capitophorus archangelskii
Eleutherococcus 175
ELAEIS Palmae
ELAEOCARPUS Elaeocarpaceae
ELEUTHEROCOCCUS Araliaceae
ENCEPHALARTOS Zamiaceae
Aphis gossypii
ENGELHARDTIA Juglandaceae
ERIOBOTRYA Rosaceae
ERVATAMIA = TABERNAEMONTANA
178 Erythrina
ERYTHRINA Leguminosae
ESCALLONIA Escalloniaceae
EUCALYPTUS Myrtaceae
EUCLEA Ebenaceae
EUCOMMIA Eucommiacee
EUGENIA Myrtaceae
EUONYMUS Celastraceae
Several subspecies or sibling species of the Aphis fabae group use Euonymus
europaeus as their primary host in Europe (Müller and Steiner, 1986). These
are suspected to hybridize in nature to some extent, and the morphological
separations attempted in couplets 6 and 7 of the key below may not be very
reliable. For biological differences between members of this group consult the
text (p. 559).
2 Antennal tubercles with inner faces convergent in dorsal view (Fig. 42I).
Aptera without any secondary rhinaria on ANT III. Alata with a dark
dorsal abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
Antennal tubercles with inner faces parallel or divergent in dorsal view.
Aptera with one or more secondary rhinaria near base of ANT III. Alata
either without dorsal abdominal markings or with transverse segmentally
divided bars ................................................. 3
3 Antennal tubercles smooth with inner faces divergent in dorsal view
(Fig. 42K). SIPH tapering, with a longer-than-wide subapical zone of
polygonal reticulation. (Longest hairs on ANT III more than 0.5 of basal
diameter of segment.) Alata without dorsal abdominal markings .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
Antennal tubercles with inner faces scabrous, parallel in dorsal view
(Fig. 42J), SIPH without a zone of polygonal reticulation. Alata with
brown transverse bars on dorsal abdomen . . . . . . . . . . . . . . . . . . . . . . . 4
4 Femora and CAUDA mainly dark. SIPH swollen on distal half . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum magnoliae
Femora and CAUDA mainly pale. SIPH tapering . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
5 ANT PT/BASE 3.5-5.0. Abdominal sternites 5 and 6 with conspicuous
sclerotic ridges, and hind tibia with a row of evenly spaced, short,
stridulatory pegs (Fig. 122) . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
ANT PT/BASE 1.7-3.5. Stridulatory ridges and pegs absent . . . . . . 6
6 ANT III 3-5 x longer than the longest hair borne upon it, which is
2-3 x basal diameter of the segment. Marginal tubercles completely
absent from ABD TERG 2-6. Males apterous, oviparae with hind tibiae
only slightly swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis euonymi
ANT III 4-20 x longer than the longest hair borne upon it, which is
0.6-2.5 (but rarely more than 2.0) X basal diameter of the segment. ABD
TERG 2-6 with a total of 0-7 (usually 1-6) marginal tubercles. Males
alate, oviparae with strongly swollen hind tibiae . . . . . . . . . . . . . . . . . . 7
7 ANT III of aptera 11-20 x longer than the longest hair borne upon it.
Hairs on marginal sclerites of ABD TERG 3 of aptera 12-49 µm long, of
alata 18-51µm long, usually less than 2 x maximum diameter of the
spiracular pore on that segment . . . . . . . . . . . . . . Aphis fabae solanella
ANT III of aptera 4-9 x longer than the longest hair borne upon it. Hairs
on marginal sclerites of ABD TERG 3 of aptera 40-85 µm long, of alata
31-83 µm long, more than 2 x maximum diameter of spiracular pore on
that segment . . . . . . . . . . . . . Aphis fabae fabae/fabae cirsiiacanthoidis
182 Euodia
EUPTELEA Eupteleaceae
EUSCAPHIS Staphyleaceae
FAGARA = ZANTHOXYLUM
Fig. 47. A, margin of abdomen of Platyaphis fagi (aptera); B, dorsal abdominal wax pore plate of
fundatrix of Grylloprociphilus imbricator; C, dorsal abdominal wax pore plate of Eriosoma crataegi,
D, ANT VI of Phyllaphis fagi (aptera); E, dorsal abdominal wax pore plate of Ph. fagi (alata).
ANT VI short and stubby, its base less than 2 x longer than wide.
Forewings of alata maculate, with pterostigma not extending to tip of
wing. BL usually less than 4.5 mm . . . . . . . . . . . . . . . . Lachnus pallipes
5 Apterae with faceted wax glands. Alatae with secondary rhinaria on ANT
III-V or III-VI, media unbranched or once-branched, and a broadly
rounded CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Apterae (if produced) with wax glands comprising groups of ring-shaped
pores (Fig. 47E). Alatae with secondary rhinaria only on ANT III, a
twice-branched media and a knobbed CAUDA . . . . . . . . . . . . . . . . . . . 7
6 Apterae (fundatrices) with 4-segmented antennae and circular groups of
wax pore plates on all segments, each with large central facets surrounded
by very much smaller marginal ones (Fig. 47B). SIPH absent. Alata with
strongly ciliated secondary rhinaria and an unbranched media .......
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grylloprociphilus imbricator
Apterae with 6-segmented antennae, wax pore plates smaller and usually
comprising a ring of large facets around a central area which may be
Ficus 185
divided into small cells (Fig. 47C). SIPH present as partly sclerotized rings
with surrounding hairs. Alatae with unciliated secondary rhinaria and a
once-branched media . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma crataegi
7 All viviparae alate. ANT PT/BASE about 1. Dorsal abdomen of alata
pale. SIPH as pale, truncate cones . . . . . . . . . . . . . . Mesocallis fagicola
Viviparae apterous or alate. ANT PT/BASE less than 0.25 (Fig. 47D).
Dorsal abdomen of alata pale or with variably developed transverse bars
and pigmented marginal sclerites bearing numerous wax pores (Fig. 47E).
SIPH pore-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Alatae with pale head and pterothorax and no dorsal abdominal cross-
bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllaphis fagifoliae
Alatae with dark head and pterothorax and pigmented dorsal abdominal
cross-bands and marginal sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllaphis fagi (and grandifoliae?)
Aphids on Ficus carica are keyed in B & E (1984). The present treatment is
extended to cover all aphids recorded from the genus Ficus.
Fig. 48. A, SIPH and adjacent cuticle of Nipponaphis ficicola (aptera); B, head of Reticulaphis mirabilis
(after Takahashi, 1939a); C, head of Reticulaphis distylii ssp. minutissima (with detail of dorsal cuticle);
D, SIPH of Greenidea formosana group (aptera); E, SIPH of G. ficicola (aptera); F, dorsal view of front of
head and processes on ANT I of Matsumuraja calorai aptera; 6, ANT I-IV of Toxoptera schlingen alata;
H, ANT I-IV of T. aurantii alata.
- SIPH of aptera usually a little shorter than hind tibiae and clearly less than
0.5 x BL. Alata with 12-22 secondary rhinaria on ANT III . . . . . . . . .
.............................................. Greenidea ficicola
8 ANT I of aptera very broad, with a finger-like distal projection, extending
beyond apex of ANT II and bearing short, capitate hairs (Fig. 48F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Matsumuraja calorai
- ANT I without a finger-like projection . . . . . . . . . . . . . . . . . . . . . . . . 9
9 SIPH with a subapical zone of polygonal reticulation . . . . . . . . . . . . 10
- SIPH without any polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . 11
10 Aptera with SIPH usually uniformly dark, and dorsal abdomen often with
a pattern of dark segmental markings. R IV+V 0.8-0.9 x HT II. Alata
with 4-14 secondary rhinaria mostly concentrated on basal half of ANT
III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion africanum
- Aptera with SIPH often pale at base, and dorsal abdomen without
dark markings. R IV+V 0.95-1.15 X HT II. Alata with 14-18 second-
ary rhinaria in spaced-out row along most of ANT III . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion halli
11 Antennae always 5-segmented, in both apterae and alatae . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis kachkouli
- Antennae 6-segmented (except sometimes in small apterae of Aphis
gossypii) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Stridulatory apparatus present, comprising stridulatory ridges on abdom-
inal sternites 5 and 6 and evenly-spaced peg-like hairs on the hind tibiae
(Fig. 122) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Stridulatory apparatus absent ... go to key to polyphagous aphids, p. 532
13 SIPH 0.4-0.7 x CAUDA, and 0.05-0.08 x BL . . . . Toxoptera odinae
- SIPH 0.9-1.9 x CAUDA, 0.11-0.18 X BL . . . . . . . . . . . . . . . . . . . . . . 14
14 CAUDA with 19-54 hairs (rarely less than 25). Hairs on ANT III of aptera
mostly longer than basal diameter of segment. Alata with forewing media
usually twice-branched . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera citricidus
- CAUDA with 7-26 hairs (rarely more than 20). Hairs on ANT III of
aptera mostly shorter than basal diameter of segment. Alata with forewing
media usually once-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Alata with 12-18 large and small secondary rhinaria on ANT III, 3-6 on
ANT IV and 0-2 on ANT V (Fig. 48G). ANT PT/BASE often less than
3.5 in aptera, and often less than 3.75 in alata . . . . . . . . . . . . . . . . . . . .
............................................ Toxoptera schlingeri
- Alata with 2-10 more evenly-sized secondary rhinaria on ANT III, 0-3
on ANT IV, 0 on ANT V (Fig. 48H). ANT PT/BASE greater than 3.5
in aptera, greater than 3.75 in alata . . . . . . . . . . . . . Toxoptera aurantii
190 Fraxinus
Fraxinus species are primary or only hosts for several members of the
pemphigine genus Prociphilus, the spring generations of which form leaf
nests. Only the alatae produced by these leaf-nest colonies are keyed here.
Of the other aphids described or recorded from Fraxinus, only one (Myzus
beybienkoi) has been collected more than once. The others are not included
in the key and are indicated by square brackets in the following list.
Fig. 49. A, antenna of Prociphilus fraxinifolii alata; B, antenna of P. probosceus alata; C, ANT I-III and
VI of P. pergandei alata; heads of spring migrant alatae of D, P. probosceus, E, P. bumeliae and
F, P. fraxini (o = ocellus, w = wax gland).
FUCHSIA Onagraceae
Aulacorthum solani
Aulacorthum (Neomyzus)
circumflexum
Macrosiphum euphorbiae
Myzus ornatus, persicae
(Use key to polyphagous aphids, p. 532)
GARCINIA Guttiferae
GARDENIA Rubiaceae
GIGANTOCHLOA Gramineae
Ceratoglyphina bambusae
Pseudoregma bambusicola
(Use key to aphids on Bambusa, p. 83)
GLEDITSIA Leguminosae
Brachycaudus helichrysi
Myzus persicae
(Use key to polyphagous aphids, p. 532)
GLIRICIDIA Leguminosae
Aphis craccivora
GLOCHIDION Euphorbiaceae
with R IV+V 1.15-1.30 x HT II. Alata with SIPH 0.12-0.18 x BL, and
1.4-1.8 x CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis eugeniae
Hind tibia without any short peg-like hairs. HT I with 2 hairs. Aptera
with R IV+V 1.0-1.16 x HT II. Alata with SIPH 0.09-0.14 x BL and
1.2-1.5 x CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
GREVILLEA Proteaceae
GREWIA Tiliaceae
GREYIA Greyiaceae
No aphids recorded
196 Guazuma
GUAZUMA Sterculiaceae
GUETTARDA Rubiaceae
HAGENIA Rosaceae
Macrosiphum euphorbiae
HAMAMELIS Hamamelidaceae
Fig. 50. Hindwings of alatae from galls of A, Hormaphis hamamelidis, B, Hamamelistes spinosus,
C, Hormaphis betulae, D, Hamamelistes miyabei.
4 ANT IV distinctly longer than (more than 1.2 x) ANT V. Hindwing with
the 2 oblique veins united or close together at their bases (e.g. Fig. 50D)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamamelistes miyabei
- ANT IV about equal in length to (0.9-1.1 x) ANT V. Hindwing with the
2 oblique veins separated at their bases . . . . . . . Hamamelistes kagamii
HARPEPHYLLUM Anacardiaceae
HARUNGANA Guttiferae
HEMIPTELEA Ulmaceae
pale areas around the hair bases. Dorsal abdomen without finger-like
tubercles but with dark markings. ANT PT/BASE less than 0.5. ANT III
with about 6-8 widely-spaced oval secondary rhinaria . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis (Pseudochromaphis) coreanus
Forewing membrane with fuscous bordering distal branches of media and
patches at ends of Cula and Cu1b (Fig. 51B). Head and prothorax rather
uniformly dark. Dorsal abdomen pale, with finger-like spinal tubercles on
ABD TERG 1 and 2. ANT PT/BASE about 1 or more. ANT III with
14-18 secondary rhinaria . . . . . . . . . . . . . . . . . Tinocallis takachihoensis
HERNANDIA Hernandiaceae
HETEROMELES = PHOTINIA
HIBISCUS Malvaceae
Hibiscus has a very similar aphid fauna to cotton (keyed in Blackman and
Eastop, 1984). The aphids are all oligophagous or polyphagous, so there is
no point in listing the host species separately.
2 Antennae with numerous long hairs. ANT II much longer than ANT I.
ANT PT/BASE less than 0.2. SIPH absent. CAUDA broadly rounded
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Smynthurodes betae
- Antennae with sparse, short hairs. ANT II similar in length to ANT I.
ANT PT/BASE more than 0.5. SIPH present. CAUDA tongue-shaped
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachyunguis harmalae
3 SIPH long, thin and pale, about 3 x CAUDA and more than 0.3 x BL
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon gossypii
SIPH pale or dark but not more than 2.5 x CAUDA or more than
0.25 x BL . . . . . . . . . . . . . . . . . . see key to polyphagous aphids, p. 532
All the aphid species found on Hippophae are also recorded from Elaeagnus,
so the key to Elaeagnus aphids can be used to discriminate the species listed
below. It is likely that all heteroecious Capitophorus species can utilize either
Elaeagnus or Hippophae as primary host, depending on availabity.
HOLMSKIOLDIA Verbenaceae
HOVENIA Rhamnaceae
HYMENOCARDIA Euphorbiaceae
HYMENODICTYON Rubiaceae
HYMENOSPORUM Pittosporaceae
HYPTIS Labiatae
INGA Leguminosae
IOZOSTE = ACTINODAPHNE
JACARANDA Bignoniaceae
Fig. 52. CAUDA and anal plate of A, Panaphis nepalensis nepalensis and B, P. juglandis; anal plates of
C, Chromaphis hirsutistibus and D, Ch. californica; E, hairs on middle section of hind tibia of
Ch. hirsutistibus; F, dorsal process of Dasyaphis onigurumi (aptera); G, SIPH of aptera of Kurisakia
onigurumi.
9 ANT PT/BASE less than 0.5. Anal plate only shallowly indented
(Fig. 52C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- ANT PT/BASE more than 0.5. Anal plate deeply bilobed (e.g. Fig. 52D)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 Hairs on hind tibia very long, up to about 3.5 x middle diameter of the
tibia (Fig. 52E). Wings with dark spots at ends of veins (use hand lens)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chromaphis hirsutustibis
- Hairs on hind tibia shorter, the longest of them less than 2 x middle
diameter of tibia. Wings without dark spots at ends of veins . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chromaphis juglandicola
11 Forewing veins Cu1a and Cu1b both bordered with fuscous, much darker
than other veins. Each antennal segment darkening over distal half of its
length. Tibiae dark . . . . . . . . . . . . . . . . . . . . . . . . Chromaphis californica
Juniperus 207
- Forewing veins Cula and Cu1b not darker than other veins, or only
a little darker. Each antennal segment ringed with black apically. Tibiae
pale or dusky . . . . . . . . . . . . . . . . . . Monelliopsis spp. (bisetosa, caryae,
nigropunctata, tuberculata) - use couplets 7-10 of key to aphids on Carya.
12 CAUDA knobbed. Dorsum of aptera bearing numerous long processes,
each tipped with a single bristle (Fig. 52F), these processes as long as or
longer than the reduced, apparently 3-segmented antennae. SIPH as
small, inconspicuous rings . . . . . . . . . . . . . . . . . . . . . . . Dasyaphis rhusae
- CAUDA rounded. Dorsum of aptera without processes, but with numer-
ous long fine hairs. SIPH on broad cones ringed with long hairs (Fig. 52G).
Antennae of both alata and aptera 5-segmented . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kurisakia onigurumi
JULBERNARDIA Leguminosae
Gootiella tremulae
Illinoia morrisoni
J. conferta Cinara fresai
Illinoia morrisoni
J. horizontalis Cinara cupressi, fresai,
manitobensis, petersoni
Gootiella tremulae
Illinoia morrisoni
J. japonica Cinara fresai
J. macrocarpa Cinara cupressi
J. monosperma Cinara tonaluca
Illinoia morrisoni
J. monticola Illinoia morrisoni
J. occidentalis Cinara fresai, rubicunda
Illinoia morrisoni
J. osteosperma Cinara burrilli, pulverulens,
wahhaca
J. oxycedrus Cinara cupressi, fresai, juniperi,
?petersoni
J. pseudosabina Cinara dahurica, tujafilina
J. sabina [Aploneura juniperina = Gootiella
alba?]
Cinara fresai
Illinoia morrisoni
J. scopulorum Cinara burrilli, cupressi, fresai,
pulverulens, wahluca
Illinoia morrisoni
Sanbornia juniperi
Siphonatrophia cupressi
J. squamata Cinara fresai
Illinoia morrisoni
J. virginiana Cinara cupressi, fresai,
juniperivora
Illinoia morrisoni
Sanbornia juniperi
Siphonatrophia cupressi
Fig. 53. A, SIPH of Illinoia morrisoni (aptera); B, hind leg of aptera of Gootiella tremulae; C, ventral
aspect of head of Sanbornia juniperi aptera; D, middle section of hind tibia of Cinara juniperivora
(aptera); E, hind tarsus of Cinara wahluca showing measurement of dorsal length and basal width;
F, same for C. juniperi; G, ANT VI of C. juniperi; H, ANT VI of C. petersoni.
Juniperus 211
- ANT PT/BASE less than 0.4. Diameter of base of SIPH cone more than
0.14 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 ANT BASE VI less than 0.5 x HT II ... Cinara burrilli*/pulverulens
- ANT BASE VI more than 0.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Dorsum with numerous hairs, not confined to transverse rows. ANT III
with a rhinarium near apex . . . . . . . . . . . . . . . . . . . Cinara manitobensis
- Dorsum with fewer hairs arranged in transverse rows. ANT III usually
without a primary rhinarium . . . . . . . . . . . . . . . . . . . . . . Cinara cognita*
12 HT I with dorsal length a little more than, or at least equal to (1.0-1.2 x)
basal width (e.g. Fig. 53E). PT with 4-5 subapical hairs . . . . . . . . . . 13
- HT I with dorsal length distinctly shorter than (0.5-0.8 x) basal width
(e.g. Fig. 53F). PT usually with 3 subapical hairs (subgenus Cupressobium)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13 Greatest diameter of SIPH cone 0.25 mm, less than R IV+V. HT II less
than 0.25 mm. Base VI less than 0.14mm . . . . . . . . . . Cinara wahluca
- Greatest diameter of SIPH cone at least 0.28mm, more than R IV+V.
HT II more than 0.25mm long. Base VI usually more than 0.14mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara juniperensis
14 Femora usually wholly pale except at point of articulation with tibia,
and tibiae only dark at apices. ANT PT/BASE usually about 0.2. R IV
0.14-0.18 mm long, HT II 0.20-0.28 mm long . . . . . . Cinara tujafilina
- Femora distally dark and tibiae either wholly dark or dark at base as well
as apex. If ANT PT/BASE is less than 0.25 then R I V is more than 0.2 mm
long and HT II is more than 0.3 mm long . . . . . . . . . . : . . . . . . . . . . . 15
15 ANT VI BASE with 4-7 hairs restricted to basal half (Fig. 44B). Tibiae
only dark at base and apex . . . . . . . . . . . . . . . . . . . . . . . Cinara cupressi
- ANT VI BASE with 7-15 hairs not restricted to basal half. Tibiae wholly
dark or paler in middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 BL more than 3.3 mm. Length of sclerotized part of stylet groove (Fig. 74A,
p. 292) more than 1.5 mm, and hind tibia more than 2.2 mm long. Hairs on
hind tibia about as long as its diameter at midpoint . . . . . . . . . . . . . . . 17
- BL usually less than 3.3 mm, rarely up to 3.5 mm. Length of sclerotized
part of stylet groove less than 1.2mm, length of hind tibia less than
2.15mm, with dorsal tibial hairs at least 1.5 x width of hind tibia at
midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17 ANT PT less than 0.05mm, ANT PT/BASE 0.17-0.22. R IV+V
0.35-0.4mm, 4.8-5.0 x its basal width . . . . . . . . . . . . Cinara dahurica*
- ANT PT more than 0.05mm, ANT PT/BASE 0.22-0.33. R IV+V
0.29-0.34 mm, 3.1-4.1 x its basal width . . . . . . . . . Cinara smolandiae
212 Kalopanax
18 Hind tibiae with pale section on basal half. ANT III usually longer than
maximum diameter of base of SIPH cone . . . . . . . . . . . . Cinara fresai
- Hind tibiae wholly dark. ANT III usually shorter than maximum diameter
of base of SIPH cone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 ANT BASE VI narrow and often slightly constricted at about its midpoint
(Fig. 53G); width at midpoint usually less than 0.2 of length (look at
several specimens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara juniperi
- ANT BASE VI broader, width at midpoint more than 0.2 of length (e.g.
Fig.53H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 ANT BASE VI longer than both ANT IV and R IV . . . . . . . . . . . . . . .
................................................ Cinara petersoni
- ANT BASE VI shorter than both ANT IV and R IV . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara mordvilkoi
KALOPANAX Araliaceae
KETELEERIA Pinaceae
Densely hairy aphid, BL more than 3 mm. Aptera with multifaceted eyes.
Secondary rhinaria rounded. SIPH as large pores on shallow pigmented
hairy cones. CAUDA rounded with numerous hairs . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara keteleeriae
Sparsely haired aphid, BL less than 2.5 mm. Aptera with 3-faceted eyes.
Secondary rhinaria of alata transversely elongate. SIPH as small pores
without surrounding hairs or pigmentation. CAUDA triangular with 2
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mindarus keteleerifoliae
Laburnum 213
KIELMEYERA Guttiferae
KIGELIA Bignoniaceae
KOELREUTERIA Sapindaceae
KRAUSSIA Rubiaceae
LABURNUM Leguminosae
LACCOSPERMA Palmae
Fig. 54. Spinal tubercles on ABD TERG 2 of A, Tinocallis kahawaluokalani and B, T. himalayensis.
LANNEA Anacardiaceae
LANSIUM Meliaceae
LAPORTEA Urticaceae
The aphid fauna of Larix seems to reflect the apparently close relation-
ship between the tree species in this genus. Host records of the best-known
larch-feeding aphids (Cinara cuneomaculata, C. laricis, Adelges laricis)
show no sign of discrimination between species of Larix, and it seems possible
that the less well-known and less widely-distributed aphids would be equally
non-specific in their choice of hosts if given the opportunity. For Adelges
species, it is not possible to provide a complete key in the present state of
knowledge.
L. kaempferi = L. leptolepis
L. kamtschatica Cinara cuneomaculata, kochiana,
laricicola, laricis
L. koraiensis (koreana?) Adeiges karamatsui
Cinara kochiana ssp. kochi
L. laricina Adelges aenigmaticus, lariciatus,
(Alaskan Larch, Tamarack) laricis
Cinara cuneomaculata, laricifex,
spiculosa, subterranea
L. leptolepis Adelges isedakii, karamatsui,
(Japanese Larch, Karamatsu) kitamiensis, laricis, torii,
viridana
Cinara chibi, cuneomaculata,
kochiana, kochiana ssp. kochi,
laricicola, laricis
Prociphilus [?kuwanai], laricis,
[ushikoroshi]
L. lyalli Adelges lariciatus
Cinara lyalli
L. occidentalis Adelges oregonensis
(Western Larch) Cinara laricifoliae
L. pontaninii Adelges laricis ssp. pontaninilaricis
L. sibirica Adelges tardoides, viridana,
(Siberian Larch) viridis, viridula
Cinara cuneomaculata
Elatobium laricis
Fig. 55. A, head of neosistens of Adelges viridis showing wax glands with double-walled facets; head
and prothorax of B, A. viridana neosistens and C, A. laricis (partly after Schneider-Orelli and Schneider,
1954); head and prothorax of alata (sexupara) showing arrangement and details of wax glands of
D, A. viridis, E, A. viridana and F, A. laricis; G, adult sistens of A. viridana (in May); H, adult sistens
of A. laricis (in April); I, adult progrediens of A. laricis (July).
220 Larix
Fig. 56. Sclerites on ABD TERG IV of apterae of A, Cinara lyalli, B, C. laricifoliae, C, C. laricifex,
D, C. spiculosa, E, C. laricis.
LAURUS Lauraceae
LEPTOSPERMUM Myrtaceae
LEUCADENDRON Proteaceae
LEUCAENA Leguminosae
LEUCOSIDEA Rosaceae
Longest hairs on ANT III less than 2 x (and usually shorter than) basal
diameter of segment. SIPH usually clearly longer than CAUDA, which
bears less than 14 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 SIPH conspicuously swollen over about 0.7 of length, and in aptera
having basal half pale and distal half dark. Alata with secondary rhinaria
on ANT IV and a black dorsal abdominal patch . . . . . Myzus ligustri
SIPH tapering or slightly swollen on distal half, but if swollen then in
aptera only dark at extreme apices. Alata without secondary rhinaria on
ANT IV, with or without a black dorsal patch . . . . . . . . . . . . . . . . . . . 6
6 SIPH long, more than 0.2 x BL, and uniformly dark in aptera, much
darker than CAUDA; without any polygonal reticulation . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum ibotum
SIPH either much less than 0.2 x BL or if longer then not much darker
than CAUDA in aptera, and with a subapical zone of polygonal reticula-
tion . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
LINDERA Lauraceae
Rostrum much shorter than body. Forewing of alata with wing veins not
or only very lightly bordered, and with pterostigma extended to wing-tip
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma liquidambarus
LIRIODENDRON Magnoliaceae
Fig. 57. A, margin of prosoma of Sinonipponaphis hispida, with hair-like processes; B, prosomal
marginal hairs of Uichancoella gabrielli; C, ventrolateral part of abdomen of Mollitrichosiphum tenuicorpus
and D, ridges on basal part of hind tibia (also present in M. lithocarpi).
LITSEA Lauraceae
Fig. 58. Marginal prosomal hairs of apterae of A, Schizoneuraphis gallarum and B, S. litseicola] C, part
of surface of prosoma of Nipponaphis litseae showing pustulate ornamentation and dorsal hairs;
abdominal plate of D, S. himalayensis, E, Schizoneuraphis gallarum and F, S. litseicola.
LIVISTONA Palmae
LONCHOCARPUS Leguminosae
Aphis craccivora
LOXOSTYLIS Anacardiaceae
MAACKIA Leguminosae
MACADAMIA Proteaceae
Aphis gossypii
Myzus persicae
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)
234 Macaranga
MACARANGA Euphorbiaceae
MACHILUS = PERSEA
MACLURA Moraceae
MAGNOLIA Magnoliaceae
MALACANTHA = POUTERIA
236 Mallotus
MALLOTUS Euphorbiaceae
MALPIGHIA Malpighiaceae
Fig. 60. A, antenna of Aphidounguis mali (aptera); dorsal abdominal wax pore plate of B, Schizoneurella
indica and C, Eriosoma lanigerum; head of spring migrant alata of D, Prociphilus caryae fitchii and
E, P. sasakii; SIPH of apterae of F, Ovatus malisuctus, G, 0. crataegarius and H, Rhopalosiphum
insertum; hairs on HT I of I, Aphis eugeniae and J, A. pomi.
- BL of s.m.a. less than 4 mm. R IV+V less than 0.16 mm long and bearing
2-5 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 S.m.a. without any clear wax pore plates on head, and secondary rhinaria
on ANT V numbering 8-10 . . . . . . . . . . . . . . . . . . . Prociphilus kuwanai
- S.m.a. with 1-2 pairs of very pale and clearly-defined wax pore plates on
head; secondary rhinaria on ANT V numbering 0-7 . . . . . . . . . . . . . . 8
8 S.m.a. with one pair of circular wax pore plates on vertex of head. ANT
I (outside length) as long as or longer than its basal width (Fig. 60E). SIPH
present as small pigmented rings . . . . . . . . . . . . . . . . Prociphilus sasakii
240 Malus
- S.m.a. with 2 pairs of cephalic wax pore plates, one pair on frons and
the other on vertex, and ANT I shorter than its basal width (Fig. 60D).
SIPH absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus caryae fitchii
9 Dorsal abdomen with paired, pigmented spinal tubercles on each segment.
Alata with maculate forewings and short pterostigma . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterochloroides persicae
- Dorsal abdomen without dark tubercles. Forewings of alata not maculate,
with a long pterostigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Forewing of alata with pterostigma curved distally, extended around wing-
tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma xizangensis*
- Forewing of alata with pterostigma almost straight, not extending around
tip of wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrolachnus pyri
11 Head with well-developed antennal tubercles, their inner faces scabrous
or spinulose and bearing short, blunt or slightly clavate hairs (e.g.
Fig.42I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- Head with antennal tubercles either undeveloped or with inner faces
smooth and divergent and bearing pointed hairs . . . . . . . . . . . . . . . . . 15
12 SIPH moderately swollen on distal half (Fig. 121H) . . . . Myzus persicae
- SIPH cylindrical or tapering on distal half, or at most slightly swollen
subapically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Genital plate enlarged and produced posteriorly. ANT PT/BASE less than
4. SIPH dark, very coarsely imbricated (scabrous), and with a small flange
(Fig. 60F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus malisuctus
- Genital plate normal. ANT PT/BASE more than 4. SIPH pale or dusky,
moderately imbricated, with a well-developed flange (Fig. 60G) . . . . . 14
14 Spring migrant alata with 60-83 secondary rhinaria on ANT III, 36-52
on ANT IV and 13-22 on ANT V . . . . . . . . . . . . . . . . . . Ovatus insitus
- Spring migrant alata with 22-49 rhinaria on ANT III, 5-20 on IV and 0-10
on V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus crataegarius
15 Head capsule of aptera spinulose or nodulose . . . . . . . . . . . . . . . . . . . 16
- Head capsule of aptera smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
16 SIPH short, of similar length to short CAUDA, with closely-spaced rows
of nodules or spinules (Fig. 42M) . . . . . . . . . . . . . . . Nearctaphis bakeri
- SIPH more than 4 x CAUDA; jet black and imbricated in aptera
(= fundatrix), dark and mainly smooth in alata . . . . Muscaphis smithi
17 CAUDA tongue- or finger-shaped, clearly longer than basal width . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
- CAUDA short, helmet-shaped, semicircular or triangular, not longer than
or about as long as its basal width in dorsal view . . . . . . . . . . . . . . . 27
Malus 241
18 Body oval, with SIPH only about half as long as the distance between
their bases, without polygonal reticulation. Antennal tubercles weakly
developed or undeveloped. Aptera without rhinaria on ANT III . . . . 19
- Body spindle-shaped, SIPH longer than the distance between their bases,
with a subapical zone of polygonal reticulation. Antennal tubercles diver-
gent in dorsal view. Aptera with a few rhinaria on ANT III near base
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
19 SIPH slightly swollen subapically and constricted before the well-
developed apical flange (Fig. 60H). Marginal tubercles on ABD TERG 7
placed posteriodorsally to spiracle, and no larger than the spiracular open-
ing (Fig. 42H) . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum insertum
- SIPH tapering from base to apex, with flange only moderately developed.
Marginal tubercles on ABD TERG 7 placed posterioventrally to spiracle
and usually larger than spiracular opening (Fig. 42H) . . . . . . . . . . . . 20
20 Stridulatory apparatus present, with sclerotic ridges on abdominal ster-
nites 5 and 6 and a row of peg-like hairs on hind tibia (Fig. 122). ANT
PT/BASE 3.5-5.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
- No Stridulatory ridges present and peg-like hairs on hind tibia only in one
species (Aphis eugeniae). ANT PT/BASE less than 3.5 . . . . . . . . . . 21
21 Femoral hairs long and fine, many exceeding the length of the trochantero-
femoral suture. CAUDA with 6-24 hairs . . . . . . . . . . . . . . . . . . . . . . . 22
- Femoral hairs mostly short, with only a few ventral hairs long and those
not exceeding the length of the trochantero-femoral suture. CAUDA with
4-8 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
22 Aptera with black transverse bands on ABD TERG 7 and 8, and at least
some dark spinopleural markings anterior to SIPH. Alata with a series
of dorsal abdominal transverse bars. CAUDA tongue-shaped, without a
trace of a constriction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis fabae
- Aptera without dark dorsal abdominal markings, and alata with only
marginal sclerites. CAUDA finger-shaped, with a slight constriction near
midpoint (e.g. Fig.121T) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 Hind tarsus with 3 hairs (a sense peg plus 2 lateral hairs), like the fore
and mid-tarsi (Fig. 60I). Basal half of hind tibia with a spaced-out row
of short peg-like hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis eugeniae
- Hind tarsus with only 2 hairs (no sense peg; Fig. 60J). Hind tibia without
peg-like hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Marginal tubercles present on ABD TERG 2-4. CAUDA with 10-19 hairs
(rarely less than 13). R IV+V more than 120 µm . . . . . . . . Aphis pomi
- Marginal tubercles usually absent from ABD TERG 2-4. CAUDA with
7-15 hairs (rarely more than 12). R IV+V less than 120 µm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
242 Malus
25 Dorsum of aptera mainly black. SIPH and CAUDA black, the CAUDA
having a rather pointed apex . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
- Dorsum of aptera pale. SIPH usually darker than CAUDA, which has a
rounded apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
26 Front of head pale or dusky. SIPH of aptera pale at least on basal half,
sometimes dusky towards apex . . . . . . . . . . . . Macrosiphum euphorbiae
- Front of head black. SIPH wholly black . . . . . . . . Macrosiphum rosae
27 Abdomen without marginal tubercles. SIPH more than 3.5 x CAUDA
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allocotaphis quaestionis
- Abdomen with marginal tubercles on most segments. SIPH less than
3.5 x CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28 Antennae of aptera at least as long as distance from frons to bases of
SIPH, and those of alata about as long as body. Aptera without pigmenta-
tion of abdominal tergites anterior to SIPH . . . . . Dysaphis plantaginea
- Antennae of aptera shorter than the distance from frons to bases of SIPH,
and those of alata less than body length. Aptera with dorsal abdominal
pigment spots or patches anterior to SIPH . . . . . . . . . . . . . . . . . . . . . 29
29 Colonies include many apterae (or alatiform apterae with sclerotized
thorax) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
- Colonies comprise mainly immature and adult alatae . . . . . . . . . . . . 31
30 Longest hairs on ANT III longer than basal diameter of segment. True
aptera (no sclerotization of thorax) with only sparse abdominal pigmenta-
tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis devecta
- Longest hairs on ANT III less than 0.5 x diameter of base of segment.
True aptera with broad dark bands or paired patches on all tergites . . . .
................................................ Dysaphis affinis
31 (Remaining couplets apply only to spring migrant alatae) Each antenna
with a total of 112-198 secondary rhinaria. Marginal tubercles usually
present on ABD TERG 1-7 inclusive, sometimes absent from ABD TERG
6. HT I with 3 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis brancoi
- Each antenna with a total of 20-113 secondary rhinaria. Marginal
tubercles present on ABD TERG 1-5, 1-5 and 7, or 1-7. HT I with 2 or
3 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 Ratio of total number of secondary rhinaria on both antennae to length
of SIPH (in µm) 1.12-1.64 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
- Ratio of total number of secondary rhinaria on both antennae to length
of SIPH (in µm) 0.26-1.00 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
33 Marginal tubercles frequently present on ABD TERG 1-7 inclusive.
Ratios of diameter of SIPH at midpoint to diameters of largest and
Malus 243
smallest marginal tubercles on ABD TERG 1-5 are 0.8-1.1 and 1.1-1.8,
respectively. ANT V with 0-6 secondary rhinaria . . . . . Dysaphis sibirica
- Marginal tubercles present on ABD TERG 1-5, or 1-5 and 7 (never on
6). Ratios of diameter of SIPH at midpoint to diameters of largest and
smallest tubercles on ABD TERG 1-5 are 1.3-2.1 and 2.7-6.0, respec-
tively. ANT V with 3-10 secondary rhinaria . . . . Dysaphis mordvilkoi
34 Posterior part of pronotum with a pair of pleural hairs . . . . . . . . . . 35
- Posterior part of pronotum without a pair of pleural hairs (although occa-
sionally one is present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
35 ABD TERG 8 with 7-10 hairs. Longest hair on ANT III 42-64 µm long,
1.6-2.7 x basal diameter of segment . . . . . . . . . . Dysaphis chaerophylli
- ABD TERG 8 with 4-5 hairs. Hairs on ANT III maximally 39µm, not
more than 1.7 x basal diameter of segment . . . . . . . . . . . . . . . . . . . . . 36
36 CAUDA with 6-9 hairs. Longest hair on ANT III is 13-18 µm, usually
shorter than basal diameter of segment . . . . . . . . . . . . . . Dysaphis flava
- CAUDA with 4-6 hairs, usually 5; rarely with 7. Longest hair on ANT
I I I is 9-39µm, longer or shorter than basal diameter of segment, but if
shorter then very b l u n t or dilated apically . . . . . . . . . . . . . . . . . . . . . . 37
37 Spinopleural hairs on ABD TERG 3 usually short and blunt, maximally
ll-23µm long. Longest hair on ANT III 9-29 µm, 0.4-1.3 x basal
diameter of segment. Marginal tubercles present on ABD TERG 1-7
inclusive (occasionally absent from one side on 6) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis radicola group
- Spinopleural hairs on ABD TERG 3 pointed, the longest being more than
25 µm long. Longest hair on ANT III in range 20-39 µm, 1.0-1.7 x basal
diameter of segment. Marginal tubercles present on ABD TERG 1-5, or
1-5 and 7; absent from 6 except rarely on one side only . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis anthrisci group
38 Each antenna with a total of 60-99 secondary rhinaria. R IV+V less than
1.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis affinis
- Each antenna with a total of 26-59 secondary rhinaria. R IV+V usually
more than 1.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39 Total number of 'spinal' tubercles (including those on head) 6-19. Each
antenna with a total of 37-59 (usually more than 46) secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis armeniaca
- Total number of 'spinal' tubercles (including those on head) 2-7. Each
antenna with a total of 26-46 secondary rhinaria . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis foeniculus malidauci
244 Mammea
MAMMEA Guttiferae
MARGARETTA Asclepiadaceae
MARKHAMIA Bignoniaceae
MAYTENUS Celastraceae
Macrosiphum euphorbiae
Myzus persicae
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)
MELALEUCA Myrtaceae
MELIA Meliaceae
MELIOSMA Meliosmaceae
MEMECYLON Melastomataceae
Brachycaudus helichrysi
Taiwanaphis memecyloni
MESUA Guttiferae
METASEQUOIA Taxodiaceae
METROSIDEROS Myrtaceae
METROXYLON Palmae
MICHELIA Magnoliaceae
Aphis spiraecola
Formosaphis micheliae
Myzus persicae
Prociphilus micheliae
Sinomegoura citricola
MICROCOS Tiliaceae
MILLETTIA Leguminosae
MIMOSA Leguminosae
MIMUSOPS Sapotaceae
MONOTES Dipterocarpaceae
MORINDA Rubiaceae
Aphis gossypii
MORINGA Moringaceae
MUNDULEA Leguminosae
MURRAYA Rutaceae
Aphis spiraecola
Myzus persicae
Sinomegoura citricola
Toxoptera aurantii, citricidus
(Use key to polyphagous aphids, p. 532)
Macrosiphum euphorbiae
Myzocallis myricae
M. rubra Aulacorthum solani
Parathoracaphis cheni
The small but very diverse aphid fauna of Myrica consists mainly of species
recorded from shrubs such as M. gale, but these are nevertheless included in
the following key for the sake of completeness.
8 SIPH dark over most or whole of length, with a small flange. HT II only
0.07-0.10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia borealis
SIPH pale or dusky, or only dark at apices, with a well-developed flange.
HT II 0.10-0.14 mm long . . . . . . . . . . . . . . . . . . . . . . Illinoia canadensis
MYRTUS Myrtaceae
Aulacorthum (Neomyzus)
circumflexum
Myzus persicae
Toxoptera aurantii
(See key to polyphagous aphids, p. 532)
NAUCLEA Rubiaceae
NECTANDRA Lauraceae
- ANT PT/BASE less than 1. Anal plate bilobed, CAUDA with an apical
knob. Body of apterae with numerous club-shaped tubercular processes
(Fig. 63B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lizerius tuberculatus
ANT PT/BASE more than 3. Anal plate entire, CAUDA tongue-shaped.
Body of aptera without club-shaped processes . . . . . Toxoptera aurantii
(or try key to polyphagous aphids, p. 532)
254 Neolitsea
NEOLITSEA Lauraceae
Although only these three species are recorded from Neolitsea, it seems pro-
bable that it may be colonized by other aphids which occur on Machilus and
Litsea. Lists and keys for those plant genera should therefore be consulted.
Fig. 62. Processes on ABD TERG 7 and 8 of aptera of A, Taiwanaphis furcifera (fundatrix),
B, T. niuginii, C, T. tasmaniae, D, T. nothofagi, E, Neosensoriaphis parva and F, Neuquenaphis chilensis',
processes on head of aptera of G, Neuquenaphis schlingen and H, Neuquenaphis essigi; processes on
ABD TERG 3 of aptera of I, Neuquenaphis essigi and J, N. sensoriata.
Nothofagus 257
- Antennae 0.6-1.1 x BL. Embryos with dorsal hairs less than 4µm long
(shorter than spinules on processes) . . . . . . . . . Neuquenaphis edwardsi
13 ANT PT/BASE 2.0-3.2. ANT III with 10-30 rhinaria. R IV+V with 7-12
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis sensoriata
- ANT PT/BASE 3.2-5.5, ANT III with 3-16 rhinaria. R IV+V with 4-8
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 ANT III with 8-16 rhinaria. Spinal processes on ABD TERG 1-3 pale and
mostly arising from unpigmented or only very weakly pigmented sclerites.
Forewing veins rather dark, the oblique veins beginning and ending in
conspicuous fuscous triangles (hand lens) . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis michelbacheri
- ANT III with 3-9 rhinaria. Spinal processes on ABD TERG 1-3 pig-
mented, arising from pigmented sclerites. Forewing veins with very little
fuscous at ends . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis edwardsi
NYSSA Cornaceae
OCHNA Ochnaceae
Sitobion ochnearum
OCHROMA Bombacaceae
Aphis gossypii
OCOTEA Lauraceae
Fig. 63. End of abdomen (ABD TERG 6-8) of aptera of A, Lizerius ocoteae and B, L. tuberculatus.
OSMANTHUS Oleaceae
OSTRYODERRIS = AGANOPE
OZOROA Anacardiaceae
PANAX Araliaceae
PANDANUS Pandanaceae
PAPPEA Sapindaceae
PARABENZOIN = UNDERA
PARINARIA Chrysobalanaceae
Aphis gossypii
PASANIA = LITHOCARPUS
PAULOWNIA Serophulariaceae
PELTOPHORUM Leguminosae
PERSEA Lauraceae
PHELLODENDRON Rutaceae
PHOEBE Lauraceae
PHOTINIA Rosaceae
2 Body and appendages with numerous long, fine hairs. SIPH pores large,
on hairy cones . . . . . . . . . . . . . . . . . . . . . . . . . . Nippolachnus bengalensis
Body and appendages with sparse, short hairs. SIPH pores indistinct or
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus sp(p).
4 SIPH short and conical; in aptera less than twice as long as their basal
widths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 CAUDA rounded, about as long as its basal width. R IV+V long, more
than twice as long as width at base . . . . . . . . . Brachycaudus helichrysi
CAUDA clearly longer than its width at base. R IV+V short; in alata
(emigrant) shorter that its basal width . . . . . . . . Melanaphis bambusae
266 Phyllanthus
6 SIPH dark, less than 1.5 x the long, pale, pointed CAUDA . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinomegoura photiniae
SIPH pale or dark only at tips, about twice length of CAUDA or more
............................................................. 7
7 SIPH with a subapical zone of polygonal reticulation. Alata without a
black dorsal abdominal patch . . . . . . . . . . . . . Macrosiphum euphorbiae
- SIPH without any polygonal reticulation. Alata with a black dorsal
abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 ANT III of aptera with a few rhinaria near base. ANT PT/BASE ratio
5 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbriaphis gentneri
- ANT III of aptera without rhinaria. ANT PT/BASE usually less than 4
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
PHYLLANTHUS Euphorbiaceae
PHYLLOSTACHYS Gramineae
Phyllostachys seems to have a much smaller aphid fauna than other genera
of bamboos, but it may nevertheless be worthwhile to consult the keys to
Arundinaria and Bambusa for specimens that do not fit the species listed here.
Astegopteryx bambucifoliae
Ceratoglyphina bambusae
Glyphinaphis bambusae
Melanaphis bambusae
Paracolopha morrisoni
Phyllaphoides bambusicola
Takecallis arundicolens,
arundinariae, sasae, taiwanus
Many of the 38 species of spruce-feeding Cinara have only been found on one
or two hosts, but the more widely-distributed and/or more common species
show little evidence of specificity to particular species of Picea, so a single,
rather long, key is required, of which couplets 36 onwards deal with this genus;
only the apterous viviparae are keyed. Many of the North American Cinara
species are inadequately known, and their separation often relies heavily on
the original descriptions. Cinara nepticula Hottes and Cinara yukona Hottes
are only known from the alate morph so cannot be included in the key. Several
other separations, especially those involving species not seen by the authors
(marked *), are probably unreliable because they are based on too few
specimens.
Picea is the primary host for Adelgidae. The taxonomy of adelgids is
difficult because of their complex polymorphism (see Fig. 1). Identification of
adelgids on spruce requires examination of immatures of the fundatrix genera-
tion (usually found overwintering in or near the buds) and/or the alate migrant
(gallicola) that emerges from the gall the following summer. Even then, species
limits are often uncertain (most notably in the laricis group). However, there
is some degree of host specificity in Adelgidae, so it should be possible to
narrow down the possibilities considerably by using the key in conjunction
with the host plant list below and the descriptions of galls, etc. in the text.
The parthenogenetic generations of species in a number of genera of
Pemphiginae in the tribe Prociphilini, Pachypappa, Pachypappella, Pro-
ciphilus, live in white wax on the roots of Picea spp. as their secondary hosts.
The couplets (29-35) separating apterae of Pachypappa and Pachypappella on
spruce roots are based on the work of Danielsson (1990a). The root-feeding
morphs of Prociphilus (subgenus Stagona) cannot at present be satisfactorily
keyed to species.
Pachypappella lactea
Pineus cembrae, konowashiyai,
pineoides
Prociphilus (Stagona) xylostei
Tsugaphis sorini ssp. piceicola
P. asperata Cinara alba, piceae
(Chinese Spruce)
P. brachytyla Adelges glandulae
P. engelmanni Adelges cooleyi
Cinara braggii, coloradensis,
?curvipes, engelmanniensis,
fornacula, glehna, hottesi,
nimbata, obscura, pruinosa,
saskensis, vandykei
Elatobium abietinum
Pineus boycei, engelmanni,
pinifoliae
Prociphilus (Stagona) xylostei
P. glauca Adelges abietis, cooleyi, lariciatus,
(White Spruce) pectinatae ssp. ishiharai, tardus
Cinara acadiana, atripes, ?bonica,
?bonita, braggii, coloradensis,
costata, ?curvipes, fornacula,
hottesi, jucunda, nigripes,
obscura, pallidipes, piceae,
piceicola, pilicornis, pinicola,
pruinosa, saskensis, soplada,
thatcheri, vandykei, [yukona]
Essigella alyeska
Mindarus obliquus
Pachypappa rosettei, vesicalis
Pineus pinifoliae
Prociphilus (Stagona) xylostei
Tsugaphis sorini ssp. piceicola
P. glehnii Adelges pectinatae ssp. ishiharai
(Sakhalin Spruce) Cinara costata, ezoana, glehna,
nopporoensis, piceae, pinicola
Pineus cembrae
Prociphilus (Stagona) konoi
P. jezoensis (= ajanensis) Adelges abietis, japonicus,
(Yeddo Spruce) karafutonis, lapponicus,
pectinatae ssp. ishiharai
Cinara costata, ezoana, horii,
?piceae, pruinosa
Picea 271
Elatobium momii/piceana
Pineus orientalis
Tsugaphis sorini ssp. piceicola
P. jezoensis var. hondoensis Adelges isedakii, torii
P. koraiensis Adelges laricis, pectinatae, viridis
Tsugaphis sorini ssp. piceae
P. koyamai Adelges abietis
P. likiangensis Adelges glandulae
Pineus sichunanus
P. mariana Adelges cooleyi, lapponicus,
(Black Spruce) lariciatus, lands
Cinara abietis, braggii, fornacula,
hottesi, mariana, nigripes,
nimbata, pruinosa, rara
Pachypappa rosettei, ?sacculi
Pineus floccus, pinifoliae, strobi
P. morrisonicola [Prociphilus formosanus]
P. obovata Adelges lapponicus, lands,
(Siberian Spruce) pectinatae, tardus
Cinara piceicola
Pineus cembrae, pineoides
[Prociphilus oriens]
P. omorika Adelges nordmannianae, prelli
Pineus pineoides
P. orientalis Adelges lands, merkeri,
nordmannianae, prelli, viridis
Cinara pilicornis, pruinosa
Pachypappa tremulae
Pineus orientalis
P. polita Adelges funitecta
(Tigertail Spruce) Pineus orientalis
P. pungens Adelges cooleyi, lapponicus,
(Colorado Spruce) lariciatus, lands, tardus
Cinara braggii, coloradensis,
costata, fornacula, glehna,
jucunda, piceae, pilicornis,
pruinosa, [wanepae]
Elatobium abietinum
Pachypappa ?sacculi, tremulae
Pineus pinifoliae, similis
Protohormaphis piceae
Tsugaphis sorini ssp. piceicola
272 Picea
Fig. 64. A, adult aptera of Pineus pineoides, dorsal (right) and ventral views; dorsal abdominal wax glands
of alate gallicola of B, Pineus boycei (after Annand, 1928) and C, P. orientalis; antenna of alate gallicola of
D, Pineus floccus (after Annand, 1928), E, P. orientalis, F, Adelges laricis, G, A. cooleyi; H, overwintering
fundatrix larva of Adelges (subgenus Dreyfusia) showing positions of wax glands on spinal sclerites, with
the 'central areas' arrowed (total number of pits in these on both sides is counted); meso- and metathoracic
spinal sclerites (on left side only), showing shape and arrangements of wax glands in overwintering fundatrix
larva of I, Adelges nordmannianae, J, A. merkeri and K, A. prelli (all after Eichhorn, 1956); L, pattern of
wax glands on thoracic sclerites of overwintering fundatrix larva of Adelges japonicus (after Inouye, 1953);
wax glands on prothorax of overwintering fundatrix larva of M, A. abietis and N, A. viridis.
276 Picea
- Immature fundatrix with wax pore plates on inner margins of meso- and
metathoracic spinal sclerites divided into 3-4 often rather angular areas,
containing relatively large pits of irregular shape (Figs 64J, K). Total
number of pits in the most central areas only of the spinal wax pore plates
of meso- and metathorax plus ABD TERG 1-3 (10 central areas) is 15-56
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Immature fundatrix with total number of pits in 10 central areas (as
defined in couplet 12) usually more than 40 (range 34-56, mean about 45)
(Fig. 64J) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges merkeri
Fig. 65. Dorsal (right) and ventral aspects of apterous vivipara of A, Protohormaphis piceae and
B, Tsugaphis sorini piceicola (both after Shaposhnikov and Gabrid, 1987).
Fig. 66. Fore tarsus of apterous exule of A, Pachypappa tremulae, B, Pachypappella lactea and
C, Pachypappa vesicalis.
- Aptera with R IV+V about 1.0 x ANT BASE VI or longer, and with
marginal wax pore plates reduced or absent, at least on ABD TERG 3-8;
large spinal and pleural wax pore plates on ABD TERG 4-7. Alata
(sexupara) with media of forewing usually unbranched (on roots) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 R IV+V without a pale subapical zone. Aptera with thick, spine-like hairs
on HT I, similar in size to those at apex of tibia. Alate sexupara with
secondary rhinaria on ANT V . . . . . . . . . . . Prociphilus (Stagona) spp.
- R IV+V with a distinct pale subapical zone. Aptera with hairs on HT I
fine or spine-like, if spine-like then often smaller than those at apex of
tibia. Alate sexupara without secondary rhinaria on ANT V . . . . . . 30
30 (Remaining couplets in this part of key refer only to aptera.) Hairs on HT
I very small and thin, or if thick basally then with finely-pointed apices
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
- Hairs on HT I thick and sometimes spine-like, with short or blunt apices
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
31 Segments I and II of all tarsi distinctly separated from each other. Seg-
ment I of fore tarsus with 2-3 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
- Segments I and II of tarsi not distinctly separated. Segment I of fore tarsus
with 2 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
32 Hairs on HT I and apical dorsal hairs on HT II very small, 4-8 µm long,
with abrupt apices. Segment I of fore tarsus usually with 2 hairs; if a third
middle hair is present it is longer than the lateral hairs . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa rosettei*
- Hairs on HT I and apical dorsal hairs on HT II longer, 10-15 µm, with
finely-pointed apices. Segment I of fore tarsus almost always with 3 hairs,
the middle hair shorter and blunter than the lateral hairs (Fig. 66A)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa tremulae
280 Picea
Fig. 67. Hairs on outer side of hind tibia of aptera of A, Cinara piceae, B, C. piceicola and C, C. pilicornis.
33 ANT PT only 0.015-0.02 mm long. Legs short, with length of hind femur
less than 4 x its maximum width. All tarsi reduced, with segments I and
II not distinctly separated (HT II less than 0.1 mm long). Second tarsal
segments with dorsal apical hairs short and spine-like (Fig. 66B) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappella lactea
- ANT PT more than 0.02 mm long. Legs longer, with hind femur more
than 4 x its maximum width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34 Antenna usually 6-segmented, with PT finger-like, 0.034-0.05 mm long.
Hairs on first tarsal segments spine-like and similar in size to those at
apices of tibiae. Hind tibia on dorsal side bearing 2-5 spine-like hairs with
short, blunt apices . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa populi*
- Antenna usually 5-segmented, with PT less than 0.035 mm long. Hairs on
first tarsal segments distinctly shorter than those at apices of tibiae (e.g.
Fig. 66C). Hind tibia with some hairs on dorsal side with long, pointed
apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35 Hairs arising from dorsal abdominal wax pore plates are more than 25 µm
long. Each wax gland facet with a dark circle inside a pale outer ring
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa vesicalis
- Hairs arising from abdominal wax pore plates are less than 25 µm long.
Wax gland facets without dark centres . . . . . . . . ?Pachypappa sacculi*
36 (Remaining couplets all refer only to apterous viviparae.) Hairs on outer
side of hind tibia short and rather thick (e.g. Fig. 67A), only more than
0.06mm long in very large aphids, mostly less than or not exceeding
0.5 x width of hind tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . 37
- Hairs on outer side of hind tibia more than 0.06 mm long and much more
than 0.5 x width of hind tibia at midpoint . . . . . . . . . . . . . . . . . . . . . 41
37 Large rotund aphid, BL 3.2-6.7 mm long, but usually more than 4.5 mm.
Coxae black. R IV more than 0.30mm . . . . . . . . . . . . . . Cinara piceae
Picea 281
- BL 2.4-4.2 mm. Coxae (where colour known) light brown. R IV less than
0.25 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38 PT less than 0.03 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara jucunda*
- PT more than 0.04 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39 R IV only about 0.17 mm (in specimen of BL about 2.5 mm) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara caudelli*
- R IV more than 0.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40 HT II about 0.29mm (in specimen of BL 3.1 mm). Hairs on SIPH cones
rather evenly distributed over entire pigmented area . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara soplada*
- HT II 0.33 mm or more. Hairs on SIPH cones concentrated on apical half
around pore, with basal part of pigmented area having only sparse hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara coloradensis
41 Hairs on outer side of middle section of hind tibia all less than 0.12mm
long (e.g. Fig.67B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
- All or many of hairs on outer side of middle section of hind tibia of length
exceeding 0.12 mm (e.g. Fig. 67C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
42 Hind tibiae uniformly dark brown to black . . . . . . . . . Cinara atripes*
- Hind tibiae with at least the basal or middle section paler . . . . . . . . 43
43 SIPH cones pale, almost concolorous with dorsal cuticle . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara fornacula
- SIPH cones pigmented, much darker than rest of dorsal cuticle . . . 44
44 HT II more than 0.32 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
- HT II less than 0.32 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
45 BL less than 4.6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara piceicola
- BL more than 5.0mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
46 R IV about 0.22mm. Antennae and legs short; ANT III less than
0.1 x BL, hind tibia less than 0.4 x BL . . . . . . . . . . . . . Cinara bonita*
- R IV 0.40-0.44 mm. Antennae and legs much longer; ANT III more than
0.15 x BL, hind tibia more than 0.6 x BL . . . . . . . . Cinara saskensis*
47 Hind tibia more than 1.65 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
- Hind tibia less than 1.65 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
48 R IV+V about 0.42 mm . . . . . . . . . . . . . . . . . . . . . . . . . Cinara acadiana*
- R IV+V 0.36 mm or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
49 R IV+V 0.24mm or less. ANT III with 1-2 secondary rhinaria distally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara vandykei
282 Picea
59 Longest hairs on ANT III about one quarter of length of ANT III. (Alata
with forewings marked with fuscous) . . . . . . . . . . . . . . . Cinara nimbata
- Longest hairs on ANT III about one half of length of ANT III. (Alata
with hyaline forewings) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara comata
60 R IV+V 1.1-1.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pruinosa
- R IV+V less than 1.1 x HT II ................................ 61
61 ANT III 3.4-4.1 x longer than the longest hair borne upon it. HT II
0.30-0.48 mm long, 1.1-1.4 x R IV+V. Coxae, femora and tibiae all
rather pale; tibiae only dark at apices . . . . . . . . . . . . . . . Cinara braggii
- ANT III 2.4-3.1 x the longest hair borne upon it. HT II 0.28-0.32 mm
long, 1.0-1.2 x R IV+V. Legs darker, with tibiae dark on about distal
third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
62 ANT III more than 0.15 x BL, bearing hairs up to 160µm long . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara glacialis*
- ANT III 0.08-0.13 x BL, with longest hairs 90-130 µm long . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara glehna/mariana*
PILGERODENDRON Cupressaceae
There are about 170 pine-feeding aphids, including more than 100 species of
Cinara. A single key including all the Cinara species would be unwieldy and
difficult to use. Fortunately there is a fairly well-established subgeneric classi-
fication of Pinus to which many (but by no means all) pine-feeding Cinara
species conform. We have used this subgeneric structure wherever possible to
key together Cinara species that feed on related pines. We have also made use
of it for separation of some closely related and morphologically similar species
of Essigella, a wholly North American genus of Lachnidae recently revised
using multivariate techniques by Sorensen (1992).
The main key that follows goes as far as possible with the identification
of all pine-feeding aphids to species level, except the genera Cinara and
Essigella. In the case of the adelgid genus Pineus, the secondary host morphs
(sistens, progrediens) on Pinus are very difficult to identify because of their
284 Pinus
lack of species-constant characters, and even the partial key provided should
be treated very warily.
The separate species of Pinus are then listed with the aphids recorded from
them, each followed where necessary by keys to Cinara and/or Essigella.
These keys can only be applied to apterous viviparae.
Fig. 68. A, CAUDA and anal plate of aptera of Prociphilus piniradicivorus; B, CAUDA of aptera of
P. pini; hind tibia of aptera of C, P. crataegistrobi and D, P. pini.
with 0-4 secondary rhinaria on ANT VI BASE, and with distal end of
pterostigma tapering, curved around tip of wing) . . . . . . . . . . . . . . . . . 8
8 R IV+V with 2 accessory hairs. Hind tibia thick, with 1-3 short, spine-like
hairs near middle on outer side (Fig. 68C). (Alate sexupara with 1-4 secon-
dary rhinaria on ANT VI BASE) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus (Stagona) crataegistrobi
- R IV+V with 3-4 accessory hairs. Hind tibia narrower, with several hairs
distributed along outer side which are fine-pointed, not distinctly spine-
like (Fig. 68D). (Alate sexupara without secondary rhinaria on ANT VI
BASE) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus (Stagona) pini
9 SIPH as large pores on usually pigmented hair-bearing cones . . . . . 10
- SIPH as small pores without associated hairs . . . . . . . . . . . . . . . . . . . 17
10 R V acutely pointed, dagger-shaped, usually twice or more as long as its
basal width (e.g. Fig. 69A). SIPH cones of aptera often large and with
numerous hairs . . . . . Cinara spp. (see separate keys in Host Plant List)
- R V short, flask-shaped, pointed only at tip, similar in length to or not
much longer than its basal width (e.g. Figs69B, C). SIPH cones always
with few hairs, in 1-3 rings around pore . . . . . . . . . . . . . . . . . . . . . . . 11
11 Hairs on ventral side of hind femur very short, thick and peg-like
(Fig. 70A). Hind legs wholly dark except for extreme base of femur, and
very elongate; total length about 2 x BL . . . . Schizolachnus flocculosus
- Hairs on ventral side of hind femur long, not peg-like. Hind legs long but
less than 2 x BL, with at least basal fifth of femur pale . . . . . . . . . 12
286 Pinus
Fig. 70. A, ventral side of hind femur of Schizolachnus flocculosus (aptera); B, middle section of hind
tibia of Schizolachnus curvispinosus (aptera); C, hair-bearing scleroites on ABD TERG 3 of
S. curvispinosus (aptera); ANT VI of D, S. pineti and E, S. parvus.
12 Hind tibia wholly dark bearing long, thick, curved hairs with blunt apices
(Fig. 70B). Dorsal abdominal hairs thick and spine-like, arising from
transverse rows of small sclerites (Fig. 70C) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizolachnus curvispinosus
- Hind tibia pale or dark, bearing mainly long, finely-pointed hairs. Dorsal
abdominal hairs also long and fine, not arising from sclerites ..... 13
13 Hind tibia more than 20 x longer than its width at midpoint ........
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizolachnus piniradiatae
Pinus 287
Fig. 71. Arrangement of spinopleural hair-bearing scleroites between muscle plates on ABD TERG 3:
A, Eulachnus nigricola; B, Eu. brevipilosa; C, Eu. tuberculostemmata; D, Eu. thunbergii; E, Eu. rileyi; f, Eu. agilis.
Pinus 289
- Several of longest hairs on ANT I I I more than 40µm long. ABD TERG
1-3 with longest hairs 25-145 µm long, but if under 40 µm then they are
often blunt or slightly capitate and dark like those on ABD TERG 7
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 Hair-bearing scleroites on ABD TERG 1-5 numerous, in two irregular
transverse rows on each tergite, the anterior row having more scleroites
than the posterior row (Fig. 71D). Fore femur dusky or dark and very
thick; its length less than 3.5 x its maximum width . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulachnus thunbergii
- Hair-bearing scleroites on ABD TERG 1-5 often in two rows, but then
the anterior 'row' often has very few scleroites (often only 2), and never
has more than the posterior row. Fore femur pale or dark, more than
3.5 x longer than its maximum width . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 ABD TERG 3 with longest hairs 100-155 µm, usually with finely pointed
apices (Fig. 71E). ANT V usually clearly longer than (1.1-1.5 x) HT II.
Hind femur uniformly dark except at base; much darker than basal half
of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulachnus rileyi
- ABD TERG 3 with longest hairs 25-110 µm, and usually with blunt apices
(Fig.71F). ANT V usually shorter than (0.6-1.2 x ) HT I I . Hind femur
pale or mottled, mainly about as dark as basal half of ANT I I I . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulachnus agilis
24 Claws normal, with simple curved acute apices. ABD TERG 2-7 mem-
branous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudessigella brachychaeta
- Claws with modified apices; bifurcate, or double (Fig. 73). ABD TERG
2-7 lightly to heavily sclerotized (but not necessarily pigmented) . . . . .
. . . . . . . . . . . . . . . . Essigella spp. (see separate keys in Host Plant List)
25 Cephaloprothoracic shield (henceforth CPS) with sutures between head
and thorax and along midline. Wax glands on CPS comprising mainly
separate, large facets that are distinctly subdivided (Fig. 72A) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus pinifoliae
- CPS entire. Wax glands on CPS consisting mainly of groups of facets that
are not individually subdivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Wax glands on anteriodorsal part of CPS (i.e. between eyes) mainly com-
prising groups of 2-10 facets that are often loosely grouped or at least
not closely pressed together, so that all except the largest retain a more-or-
less circular shape (e.g. Figs 72B, C) . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
- Wax glands between eyes mainly comprising more extensive groups or net-
works of 10 to more than 100 closely packed facets, mostly pressed
together so that they tend to be polygonal in shape (e.g. Figs. 72D, E) or
look like overlapping tiles when viewed obliquely . . . . . . . . . . . . . . . . 30
27 Wax glands between eyes comprising groups of 2-6 large facets that are
mostly much larger than the ocelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
290 Pinus
Fig. 72. Dorsal (right) and ventral aspects of A, Pineus pinifoliae (after Annand, 1928) and B, P. pini,
showing details of wax gland structure and (for pini) antenna; wax gland facets mesiad to ocelli of
C, P. boerneri, D, P. cembrae and E, P. strobi.
Fig. 74. A, rostrum of Cinara showing measurement of sclerotized part of stylet groove; B, SIPH of
Cinara anzai (aptera); C, HT I of C. atra; 0, SIPH of C. pinona (aptera); hairs on dorsal side of hind tibia
of aptera of E, C. apacheca and F, C. edulis.
BL less than 4.5 mm. Tibiae with paler section on basal half. Length of
sclerotized part of stylet groove less than 2.4 mm . . . . . . . . . . . . . . . . . 2
2 Hairs on dorsal side of hind tibia only about 0.5 or less of width of tibia
at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Hairs on dorsal side of hind tibia of length similar to width of tibia at
midpoint, or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 ABD TERG 1-7 each with a pair of irregularly-shaped or broken pig-
mented areas, of similar extent on all tergites . . . . . . . . . . . . inscripta*
At least ABD TERG 3-6 without pigmented sclerites . . . . . . . . . . apini
4 SIPH bases rather small (maximum diameter less than 0.35 mm) and very
asymmetrical, extending about twice as far anteriorly as posteriorly
(Fig. 74B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . anzai
SIPH bases large and fairly symmetrical; maximum diameter more than
0.4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 BL 4.0-4.4 mm. R V very short, 0.06 mm or less, only about one third
of length of R IV. HT II about 0.4mm . . . . . . . . . . . . . . . . . . oregoni*
- BL 2.6-3.8 mm. R V 0.08-0.10 mm, more than one third of length of
Pinus 293
Key to Cinara spp. on Pinus cembra and other pines of subsect. Cembrae
(koraiensis, pumila, sibirica)
1 Hairs on ABD TERG 5 long (80-180µm). ANT III shorter than IV + V
together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Hairs on ABD TERG 5 short (4-40 µm). ANT III as long as or longer
than IV + V together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Head, pronotum and mesonotum dark. Total antennal length about
0.45 x BL. ABD TERG 8 with about 30 hairs . . . . . . . . . . . mongolica
Head and thorax pale dorsally. Total antennal length about 0.35 x BL.
ABD TERG 8 with about 20 hairs . . . . . . . . . . . . . minoripinihabitans*
3 R IV 0.14-0.22 mm long, about 2 x R V. Hairs on ANT VI BASE max-
imally about 1.5 x basal diameter of segment . . . . . . . . . . . . . . . . . pini
- R IV 0.21-0.29 mm, more than 2 x R V. Hairs on ANT VI BASE
maximally about 2 or more x basal diameter of segment . . . . . . . . . 4
4 R IV less than 0.25 mm long, bearing 8-10 accessory hairs . . . . cembrae
R IV more than 0.25 mm long, bearing 14-22 accessory hairs . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . watanabei
Key to Cinara spp. on Pinus cembroides and other pines of subsect. Cem-
broides (edulis, monophylla, quadrifolia)
Parts of this key are based on Hottes' (1960b) key to Cinara species on
P. edulis. Most of the Cinara species described from pinyon pines are based
on inadequate material, and further study is likely to reveal synonymies.
l R IV more than 0.28mm long, with more than 25 accessory hairs . . . 2
- R IV less than 0.27 mm long, with less than 25 accessory hairs . . . . 3
Pinus 295
2 R IV 0.5 mm or more long, much longer than ANT III, with more than
70 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . puerca
- R IV less than 0.4 mm long, shorter than ANT III, with less than 40 acces-
sory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tanneri
3 Diameter of SIPH base 0.25 mm or less . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Diameter of SIPH base 0.27 mm or more . . . . . . . . . . . . . . . . . . . . . . . 5
4 BL 1.6-2.2 mm. Legs with contrasting pale and dark sections . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . caliente
- BL about 2.6mm. Legs dark, without pale sections . . . . . . . . . poketa*
5 Ventral hairs on HT I longer than maximum width of segment, with apices
attenuated, the more distal hairs often having their apices sharply crooked
(Fig.74C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atra
- Ventral hairs on HT I mostly not longer than maximum width of HT I,
or if slightly longer then without attenuated and crooked apices . . . . . . 6
6 Hind tibia densely clothed in hairs (more than c. 300 per mm), with those
on dorsal side mostly arising at an angle of less than 45° and curved so
that distally they are almost parallel to the tibia (Figs 74E, F) . . . . . 7
- Hind tibial hairs usually less dense, but if at a density of more than
c. 300 per mm then those on dorsal side arise at an angle of 45° or more
and with apices pointing away from tibia . . . . . . . . . . . . . . . . . . . . . . . 10
7 Hairs on hind tibia short and spine-like (Fig. 74E), maximally about 50 µm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . apacheca
- Hairs on hind tibia fine (e.g. Fig. 74F), maximally more than 50 µm long
............................................................. 8
8 SIPH cone uniformly pigmented, with maximum diameter 0.3-0.5 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . edulis
- SIPH cone two-toned, with only the upper, constricted, part deeply
pigmented (e.g. Fig. 74D); maximum diameter about 0.35mm . . . . . 9
9 Length of pale section on basal half of hind tibia less than one quarter
of total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinona
- Length of pale section on basal half of hind tibia more than one quarter
of total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . metalica*
10 Hairs on hind tibia small and fine, maximally about 85 µm long, with
finely pointed apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Hairs on hind tibia longer or, if less than 85 µm long, then coarse with
blunt or abruptly acute apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Hairs on dorsal side of hind tibia mainly standing almost at right angles
to tibia. ANT VI BASE with less than 12 hairs . . . . . . . . . . . . rustica*
296 Pinus
- Hairs on dorsal side of hind tibia set at a more acute angle to tibia. ANT
VI BASE with more than 12 hairs . . . . . . . . . . . . . . . . . . . . . . . . . pinata
12 ANT III less than 0.33 mm long, much shorter than maximum diameter
of SIPH cone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nitidula
- ANT III more than 0.35 mm long, usually about as long as or slightly
longer than maximum diameter of SIPH cone . . . . . . . . . . . . . . . . . . 13
13 ABD TERG 5-7 (or 6-7) each with a single pair of small dark sclerites.
Hairs on hind tibia maximally 70-150 µm . . . . . . . . . . . . . . . . . wahtolca
- ABD TERG 5-7 with or without sclerites, but not each with a single pair.
Hairs on hind tibia maximally 50-80 µm . . . . . . ponderosae/terminalis
Key to Cinara spp. on Pinus contorta and other pines of subsect. Contortae
(banksiana, clausa, virginiana)
1 Dorsal abdomen almost completely covered by an extensive pigmented
area that is fused at least between tergites 4 and 6 and usually merges with
SIPH bases (Figs 75A, B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Dorsal abdomen with any dark markings divided segmentally and not
usually merged with SIPH bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 Subgenital plate (of vivipara) with only 6-14 hairs. R IV with only 4-7
accessory hairs. SIPH with only 10-12 short hairs (maximally 40 µm)
....................................................... russellae
- Subgenital plate with 30-40 hairs. R IV with 8-16 accessory hairs. SIPH
with 20-35 longer hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 ANT VI BASE with 10-15 hairs. R IV with 10-16 accessory hairs,
including several on side away from stylet groove . . . . . . . . . . . gracilis
Pinus 297
- ANT VI BASE with 8-11 hairs. R IV with 7-11 accessory hairs in two
rows alongside stylet groove, rarely with 1-2 on opposite side . . . . . 4
4 Dorsal cephalic hairs rather sparse, situated mainly alongside dorsal
suture and near antennal bases, maximally 60 µm long, often shorter than
the distances between their bases. SIPH cones shallow with their bases
very extensive, often extending over half the length of the abdomen
(Fig.75A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . canatra
- Dorsal cephalic hairs numerous, rather evenly distributed, maximally
70-80 µm long, much longer than the distances between their bases. SIPH
cones higher, with bases much less extensive (Fig. 75B) . . . . . . . . nigra
5 HT I (measured along ventral side) at least 0.5 x HT II (e.g. Fig. 76B).
Longest hairs on dorsal side of hind tibia 110-230 µm long, 1.5 or more
x width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- HT I less than 0.5 x HT II. Longest hairs on dorsal side of hind tibia
30-150 µm long, rarely up to 1.5 x middle width of tibia . . . . . . . . . 9
6 ABD TERG 2-7 each with a variably developed but usually large pair of
dark sclerites (or diffuse dusky areas) including the bases of several hairs.
HT II 0.20-0.28 mm. Eyes sessile, always with a distinct ocular tubercle
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinivora
- ABD TERG 2-7 with separate small dark hair-bearing sclerites (scleroites).
HT II 0.27-0.53 mm. Eyes on short lateral projections of head, usually
without any evident ocular tubercle (e.g. Fig. 75G) . . . . . . . . . . . . . . . 7
7 Fore tibia uniformly dark, mid- and hind tibia with pale basal sections.
First tarsal segments usually with 2 sense pegs, one behind the other
(Fig. 75H). Largest scleroites on ABD TERG 2-5 of greatest diameter
25-70 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . watsoni
- If the fore tibia is uniformly dark, then so are the mid- and hind tibiae.
First tarsal segments with 1 sense peg. Largest scleroites on ABD TERG
2-5 of greatest diameter 75-300 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 All tibiae with pale section on basal half. First tarsal segments usually with
1-2 dorsal hairs, and with the sense peg some distance from the apex.
ABD TERG 5 with 28-49 hairs between SIPH bases, most of them on
large scleroites (Fig. 80C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinea
- Tibiae usually rather uniformly dark (sometimes slightly less so near base).
First tarsal segments without dorsal hairs, and with sense peg close to
apex. ABD TERG 5 with 50-110 hairs between SIPH bases, a minority
of which are on large scleroites (Fig. 80D) . . . . . . . . . . . . . . . pergandei
9 Hairs on dorsal side of hind tibia near its midpoint mostly less than 60 µm
in length, rarely up to 70 µm, the longest of them 0.3-0.8 x diameter of
tibia at this point . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Most or all of hairs on dorsal side of hind tibia near its midpoint longer
298 Pinus
Fig. 75. Pattern of dorsal sclerotization of aptera of A, Cinara canatra, B, C. nigra, C, C. brevispinosa
(spring), D, C. contortae, E, C. banksiana, f, C. medispinosa (spring); G, stalked eye of C. pinea;
H, HT I of C. watsoni; hairs on dorsal side of middle part of hind tibia of aptera of I, C. murrayanae
and J, C. atlantica.
Pinus 299
than 60 µm, the longest of them 0.7-1.5 x diameter of tibia at this point
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
10 ABD TERG 4-6 with many scleroites of width more than 50 µm, or
bars or patches forming part of a more extensive pattern of markings
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- ABD TERG 5 and 6, and usually 4 also, without any bars or patches or
scleroites more than 4 x wider than the hair bases . . . . . . . . . . . . . . 13
11 Length of sclerotized part of stylet groove (Fig. 74A) 1.55-1.76 mm.
Dorsal abdominal markings comprising paired patches of irregular outline
on ABD TERG 1-8, sometimes reduced or broken into groups of sclerites
on 2-6 (Fig. 75D). All tibiae with pale section on basal half . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . contortae
- Length of sclerotized part of stylet groove 1.25-1.54 mm. Dorsal abdomen
either with broad transverse bands on all tergites or with paired patches
on ABD TERG 1-3 and scattered sclerites on 4-5. Fore and hind tibiae
often wholly dark, or with shorter pale section than mid-tibia . . . . . . 12
12 Dorsal abdomen usually (in spring, at least) with a complete pattern of
broad transverse bars on all tergites (Fig. 75C); if reduced on ABD TERG
4-6 then the sclerites are still arranged in single transverse rows across the
midline. Hairs on dorsal side of hind tibia short and spine-like, all less
than 50 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . brevispinosa
- Dorsal abdomen with well-developed paired patches on ABD TERG 1 and
2 (or 1-3) and 7; only scattered small sclerites on 4-5, and usually none
at all on 6 (Fig. 75E). Hairs on dorsal side of hind tibia variable, but at
least some are usually more than 50 µm long . . . . . . . . . . . . . banksiana
13 Length of sclerotized part of stylet groove (see Fig. 74A) 1.9 mm or more
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Length of sclerotized part of stylet groove less than 1.8mm . . . . . . 15
14 PT finger-like, parallel-sided, 0.07-0.09 mm long. Length of sclerotized
part of stylet groove 2.3-3.2 mm . . . . . . . . . . . . . . . . . . . . . . . piniradicis
- PT tapering, less than 0.06mm long. Sclerotized part of stylet groove
1.9-2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . schwartzii
15 Length of sclerotized part of stylet groove 1.12-1.37 mm . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ontarioensis
- Length of sclerotized part of stylet groove 1.45-1.65 mm . . . . . taedae
16 Hairs on SIPH cone sparse and very short, the longest about 15 µm long
(only the sexuales of this species are described) . . . . . . . . . parvicornis*
- Hairs on SIPH cone more than 50 µm long . . . . . . . . . . . . . . . . . . . . . 17
17 R IV with 15-28 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
300 Pinus
Key to Cinara spp. on Pinus coulteri and other pines of subsect. Sabinianae
(sabiniana, torreyana)
1 BL less than 4.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
BL more than 4.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Longest hairs on ANT III 120-200µm, and on hind tibia 140-310 µm
..................................................... tujafilina*
Longest hairs on ANT III 30-60 µm, and on hind tibia 45-80µm . . . . . 3
3 R IV 0.25-0.30 mm. SIPH less than 0.2mm in diameter, bearing a few,
short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . diabola*
R IV 0.17-0.22 mm. SIPH more than 0.5mm in diameter, bearing many
(25 +) long hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae
302 Pinus
4 Hairs on dorsal body surface, ANT III-IV and legs all rather sparse,
short, thick and slightly capitate. ANT III longer than IV + V together
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . arizonica
Hairs not capitate; those on hind tibia short (about 50 µm) and dense.
ANT III shorter than IV + V together . . . . . . . . . . . . . . . . . . montanesa
Key to Cinara spp. on Pinus densiflora and other oriental pines of subsect.
Sylvestres (massoniana, tabulaeformis, taiwanensis, thunbergii)
If the outcome of using this key is unsatisfactory, try the key to western
palaearctic species under Pinus sylvestris
1 HT I elongate, 0.16-0.33 mm long (measured on ventral side), at least
0.5 x HT II or longer (Fig. 76B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- HT I less than 0.17 mm long, less than 0.5 x HT II . . . . . . . . . . . . . . 5
2 Hairs on dorsal body and appendages short and thick; longest hairs on
ANT III 55-65 µm, on ABD TERG 3 20-40 µm, on hind tibia 75-85 µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . formosana
Pinus 303
- Hairs much longer; longest on ANT III 90-210 µm, on ABD TERG 3
80-210 µm, on hind tibia 90-230 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Tibiae with pale section on basal half. HT I with 1-2 dorsal hairs
(Fig. 76B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinea
Tibiae wholly dark, although sometimes slightly less dark near base. HT
I usually without dorsal hairs, rarely with 1 . . . . . . . . . . . . . . . . . . . . . 4
4 ABD TERG 5 with less than 50 hairs between SIPH bases, the longest
80-160 µm. Longest hairs on hind tibia 80-160 µm . . . . . piniformosana
ABD TERG 5 with more than 50 hairs between SIPH bases, the longest
120-200 µm (Fig. 80D). Longest hairs on hind tibia 150-190 µm . . . . . .
...................................................... pergandei
5 R IV 0.44-0.54 mrn long, about as long as ANT III or longer, bearing
28-35 accessory hairs in 4 rows. Longest hairs on dorsal side of hind tibia
shorter than or similar to width oi tibia at midpoint . . . . . . . . . . . . . 6
R IV 0.14-0.22 mm long, less than half as long as ANT III and bearing
7-13 accessory hairs. Longest hairs on dorsal side of hind tibia 1.5 or more
x width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6 Hairs on dorsal side of hind tibia small, fine and curved distally so that
they are often almost parallel to tibia; maximally 60-70 µm long, about
0.5 x width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . sorini
Hairs on dorsal side of hind tibia fine but erect, the longest 70-80 µm long,
about equal to width of tibia at midpoint . . . . . . . . . . . . . . . . . . etsuhoe
7 ABD TERG 1-2 (or 1-3) with marginal sclerites as well as spinal ones
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . orientalis
- ABD TERG 1-3 without marginal sclerites . . . . . . . . . . . . . . . . . . . . . . 8
8 HT II less than 0.3 mm long. ABD TERG 1-7 each with a pair of large
spinal sclerites (sometimes reduced or absent particularly on 2-3). Hind
tibia pale on basal half, contrastingly dark distally . . . . . . . . . . . . . . . . .
................................................. pinidensiflorae
HT II more than 0.3 mm long. ABD TERG 2-6 with only small sclerites.
Hind tibia mainly or wholly dark . . . . . . . . . . . . . . . . . . . . . . californica
Key to Cinara spp. on Pinus echinata and other pines of subsect. Australes
(caribaea, elliottii, glabra, occidentalis, palustris, pungens, rigida, serotina,
taeda)
1 HT I (measured on ventral side) at least 0.5 x HT II (Fig. 76A, B)
............................................................. 2
- HT I less than 0.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Longest hairs on dorsal side of hind tibia 70-130 µm, similar to width of
tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maritimae
- Longest hairs on dorsal side of hind tibia 110-230 µm, 1.5 or more x
width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . 4 species (pergandei, pinea, pinivora and watsoni)
thai are separated in couplets 6-8 of the key under P. contorta
3 BL 4.3-6.1 mm. Hairs on dorsal body surface, ANT II1-IV and legs all
short, thick and slightly capitate . . . . . . . . . . . . . . . . . . . . . . . . . arizonica
- BL up to 4.2 mm. Hairs short or long, not capitate . . . . . . . . . . . . . . 4
4 R IV 0.23-0.33 mm long, bearing 19-28 accessory hairs . . . . . . . . . . . 5
- R IV 0.16-0.23 mm long, bearing 6-14 accessory hairs . . . . . . . . . . . . 6
5 ANT VI BASE with 11-16 hairs. Longest hairs on hind tibia 80-110 µm.
Dorsal abdominal hairs long, many of them arising from sclerites of vary-
ing size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cronartii
- ANT VI BASE with 20-23 hairs. Longest hairs on hind tibia 50-75 µm.
Dorsal abdominal hairs short, and at least those on ABD TERG 3-6 not
arising from sclerites (bases sometimes encircled by minute scleroites)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . newelli
6 Subgenital plate with only 2-6 hairs (more in fundatrices). First tarsal
segments with either 2 or 3 sense pegs (ventrally, near apex; see Fig. 75H)
............................................................. 7
- Subgenital plate with 25-50 hairs. First tarsal segments with only 1 sense
peg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 First tarsal segments with 2 sense pegs (as in Fig. 75H) . . . . . . . . westi
- First tarsal segments with 3 sense pegs . . . . . . . . . . . . . . . . . . . . . rigidae
8 ANT VI BASE with 5-8 hairs. R IV with 4-6 accessory hairs. Hind tibia
darker than fore or mid-tibiae. Longest hairs on hind tibia 40-50 µm,
about 0.5 x width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . taedae
Pinus 305
- ANT VI BASE with 8-16 hairs. R IV with 6-10 accessory hairs. Hind tibia
sometimes with a shorter pale section than fore and mid-tibiae, but not
darker overall. Longest hairs on hind tibia 50-100 µm, 0.75 or more x
width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 ABD TERG 5 with more than 20 hairs, the longest 60-180 µm, between
SIPH bases (Fig. 80E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atlantica
- ABD TERG 5 with less than 20 hairs, maximally 10-80 µm, between SIPH
bases (e.g. Fig. 80F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Longest hairs on ANT III 25-60 µm, and on dorsal side of hind tibia
50-70 µm. ANT III with 1-3 (non-protruberant) rhinaria near apex
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae
- Longest hairs on ANT III 50-70 µm, and on dorsal side of hind tibia
60-100 µm. ANT III with 0-1 rhinaria near apex (if present, the rhinarium
is small and protuberant) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . melaina
2 Hairs on hind tibia rather spine-like, those on dorsal side less than 55 µm
long, about 0.5 or less x width of tibia at midpoint . . . . . . . . . . apini
Hairs on hind tibia finer and more numerous, up to 65 µm long, clearly
more than 0.5 x width of tibia at midpoint . . . . . . . . . . . . . . . . . villosa
P. funebris = P. tabulaeformis
P. glabra Cinara arizonica, allantica,
(Spruce Pine) melaina, pergandei, taedae,
watsoni
Eulachnus rileyi
Schizolachnus curvispinosus,
parvus
(Use key to Cinara spp. on Pinus subsect. Australes under P. echinata)
P. gregii Cinara atlantica, maritimae,
ponderosae
(Use key to Cinara spp. on Pinus subsect. Oocarpae under P. oocarpa)
P. griffithii (=P. excelsa = Cinara eastopi, lachnirostris,
P. wallichiana) maculipes, maritimae
(Himalayan Blue Pine) Essigella californica
Eulachnus rileyi
Pineus ghanii, wallichianae
Prociphilus himalayensis
Pseudessigella brachychaeta
Key to Cinara spp. on Pinus oocarpae and other pines of subsect. Oocarpae
(attenuata, greggii, muricata, patula, radiata)
1 R IV 0.23-0.33 mm long, bearing 14-28 accessory hairs . . . . . . . . . . . 2
R IV 0.14-0.23 mm long, bearing 6-10 accessory hairs . . . . . . . . . . . . 3
2 Longest hairs on ANT III 60-80 µm, on hind tibia 50-80 µm . (ABD TERG
2-6 without sclerites?) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . essigi*
Longest hairs on ANT III and hind tibia 80-110 µm, all very fine. (ABD
TERG 2-6 often with numerous small sclerites) . . . . . . . . . . . . cronartii
3 HT I 0.16-0.22 mm long, more than 0.5 x HT II (Fig. 76A) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maritimae
312 Pinus
mediterraneus, rileyi,
tuberculostemmata
Pineus harukawai
(Use key to Cinara spp. on western palaearctic pines of subsect. Sylvestres,
under P. sylvestris)
P. pinea Cinara acutirostris, maghrebica,
(Italian Stone Pine) maritimae, schimitscheki
Essigella californica
Eulachnus rileyi,
tuberculostemmata
Pineus boerneri, havrylenkoi
Schizolachnus obscurus
(The three Cinara species are separated in the key under P. sylvestris)
P. ponderosa (incl. var. arizonica, Cinara arizonica, glabra,
var. scopularum) nigrita(?), oregonensis, pinea,
(Ponderosa Pine) pini, ponderosae,
pseudoschwartzii, schwartzii,
solitaria, thatcheri
Essigella californica, essigi, fusca
Eulachnus rileyi
Pineus boerneri, coloradensis
Schizolachnus curvispinosus,
flocculosus, piniradiatae
Key to Cinara spp. on Pinus ponderosa and other pines of subsect. Ponderosae
(cooperi, douglasiana, durangensis, jeffreyi, michoacana, pseudostrobus)
1 BL 4.6-6.Omm. Hairs on dorsal body surface, ANT III-IV and legs all
rather widely spaced, short, thick and slightly capitate . . . . . arizonica
- BL less than 4.6mm. Hairs sometimes short and thick but not widely
spaced and not capitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 HT I 0.15-0.23 mm long, more than 0.5 x HT II (Fig. 76B). Hairs on hind
tibia 120-230 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinea
- HT I 0.08-0.15 mm long, less than 0.5 x HT II. Hairs on hind tibia
20-120 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 R IV with 12-28 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- R IV with 6-10 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
4 Longest hairs on hind tibia 30-60 µm, about 0.5 x width of tibia at mid-
point or shorter. Hairs on ABD TERG 5 mostly short and blunt, less than
30 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Longest hairs on hind tibia 70-120 µm, clearly more than 0.5 x width of
tibia at midpoint. Hairs on ABD TERG 5 with acute apices, the longest
at least 70 µm in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
314 Pinus
5 Dorsum with a large dark sclerite fused across midline and extending
across ABD TERG 2-1, but not fused with SIPH bases (Fig. 77). ABD
TERG 5 with more than 50 hairs between SIPH bases. Ventral surface
covered with dense rows of pointed spicules . . . . . . . . . . . . . . . . . glabra
- Dorsum only lightly sclerotized, often with dusky areas on each side of
midline. ABD TERG 5 with less than 20 hairs between SIPH bases. Ven-
tral surface only faintly and minutely spiculose . . . . . . . . . . . . . . . . . . 6
6 R IV 0.20-0.25 mm long, with 15-16 accessory hairs. ANT V about equal
in length to ANT IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . schwartzii
- R IV 0.26-0.30 mm long, with 19-21 accessory hairs. ANT V clearly
longer than ANT IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . thalcheri
7 ABD TERG 2-6 without distinct sclerites but often with a dusky area on
each side of midline. ABD TERG 5 with less than 20 hairs between SIPH
bases. R IV 0.22-0.24 mm long . . . . . . . . . . . . . . . . . . . pseudoschwartzii
- ABD TERG 2-6 with scattered separate, distinct sclerites. ABD TERG 5
with more than 50 hairs between SIPH bases. R IV 0.24-0.29 mm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cronartii
8 SIPH cones with two distinctly different types of hair, one type long
(about 70 µm) and spine-like, the other small (about 12 µm), fine and
numerous. R IV 0.22-0.26 mm long. Body rounded, tick-like, tibiae short
and black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . solitaria
- SIPH cones with one type of hair. R IV 0.14-0.22. Body oval . . . . 9
Pinus 315
9 Longest hairs on hind tibia 80-110 µm, and on ABD TERG 5, 60-110µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Longest hairs on hind tibia 40-95 µm, and on ABD TERG 5, 10-70 µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 Length of sclerotized part of stylet groove (see Fig. 74A) 1.8-2.2 mm.
ABD TERG 5 with less than 20 hairs between the rather small SIPH cones,
each bearing 15-20 hairs. ABD TERG 7 and 8 with broad transverse bars
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . oregonensis
- Length of sclerotized part of stylet groove 1.3-1.7 mm. ABD TERG 5 with
more than 25 hairs (Fig. 80E), between large SIPH cones each bearing
more than 50 hairs. ABD TERG 7 and 8 with paired sclerites not joined
or only tenuously linked across midline . . . . . . . . . . . . . . . . . . . atlantica
11 Length of sclerotized part of stylet groove (see Fig. 74A) 1.5-2.0mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae/terminalis
- Length of sclerotized part of stylet groove 1.2-1.4 mm . . . . . . . . . pini
ABD TERG 2-4 each with 8-12 spinopleural hairs (rarely 7), and 3-5
(rarely 2) marginal hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Metathoracic tergite fused with ABD TERG 1, either entirely across dor-
sum (pale individuals) or at least laterally (dark individuals). All tibiae
dusky to dark, concolorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . essigi
Metathoracic tergite not fused to ABD TERG 1, even laterally. Fore and
hind tibiae clearly darker than middle tibia . . . . . . . . . . . . . . . . . . . . . . 3
3 BL 2.10-2.64 mm. ANT III 0.16-0.23 mm. Hind tibia 1.06-1.56 mm.
Dorsal body pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hillerislambersi
- BL 1.79-2.39mm. ANT III 0.12-0.18mm. Hind tibia 0.76-1.13mm.
Dorsal body pale or dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . fusca
Key to Cinara spp. on Pinus sylvestris and other western palaearctic pines of
subsect. Sylvestres (mugo, nigra, pinaster); also including species on P. pinea.
(Cinara spp. on P. halepensis, including var. brutia, are keyed separately
under that host plant.)
1 ABD TERG 5-7 usually with an extensive dark sclerite encompassing
SIPH cones (e.g. Fig. 78; the lateral extent of the sclerite varies con-
siderably, as does the occurrence of clear areas within it) . . . . . brauni
- ABD TERG 5-7 with or without sclerites but these are never fused
between segments or with SIPH cones . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Pinus 319
Fig. 79. Hair-bearing scleroites on ABD TERG 5 of aptera of A, Cinara schimitscheki and B, C. piniphila.
Fig. 80. Region of ABD TERG 5 between SIPH of aptera of A, Cinara eastopi, B, C. maritimae,
C, C. pinea, D, C. pergandei, E, C. atlantica, f, C. ponderosae, 6, C. pini, H, C. guadarramae.
Pineus boerneri
Prociphilus carolinensis,
piniradicivorus
Schizolachnus parvus
(Use key to Cinara spp. on Pinus subsect. Australes under P. echinata)
(To separate the two Essigella species use the key under P. strobus)
324 Pinus
PIPTOCARPHA Compositae
PISONIA Nyctaginaceae
PISTACIA Anacardiaceae
Pistacia spp. are the primary hosts of Fordina (Pemphiginae), which produce
a wide variety of galls, sometimes very large and elaborate (Fig. 126). No other
aphids (apart from the polyphagous Aphis gossypii) are recorded from this
genus. Following the host plant list, two keys are provided, both taking the
genus Pistacia as a whole: the first key is to the emigrant alatae produced in
the galls, which usually open in autumn, and the second is to the alate
sexuparae, which may be collected on the trunks when they return to Pistacia
in spring. Confirmation of identity will often be possible from the description
of the gall given in the systematic section of the book; many of the galls are
illustrated in Fig. 126.
Geoica utricularia
Paracletus cimiciformis
Slavum lentiscoides, mordvilkoi
Smynthrodes betae
Fig. 81. Antenna of spring migrant alata (from gall) of A, Aploneura lentisci, B, Slavum lentiscoides,
C, Rectinasus buxtoni, D, Asiphoniella cynodonti, E, Chaetogeoica foliodentata, f, Aloephagus myersi,
G, Baizongia pistaciae, H, Geoica utricularia, I, G. setulosa, J, Forda formicaria, K, F. marginata,
L, F. riccobonii, M, F. hirsuta, N, Paracletus cimiciformis, 0, Smynthurodes betae.
332 Pistacia
PITTOSPORUM Pittosporaceae
PLAGIANTHUS Malvaceae
PLANERA Ulmaceae
PLATYCARYA Juglandaceae
Kurisakia sinoplatycaryae
PODOCARPUS Podocarpaceae
- ANT III of aptera less than 2.6 x ANT IV, and ANT III of alata less
than 3.5 x ANT IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 ANT III of aptera less than 0.26 mm long, less than 0.8 of length of fore
tibia. ANT III of alata less than 0.4mm long, bearing 10-25 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
ANT III of aptera more than 0.26mm long, more than 0.8 of length
of fore tibia. ANT III of alata more than 0.4mm long, bearing 30-58
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 Aptera with many dorsal abdominal hairs arising from pigmented
scleroites of varying size and shape. Pigmented areas of head without a
reticulate pattern and localized, the dorsal and lateral areas being well
separated, with a clear area posteriolaterad to antennal base. Alata with
10-22 seconday rhinaria on ANT I I I , usually not extending to apex of
segment, and ANT IV without rhinaria . . . . . . . . . . . . . . . . . gingerensis
Aptera with dorsal abdominal hairs not based on pigmented scleroites.
Pigmented areas of head with a reticulate pattern, the dorsal and lateral
areas being contiguous, and the lateral pigmentation extending forward
from triommatidium to antennal base. Alata with 20-25 secondary
rhinaria on ANT I I I , distributed over whole length of segment, and
ANT IV often with a few rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . totarae
8 Aptera with ANT III 0.35-0.62 mm long; ANT I I I and ABD TERG
3-5 with longest hairs 9-18 µm long. Alata w i t h ANT III a little longer
than IV+V+VI together, bearing 39-58 secondary rhinaria, and knob
of CAUDA pale and without spinules on proximal half of dorsal
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . brimblecombei/lanata
- Aptera with ANT I I I 0.25-0.38 mm long; ANT III and ABD TERG
3-5 with longest hairs 20-30 µm long. Alata with ANT III usually a
little shorter than IV+V+VI together, bearing 30-44 rhinaria, and knob
of CAUDA entirely pigmented and spinulose . . . . . . . . . . . . podocarpi
POLYSCIAS Araliaceae
Pachypappa sacculi
Pemphigus betae, know1toni,
monophagus, populiglobuli,
populitransversus, populivenae
Pterocomma populifoliae,
pseudopopuleum, smithiae
Thecabius gravicornis,
populimonilis
P. balsamifera Aphis maculatae
(Tacamahac or Balsam Poplar) Chaitophorus leucomelas,
populialbae, populicola,
populifolii
Clydesmithia canadensis
Epipemphigus niisimae
Pachypappa sp.
Pemphigus betae, borealis,
monophagus, populicaulis,
populiglobuli, populiramulorum,
populitransversus, populivenae
Phylloxerina salicicola
Pterocomma bicolor, populeum,
populifoliae, [ rufipes]
Thecabius affinis, gravicornis,
populiconduplifolius,
populimonilis
P. x berolinensis Pemphigus bursarius, passeki,
phenax
Thecabius affinis
P. bolleana Chaitophorus populeti, populialbae
Neopemphigus turajevi
Pterocomma kozhuchovae
P. x canadensis - P. x euroamericana
P. candicans = P. balsamifera
P. x canescens Chaitophorus populeti, populialbae
(Grey Poplar) Pachypappa tremulae, vesicalis,
warshavensis
P. cathayana Epipemphigus sanpupopuli
Pemphigus tibetensis
Pterocomma neimongolense,
sanpunum
P. ciliata Chaitophorus kapuri
(Bangikat Poplar) Doraphis populi
Epipemphigus imaicus, marginalis
340 PopuIus
orientalis Mordvilko ex
Shaposhnikov, 1955]
Pemphigus borealis, dorocola,
matsumurai, microsetosus,
semenovi
Phloeomyzus passerinii
Pterocomma yezoense
Thecabius affinis, latisensorius,
orientalis
Tuberolachnus salignus
P. monilifera = P. deltoides var. occidentalis
P. nigra (incl. var. italica) Aphis maculatae
(Black Poplar, Lombardy Poplar) Aulacorthum solani
Chaitophorus euphraticus,
leucomelas, nassonowi,
neglectus, populeti, populialbae,
populicola, pruinosae
Pachypappa marsupialis
Pemphigus bursarius, gairi,
immunis, passeki, populi,
populinigrae, protospirae,
spyrothecae, vesicarius
Phloeomyzus passerinii
Phylloxerina populi
Pterocomma bicolor, populeum,
tremulae, yezoense
Stomaphis longirostris
Thecabius affinis
Thecabius (Parathecabius)
lysimachiae
Tuberolachnus salignus
P. occidentalis = P. deltoides var. occidentalis
P. pruinosa Chaitophorus pruinosae
Lambersaphis pruinosae
Pachypappa warshavensis
P. rasumowskyana Pemphigus bursarius, ?populinigrae
P. x robusta Chaitophorus leucomelas,
nassonowi
P. sargentii = P. deltoides var. occidentalis
P. sieboldii Chaitophorus dorocolus, populeti,
populialbae ssp, yomefuri,
tremulae
Doraphis populi
Pachypappa tremulae
Populus 343
3 Eyes with only 3 facets. ABD TERG 7 with a pair of large wax pore plates.
Antennae with spinules in a reticulate pattern . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phloeomyzus passerinii
- Eyes multifaceted. Abdomen without wax glands. Antennae not orna-
mented with a reticulate pattern of spinules . . . . . . . . . . . . . . . . . . . . . . 4
4 CAUDA tongue- or finger-like, tapering, much longer than its basal width
............................................................. 5
- CAUDA knobbed, rounded or triangular . . . . . . . . . . . . . . . . . . . . . . . . 7
5 Antennal tubercles undeveloped. SIPH and CAUDA black. ANT III with
15-22 secondary rhinaria over distal 0.8 of length. Tibiae with numerous
long, fine hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis maculatae
- Antennal tubercles well developed. SIPH and CAUDA mainly pale. ANT
III with 1-9 secondary rhinaria on basal 0.4. Tibiae with short hairs
............................................................. 6
6 Antennal tubercles w i t h inner faces approximately parallel (Fig. 42J).
ANT III with a single rhinarium near base. SIPH without an apical zone
of polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
- Antennal tubercles with inner faces clearly divergent (Fig. 42K). ANT III
with 2-9 rhinaria. SIPH with a subapical zone of polygonal reticulation
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
7 Very large aphid with ANT PT/BASE 0.6 or less. SIPH pores placed on
broad dark hairy cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Small or large aphid with ANT PT/BASE more than 0.6. SIPH truncate
conical (with reticulate sculpturing) or tubular, or merely pores, or
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
8 Rostrum much longer than body, which is elongate oval. ANT PT/BASE
about 0.1, with PT broad and rounded . . . . . . . Stomaphis longirostris
- Rostrum much shorter than body, which is broadly oval. ANT PT/BASE
0.3-0.5, with PT tapering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Dorsal abdomen with a large dark spinal tubercle. BL usually less than
5.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberolachnus salignus
- Dorsal abdomen without a tubercle. BL usually more than 5.5 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 HT II less than 2.2 x HT I. ANT III with longest hairs much longer than
diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- HT II more than 2.2 x HT I. ANT III with longest hairs shorter than
diameter of segment . . . . . . . . . . . . . . . . . . . . . Longistigma xizangensis*
11 SIPH in form of small truncate cones or short cylinders (usually at least
0.5 x basal width), with reticulate sculpturing at least distally. CAUDA
knobbed or rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
346 Populus
Fig. 82. A, hind tibia of apterous vivipara of Chaitophorus populeti with scent plaques; CAUDA of aptera
of B, Ch. nassonowi, C, Ch. populicola and D, Ch. populialbae; HT I of E, Ch. populialbae and F, Ch. tremulae.
16 ABD TERG 1-7 usually all free, with separate spinopleural and marginal
sclerites. Tibiae with pale middle sections . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus miyazakii *
- ABD TERG 2-7 wholly sclerotized, fused into a solid tergum. Tibiae
wholly dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 ABD TERG 2-7 each with only 3-5 spinal hairs (plus marginals), the
pleural regions being devoid of hairs . . . . . . . . . . . Chaitophorus nudus
- ABD TERG 2-7 each with pleural as well as spinal and marginal hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 Abdominal tergum evenly spinulose or nodulose . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus nodulosus
- Dorsal cuticle without evident spinules or nodules . . . . . . . . . . . . . . . 19
19 Many of dorsal abdominal hairs blunt or w i t h square or furcate apices
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus populicola
- All dorsal abdominal hairs long, fine and acute . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus populicola ssp. patchae
20 First tarsal segments usually with 5 hairs (e.g. Fig. 82E; rarely 6 or more
on some tarsi) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- First tarsal segments usually with 7 hairs (e.g. Fig. 82F; rarely 6 on some
tarsi) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
21 ABD TERG 1-6 fused, wholly dark, distinctly ornamented with minute
spinules, and with numerous smaller pointed hairs between the very long
spinal, pleural and marginal hairs on each segment, which have usually
pointed but sometimes blunt or bifurcate apices (Fig. 83A) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus longisetosus
- Dorsal abdomen pale, or with separate dark segmental bands; more-or-
less smooth or ornamented with blunt nodules or spinules, or polygonal
reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 ANT PT/BASE 1.5 or less (fundatrix has ANT PT/BASE 0.6-0.9) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus pruinosae*
- ANT PT/BASE 1.7 or more (1.3 or more in fundatrix) . . . . . . . . . . 23
23 R 1V+V shorter than HT II, with 2 accessory hairs . . . . . . . . . . . . . . 24
- R IV+V as long as or a little longer than HT II, with 2-4 accessory
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
24 Spinopleural hairs on ABD TERG 2-7 very short, maximally about
20 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus clarus
- Spinopleural hairs long, maximally 100-150 µm . . . . . . . . . . . . . . . . . 25
25 ABD TERG 2-6 with weakly spinulose ornamentation, and long thick hairs
348 Populus
Fig. 83. Hairs in spinal region of ABD TERG 3 of aptera of A, Chaitophorus longisetosus, B, Ch. inouyei,
C, Ch. variegatus, D, Ch. hillerislambersi, E, Ch. tremulae, F, Ch. euphraticus, 6, Ch. stevensis, H, Ch. neglectus.
maximally 150 µm. ANT III with 3-9 hairs, the longest 1.7-2.2 x basal
diameter of segment . . . . . . . . Chaitophorus populialbae ssp. yomefuri
- ABD TERG 2-6 with weakly to strongly nodulose ornamentation, and
hairs up to 120 µm long (Fig. 83F). ANT III with 6-13 hairs, the longest
0.8-1.7 x basal diameter of segment . . . . . . Chaitophorus euphraticus
26 ANT PT/BASE 3.3-4.2. SIPH dark .... Chaitophorus melanosiphon
- ANT PT/BASE 1.7-3.2. SIPH pale . . . . . . . Chaitophorus populialbae
27 Empodial hairs enlarged and flattened, with expanded apices. Abdominal
tergum pale, with long, pointed or spine-like spinal, pleural and marginal
hairs interspersed with smaller hairs, some of which are much smaller
(0.25 x length of longest) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
- Empodial hairs setaceous, if somewhat flattened then the abdominal
tergum is dark and/or the spinopleural hairs are often blunt or furcate
and/or their variation in size is much less . . . . . . . . . . . . . . . . . . . . . . 30
28 ABD TERG 2-6 with smallest (accessory) hairs mainly thick and spine-
like, and very numerous (30-40 per segment; Fig. 83B) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus inouye
Populus 349
- ABD TERG 2-6 with accessory hairs finer and less numerous (less than
25 per segment; e.g. Fig. 83C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 Longest hairs on ANT III 3.5-4.9 x basal diameter of segment. ANT PT/
BASE 3.5-4.5. ABD TERG 1-7 mostly smooth, without ornamentation
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus variegatus
- Longest hairs on ANT III 1.7-2.5 x basal diameter of segment. ANT
PT/BASE 2.7-3.4. ABD TERG 1-7 with weak squamous or nodulose
ornamentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus indicus*
30 ABD TERG 1 frequently fused with ABD TERG 2-6 (examine several
specimens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
- ABD TERG 1 almost always free from ABD TERG 2-6 . . . . . . . . . 37
31 R IV+V 0.17-0.20mm, 1.2-1.4 x HT I I . Abdominal tergum smooth,
bearing mostly long hairs w i t h bifurcate apices (Fig. 83D) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus hillerislambersi
- R IV+V 0.07-0.14 mm, 0.7-1.2 x HT II. Abdominal tergum usually
ornamented with spicules or nodules . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 Dorsal ornamentation consisting of close, irregular rows of posteriorly-
directed, bluntish spinules, often united basally to form partial transverse
rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3
- Dorsal ornamentation consisting of rather evenly distributed separate
roundish nodules (e.g. Fig. 83E; sometimes weakly developed) . . . . 35
33 Dorsum pale. ANT PT/BASE 1.5-2.4 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus tremulae ssp. sorini
- Dorsum usually deeply pigmented, sometimes dark only on head and ABD
TERG 8. ANT PT/BASE 2.2-3.2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34 Longest hairs on ANT III 1.7-3.2 x basal diameter of segment . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus tremulae s. str.
- Longest hairs on ANT III with wavy, attenuate apices, 3.0-4.5 x basal
diameter of segment . . . . . . Chaitophorus tremulae ssp. shantungensis
35 Hairs on each of ABD TERG 1-7 very variable in size and form, arranged
more or less in a single row (Fig. 83G; sometimes with 1-2 smaller hairs
out of line). ABD TERG 8 with 7-12 hairs. (Alata with SIPH dark)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus stevensis
- Hairs on each of ABD TERG 1-7 arranged in two irregular rows, the
anterior row consisting of smaller hairs which are sometimes absent in
spinal region (e.g. Fig. 83H). ABD TERG 8 usually with more than 12
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 Head and ANT I-II often dark, and tergum often with a pair of dark
longitudinal pleural stripes. Most of long spinal and pleural hairs on ABD
350 Populus
Fig. 84. Right SIPH of aptera of A, Plerocomma smithiae, B, Pt. bicolor and C, Pt. beulahense.
Fig. 85. A, ANT V and VI of emigrant alata of Gootiella tremulae; B, SIPH and surrounding wax gland
of alata of Pachypappella lactea; C, SIPH of alata of Pachypappa populi; D, hairs on forewing membrane
of Pachypappa vesicalis; ANT III of emigrant alata of E, P. warshavensis and F, P. pseudobyrsa;
G, front of head of alata of Cornaphis populi; ANT VI of emigrant alata of H, Mordvilkoja vagabunda,
I, Epipemphigus niisimae, J, E. sanpopuli, K, E. imaicus.
6 CAUDA with more than 40 hairs. SIPH pores present. Secondary rhinaria
on ANT III 8-13, on IV 2-4 . . . . . . . . . . . . . . . . . . Pachypappa populi
- CAUDA with less than 30 hairs. SIPH pores absent. Secondary rhinaria
on ANT III 5-10, on IV 0-3 . . . . . . . . . . . . . . . . . . Pachypappa sacculi
7 R IV+V 0.60-0.75 x HT II. Media sometimes unbranched . . . . . . . . 8
- R IV+V 0.45-0.55 x HT II. Media always once-branched ......... 9
PopuIus 355
Fig. 86. ANT III of emigrant alata of A, Parathecabius latisensorius and B, Thecabius affinis; antenna of
emigrant alata of C, Pemphigus matsumurai and D, P. populi.
21 R IV with usually 1-6 small accessory hairs (rarely with none) and first
tarsal segments with 2-6 hairs (usually at least 3 on most tarsi). Secondary
rhinaria very narrow, slit-like (Fig. 86B), with 2-9 on ANT BASE VI
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Thecabius affinis/populiconduplifoliae
- R IV usually without accessory hairs, sometimes with 1-4 (but one of the
3 apical pairs is sometimes displaced to a more proximal position). First
tarsal segments usually with 3 hairs on fore tarsus and 2 hairs on mid-
and hind tarsi. If more, then ANT BASE VI is without secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 SIPH pores absent and embryos with mouthparts (the curled stylets are
especially evident through wall of abdomen) . . . . . . . . . . . . . . . . . . . . 23
- Either SIPH present as small pores or embryos without mouthparts
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
23 R IV+V broadly triangular, less than 2 x its basal width. (Secondary
rhinaria on ANT I I I 5-19, IV 3-7, V 3-4) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . Thecabius populimonilis (fundatrigeniae)
- R IV+V acute, more than 2 x its basal width . . . . . . . . . . . . . . . . . . . 24
24 Secondary rhinaria present on ANT VI. (On III 10-18, IV 4-7, V 3-7,
BASE VI 3-7) . . . . . . . . . . . . . . . . . . . . Pemphigus napaeus/iskanderkuli
- Secondary rhinaria absent from ANT VI . . . . . . . . . . . . . . . . . . . . . . . 25
25 Secondary rhinaria strap-like, parallel-sided (Fig. 86C), many of them
extending more than half-way around antenna, with 2-5 on ANT V
(III 8-13, IV 2-5, V 2-5). ABD TERG 8 with one large wax pore plate
extending more than half-way across tergite, and including 4 hairs . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus matsumurai
- Secondary rhinaria transversely elongate oval, not parallel-sided, extend-
ing up to half-way around antenna but not more, with 0-2 on ANT V
(e.g. Fig. 86D). ABD TERG 8 with a pair of small wax pore plates or a
single median wax pore plate including only 2 hairs . . . . . . . . . . . . . 26
26 First tarsal segments almost always with 2 hairs, rarely with 3. HT II
0.17-0.21 mm long. Secondary rhinaria on ANT III 3-9, IV 1-3,
V 0-2 (Fig. 86D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populi
- First tarsal segments usually with 3-4 hairs on at least some tarsi. HT II
0.20-0.25 mm long. Secondary rhinaria on ANT III 6-11, IV 1-4, V 0-2
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus vesicarius
27 Embryos without mouthparts. Secondary rhinaria always absent from
ANT V and VI. SIPH pores absent. (Sexuparae of monoecious species,
produced in the gall) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
- Embryos with mouthparts. Secondary rhinaria usually present on ANT
V and VI. SIPH present as small pores . . . . . . . . . . . . . . . . . . . . . . . . 31
358 Populus
28 First tarsal segments usually all with 3-5 hairs, rarely with only 2. Spinal
wax pore plates often present on all abdominal tergites . . . . . . . . . . 29
- First tarsal segments most commonly with 3 (-4) hairs on fore tarsus and
2 (-3) hairs on mid- and hind tarsi. Spinal wax pore plates usually absent
from ABD TERG 4-7. (Secondary rhinaria distributed III 3-6, IV 1-3)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
29 R IV+V about 0.08 mm long, slightly shorter than ANT V. Secondary
rhinaria on ANT III 4-7, on IV 1-4 . . . . . . . Thecabius populimonilis
- R IV+V 0.09-0.11 mm long, as long as or slightly longer than ANT V.
Secondary rhinaria on ANT III 6-10, on IV 2-5 . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus mongolicus
30 R IV+V usually without accessory hairs. CAUDA with 4-6 hairs . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus monophagus
- R IV+V usually with 1-4 accessory hairs, rarely with none. CAUDA with
2-4 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus spyrothecae
31 ANT BASE VI usually without separate secondary rhinaria (rarely with
1, in which case there are usually none on V) . . . . . . . . . . . . . . . . . . . 32
- ANT BASE VI usually with 2 or more secondary rhinaria separate from
the primary rhinarium (rarely with 1 only, in which case there is usually
1 or more on V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
32 ANT VI with an enlarged primary rhinarium (due to coalescence with
secondary rhinaria), with islands, like that on V . . . . . . . . . . . . . . . . 33
- ANT VI with primary rhinarium of normal size, no longer than wide
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
33 ANT VI with primary rhinarium 0.07-0.10 mm long, occupying more
than half of length of BASE VI (Fig. 87E), or with a separate secondary
rhinarium immediately adjacent to it. Secondary rhinaria on III 3-9, IV
1-3, V 0(-1) and BASE VI 0(-1) . . . . . . . . . . . . . Pemphigus knowltoni
- ANT VI with primary rhinarium less than 0.07 mm long, occupying less
than half of length of BASE VI (Fig. 87F). Secondary rhinaria on III 2-7,
IV 0-3, V 0(-1), BASE VI 0 . . . . . . . . . . Pemphigus populitransversus
34 ANT V with primary rhinarium very large, more than 0.04mm long
and more than half encircling segment, often with small round islands
(Fig. 87G). (Secondary rhinaria on III 7-9, IV 2-4, V 0-1, BASE VI 0)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus immunis
- ANT V with primary rhinarium either like that on VI or, if enlarged,
broader than its axial length which is less than 0.03 mm . . . . . . . . . 35
35 First tarsal segments with two spine-like hairs between the 2 long apical
hairs. ABD TERG 8 usually without a wax pore plate. ANT PT more than
2 x its basal width. (Secondary rhinaria on III 10-14, IV 2-3, V 0(-1),
BASE VI 0) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus microsetosus
Populus 359
- First tarsal segments with 0-1 spine-like hairs between the two long apical
hairs. ABD TERG 8 usually with wax pore plates. ANT PT less than
2 x its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 R IV+V about 0.1 mm, more than 0.55 x HT II. Rhinaria on ANT III
9-13, IV 3-6, V 0-1, BASE VI 0 . . . . . . . . . . . . Pemphigus circellatus*
- R IV+V 0.07-0.085 mm, less than 0.55 x HT II. Rhinaria on ANT III
5-11, IV 1-4, V 0-3, BASE VI 0(-1) . . . . . . . . . . . . . Pemphigus betae
37 Total antennal length 0.9mm or more, about 0.5 x BL . . . . . . . . . . 38
- Total antennal length less than 0.9 mm, less than 0.45 x BL . . . . . 39
38 Secondary rhinaria on ANT I I I 19-27, on IV 6-7, V 9-12, VI BASE 10-12
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus longicornus
- Secondary rhinaria on ANT I I I 15, IV 4, V 4, VI 5 (one specimen only)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus nainitalensis*
39 R IV+V 0.10-0.12 mm long, w i t h R IV straight-sided (Fig. 87A), usually
more than 0.55 X HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
- R IV+V 0.07-0.10 mm long, R I V with sides slightly convex (e.g.
Fig. 87B), usually less than 0.55 x HT I I . . . . . . . . . . . . . . . . . . . . . . . 41
40 Secondary rhinaria narrow, s l i t - l i k e , averaging 8 µm or less in width,
extending almost to base of ANT I I I , with most proximal rhinarium
usually basad to the small tooth on inner side of III near base (Fig. 87C);
on I I I 12-18, IV 3-7, V 4-7, VI BASE 4-7 . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populinigrae
- Secondary rhinaria broader, averaging more than 10µm in width, with
most proximal rhinarium on ANT III distal to tooth (Fig. 87D); on
III 10-14, IV 2-5, V 2-4, VI BASE 2-8 . . . . . Pemphigus protospirae
41 Primary rhinarium on ANT V usually much enlarged, its axial length
35 µm or more, with small islands (Fig. 87H). Secondary rhinaria on
ANT I I I 6-12, IV 2-3, V 0-3, VI BASE 2-5 . . . . . Pemphigus phenax
- If primary rhinarium of ANT V is enlarged then its axial length is less than
35 µm and the antennal sensoriation is different . . . . . . . . . . . . . . . . . 42
42 Total number of secondary rhinaria on ANT V+VI BASE (both antennae)
24 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
- Total number of secondary rhinaria on ANT V+VI BASE (both antennae)
23 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
43 ANT VI BASE slightly shorter than HT I I . (Secondary rhinaria on ANT
III 11-19, IV 4-8, V 4-10, VI BASE 6-11) . . . . . Pemphigus trehernei
- ANT VI BASE slightly longer than HT II . . . . . . . . . . . . . . . . . . . . . . 44
44 ABD TERG 8 with 2-3 hairs. (Secondary rhinaria on I I I 13-19, IV 4-5,
V 6-9, VI BASE 6-10) . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus borealis
- ABD TERG 8 usually with 3-5 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 45
360 Populus
Fig. 87. R IV+V of alata of A, Pemphigus populinigrae and B, P. bursarius; ANT III of emigrant alata of
C, P. populinigrae and D, P. protospirae, with small tooth on inner side arrowed; ANT VI of
E, P. knowltoni and F, P. populitransversus; ANT V and VI of G, P. immunis, H, P. phenax and
I, P. passeki; basal part of ANT III of J, P. populicaulis and K, P. laurifoliae.
48 ANT III 0.22-0.38 mm, but if less than 0.23 mm (small bursarius) then
ANT PT is short and stubby, less than 1.5 x its basal width . . . . . 49
- ANT III 0.15-0.25 mm, but if more than 0.22 mm then ANT PT is more
than 1.5 x its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
49 Primary rhinarium on ANT VI much larger than adjacent secondary
rhinaria and forming a broad band that extends over half-way around
antenna (Fig.87I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
- Primary rhinarium on ANT VI normal or irregular in shape but not
extending more than half-way around antenna . . . . . . . . . . . . . . . . . . 52
50 ANT VI BASE with 6-8 secondary rhinaria (Fig. 87I). (III 12-16, IV 3-5,
V 4-6, VI BASE 6-8) . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus passeki
- ANT VI BASE with 3-6 secondary rhinaria . . . . . . . . . . . . . . . . . . . . 51
51 ANT PT 0.03-0.04 mm. (Secondary rhinaria on I I I 10-15, I V 4-5, V 2-4,
VI BASE 3-6) . . . . . . . . . . . . . . . . . . . . . . . Pemphigus junctisensoriatus*
- ANT PT 0.02-0.03 mm. (Secondary rhinaria on I I I 12-16, I V 4-6, V 3-5,
VI BASE 3-6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus dorocola*
52 Tooth on inner side of ANT I I I often very close to base of segment,
usually without a rhinarium basad to it (Fig. 87J). (Secondary rhinaria on
III 8-13, IV 2-5, V 2-5, VI BASE 3-7) . . . . . Pemphigus populicaulis
- Tooth on inner side of ANT III almost always with 1-2 secondary rhinaria
basad to it . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
53 Secondary rhinaria on ANT I I I 8-17, IV 2-8, V 2-6, VI BASE 3-8. PT
usually less than 1.5 x basal width . . . . . . . . . . . . Pemphigus bursarius
- Secondary rhinaria on ANT III 9-13, IV 3-5, V 3-5, VI BASE 3-5. PT
usually more than 1.5 x basal width . . . . . . . . . Pemphigus mordvilkoi
54 Tooth on inner side of ANT III usually more than 0.07 mm from base
of segment, with always 1, and sometimes 2-3, secondary rhinaria basad
to it (Fig. 87K). (Secondary rhinaria on III 9-12, IV 2-4, V 2-4, VI BASE
3-5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus laurifoliae
- Tooth on inner side of ANT III usually less than 0.06 mm from base of
segment, with 0-1 secondary rhinaria basad to it . . . . . . . . . . . . . . . . 55
55 BL 2.1-2.5 mm. ANT V 0.8-1.0 x ANT IV, and more often than not
also bearing fewer secondary rhinaria than IV (examine several specimens;
secondary rhinaria distributed III 9-11, IV 0-5, V 0-4, VI 0-5) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populiramulorum
- BL 1.1-2.3 mm (only P. populiglobuli more than 2.0mm). ANT V
1.0-1.2 x ANT IV, and usually with at least as many sensoria as IV
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
56 ANT PT short, maximally about 0.03 mm long and usually less than
3 . 5 x its basal width. Tooth on ANT III usually without a rhinarium
362 Pouteria
PREMNA Verbenaceae
PROSOPIS Leguminosae
PRUNUS Rosaceae
Tinocalloides montanus
Tuberocephalus higansakurae,
sakurae
P. cornuta Avicennina indica
Eumyzus pruni
Myzus cornutus
Pyrolachnus imbricatus
[Rhopalosiphum maidis]
Vesiculaphis pruni
P. davidiana Myzus persicae
P. divaricata Brachycaudus cardui, divaricatae,
[divaricatellus], helichrysi,
persicae, prunicola, schwartzi
Hyalopterus pruni
Phorodon humuli
Pterochloroides persicae
Rhopalosiphum nymphaeae
P. domestica Aphis longicauda, spiraecola
(Common or European Plum) Brachycaudus [almatinus], cardui,
divaricatae, helichrysi, persicae,
prunicola, semisubterraneus
Hyalopterus pruni
Hysteroneura setariae
Macrosiphum euphorbiae
Myzus ornatus, persicae
Nearctaphis baker i
Phorodon humuli
Pterochloroides persicae
Rhopalosiphum [musae],
nymphaeae, padi
(For a key to common aphids on Plum see B & E, 1984)
P. donarium = P. lannesiana
P. donarium var. sachalinensis = P. sachalinensis
P. donarium var, speciosa = P. jamasakura var. speciosa
P. donarium var. spontanea - P. serrulata var. spontanea
P. dulcis Aphis fabae, gossypii
(Almond) Brachycaudus amygdalinus,
helichrysi, persicae
[Capitophorus prunifoliae Shinji,
1924]
Hyalopterus amygdali
Myzus amygdalinus, persicae
Prunus 365
Pterochloroides persicae
Rhopalosiphum nymphaeae
(For a key to common aphids on Almond see B & E, 1984)
P. emarginata Myzus cerasi, lythri
(Bitter Cherry)
P. erythrocarpa Aphidura bozhkoae
Brachycaudus pilosus
P. glandulosa Hyalopterus pruni
Myzus persicae, varians
Phorodon japonensis
Rhopalosiphum nymphaeae,
rufiabdominalis
Tuberocephalus momonis
P. grayana Myzus persicae
Tuberocephalus sp. (Sorin, 1993)
P. hortulana Hyalopterus pruni
P. humilis Tuberocephalus jinxiensis
P. incana Aphidura bozhkoae
P. insititia Brachycaudus cardui, helichrysi,
(Bullace, Damson) persicae, semisubterraneus
Brachycaudus (Appelia) prunicola
Hyalopterus pruni
Phorodon humuli
Rhopalosiphum nymphaeae
(For a key to common aphids on Damson see B & E, 1984)
P. itosakura var. ascendens Tuberocephalus sasakii
P. jamasakura (incl. var. speciosa) Myzus cerasi, mushaensis, persicae
(Yamasakura) Myzus (or Tuberocephalus) spp.
(Moritsu and Tokumotu, 1972)
Rhopalosiphum rufiabdominalis
Tuberocephalus sakurae, sasakii
P. japonica Phorodon japonensis
P. lannesiana Myzus mushaensis
(Oshima-zakura) Myzus (or Tuberocephalus) spp.
(Moritsu and Tokumotu, 1972)
Tuberocephalus sakurae, sasakii
P. laurocerasus Aulacorthum solani
(Cherry Laurel) Macrosiphum euphorbiae
Myzus persicae
P. lyonii Aphis spiraecola
366 Prunus
1 CAUDA knobbed, anal plate bilobed. SIPH short dark truncate cones,
shorter than their basal width, with a single hair on apical half. (All
viviparous females alate) . . . . . . . . . . . . . . . . . . . Tinocalloides montanus
- CAUDA rounded, triangular or finger-like, sometimes with a constriction
but not distinctly knobbed. Anal plate entire. SIPH of various forms, with
or without hairs, or absent. (Viviparae apterous or alate, rest of key
applies to apterae except where otherwise stated) . . . . . . . . . . . . . . . . . 2
2 SIPH as large pores on broad dark hairy cones . . . . . . . . . . . . . . . . . . 3
- SIPH tubular or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3 Dorsum with a double row of large pigmented spinal tubercles. (Forewings
of alata maculate with a short blunt pterostigma) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterochloroides persicae
- Dorsum without spinal tubercles. (Forewings of alata not maculate,
pterostigma elongate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 EL less than 5.5 mm. (Alata with straight pterostigma, not reaching tip of
wing) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrolachnus imbricatus
- BL more than 5.5 mm. (Alata with pterostigma curved around tip of wing)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma xizangensis*
5 ANT PT/BASE less than 0.3. SIPH absent, CAUDA broadly rounded.
(Forewing of aiata with an unbranched media) . . . . Prociphilus oriens
- ANT PT/BASE more than 0.8. SIPH present or absent, if absent then
CAUDA not broadly rounded. (Forewing of alata once- or twice-branched)
............................................................. 6
6 SIPH absent. Large lateral tubercles on thorax and ABD TERG 1-7
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asiphonaphis pruni
- SIPH present. Lateral tubercles present or absent . . . . . . . . . . . . . . . . 7
7 Head capsule smooth, wrinkled or rugose, but not ornamented with
spicules or nodules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Head capsule with spiculose or nodulose ornamentation . . . . . . . . . . 30
8 SIPH small and thin, less than 0.7 x CAUDA, lacking an apical flange
(Figs 88A, B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- SIPH more than 0.7 x CAUDA, usually with an apical flange . . . . 10
9 SIPH very thin, 2.5-4.0 x their maximum diameter (Fig. 88A). (Alata
with total secondary rhinaria on both ANT III 41-74, on ANT IV 9-21)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalopterus pruni
- SIPH 1.6-2.5 x their maximum diameter (Fig. 88B). (Alata with total
secondary rhinaria on both ANT III 21-41, on ANT IV 0-10) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalopterus amygdali
370 Prunus
Fig. 88. SIPH of aptera (in spring populations) of A, Hyalopterus pruni, B, H. amygdali, C, Avicennina
indica, D, Rhopalosiphum nymphaeae, E, Rh. cerasifoliae, F, Rh. padi.
10 SIPH dark, smooth and conspicuously inflated over more than half
length, with maximum diameter of swollen part 2 or more x minimum
diameter near base (Fig. 88C) . . . . . . . . . . . . . . . . . . . . Avicennina indica
- SIPH tapering, cylindrical, or if swollen then swelling is confined to distal
half, of maximum diameter less than 2 x minimum diameter of basal
half, and surface is rugose or imbricated . . . . . . . . . . . . . . . . . . . . . . . 11
11 Body spindle-shaped, SIPH long, 1.5-2.1 x the long finger-like CAUDA,
and with subapical zone of polygonal reticulation (Fig. 121C). ANT III
with 2-7 secondary rhinaria near base . . . . . Macrosiphum euphorbiae
- Body oval, if CAUDA is finger-like then SIPH are usually relatively
shorter, and always without polygonal reticulation. ANT III without
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 CAUDA tongue- or finger-like or tapering, usually more than 1.2 x its
basal width, but if shorter then with pointed apex . . . . . . . . . . . . . . . 13
- CAUDA bluntly conical, semicircular or broadly rounded, rarely up to
1.2 x its basal width, and never pointed . . . . . . . . . . . . . . . . . . . . . . . . 20
13 CAUDA long (more than 0.2 mm), conspicuously pale compared with the
black siphunculi, and bearing 4 hairs. Femora with contrasting black
distal and pale basal halves . . . . . . . . . . . . . . . . . . Hysteroneura setariae
Prunus 371
- CAUDA usually dusky or dark, if pale then shorter than 0.2 mm and with
more than 4 hairs. Femora without contrasting black and pale sections
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- ANT III always without secondary rhinaria. If antennal tubercles are well
developed then the inner faces are usually gibbous in dorsal view or bear
projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
34 Head, legs and antennae mainly dark; femora basally pale but with distal
part black. SIPH clavate (Fig. 121E). CAUDA with a constriction . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum magnoliae
- Head, legs and antennae mainly pale. SIPH gradually tapering (Fig. 121K).
CAUDA without a constriction . . . . . . . . . . . . . . . . Aulacorthum solani
35 ANT I with a scabrous protruberance on inner side near apex. Antennal
tubercles (except in fundatrix) with a forwardly-pointed projection as long
as or longer than its basal width. SIPH long, pale and tapering . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
- ANT I with inner side scabrous or smooth but not protruberant. Antennal
tubercles without a finger-like projection. SIPH variable . . . . . . . . . 37
36 R IV+V more than 1.3 x HT II . . . . . . . . . . . . . . Phorodon japonensis
- R IV+V less than 1.3 x HT I I . . . . . . . . . . . . . . . . . . . Phorodon humuli
37 Antenna of aptera (= fundatrix) 4-segmented, w i t h ANT PT/BASE less
than 1. (Apterous fundatrigeniae not produced) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberocephalus sasakii
- Antenna of aptera (including f u n d a t r i x ) 5- or 6-segmented, with ANT
PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38 SIPH with one or more hairs (not always in fundatrix) . . . . . . . . . . 39
- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
39 Dorsum pigmented (except fundatrix) . . . . . . . . . . . . . . . . . . . . . . . . . . 40
- Dorsum of cleared specimens pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
40 Femora spinulose . . . . . . . . . . . . . . . . . . . . . . . . . Tuberocephalus sakurae
- Femora without spinules . . . . . . . . . . . . . . Tuberocephalus liaoningensis
41 Tibiae scabrous along entire length . . . . . . . . Tuberocephalus momonis
- Tibiae with at least basal 0.75 of length smooth . . . . . . . . . . . . . . . . 42
42 SIPH about 2 x longer than basal width, shorter than ANT PT . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberocephalus jinxiensis*
- SIPH more than 3 x basal width, a little longer than ANT PT . . . 43
43 SIPH with 1-3 hairs in aptera (but not fundatrix), and 0-2 hairs in spring
migrant alata, which has secondary rhinaria distributed ANT III 44-60,
IV 14-22, V 1-5 . . . . . . . . . . . . . . . . . . . . . . . Tuberocephalus misakurae
- SIPH with 5-12 hairs in aptera (but not fundatrix), and 5-12 hairs in
spring migrant alata, which has secondary rhinaria distributed ANT III
15-25, IV 5-7, V 0-2 . . . . . . . . . . . . . . . . Tuberocephalus higansakurae
Prunus 375
44 SIPH jet black distally with basal half contrastingly pale, the transition
between the two tones being rather abrupt. Antennae also contrastingly
banded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus varians
- SIPH and antennal segments pale or dark, sometimes pale basally and
becoming darker distally, but not contrastingly two-toned . . . . . . . . 45
45 Longest hairs on ABD TERG 1-6 more than 20 µm long, with pointed
apices, 0.8 or more x basal diameter of ANT III. Hairs on ANT III with
blunt or pointed apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
- Longest hairs on ABD TERG 1-6 less than 20 µm long, with blunt or
expanded apices, usually less than 0.8 x basal diameter of ANT III. Hairs
on ANT III always with blunt apices . . . . . . . . . . . . . . . . . . . . . . . . . . 51
46 Hairs on ABD TERG 1-6 t h i c k , rather spine-like, with distinctly tuber-
culate bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
- Hairs on ABD TERG 1-6 fine, w i t h o u t or with only very small tuberculate
bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
47 Dorsal cuticle strongly corrugated. R IV+V 0.8-0.9 x HT II. ANT
PT/BASE 2.20-2.42 (except in fundatrices) . . . . . . . . . Eumyzus pruni
- Dorsal cuticle not strongly corrugated. R IV+V 1.0-1.1 x HT I I . ANT
PT/BASE 2.60-3.22 (except in fundatrices) . . . Eumyzus prunicolus
48 SIPH black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus padellus
- SIPH pale or dusky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
49 SIPH less than 2.3 x CAUDA, which is more than 1.25 x its basal width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae (part)
- SIPH more than 2.3 x CAUDA, which is less than 1.25 x its basal width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
50 R IV+V 0.85-0.95 x HT II. Tibiae smooth. (Hind tibia of immatures
weakly spinulose) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus cornutus
- R IV+V 1.1-1.2 x HT II. Tibiae with spinules or imbrication at base.
(Hind tibiae of immatures smooth) . . . . . . . . . . . . . . . Myzus mumecola
51 SIPH dark, rather thin, narrow-based (length only about 6 x basal
diameter which is less than 0.06mm), with a slight subapical swelling
(Fig. 89C). Dorsal cuticle strongly corrugated . . . . . . Myzus yamatonis
- SIPH pale or dark, broad-based (length less than 6 x basal diameter,
and/or basal diameter more than 0.06mm). Dorsal cuticle variable,
sculptured or smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
52 Dorsum fully covered with a solid blackish sclerotic shield with warty
ornamentation. SIPH jet black, long and tapering (Fig. 89D) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus cerasi/umefoliae
376 Prunus
Fig. 89. A, CAUDA of aptera of Roepkea marchali; B, mesosternal processes of Aphidura bozhkoae;
SIPH of aptera of C, Myzus yamatonis, D, M. cerasi and E, M. mushaensis.
PSEUDOPANAX Araliaceae
PSEUDOTSUGA Pinaceae
Antenna with hairs shorter than basal diameter of ANT III. R IV+V
usually without any accessory hairs. Head without wax pore plates.
CAUDA not protruberant, with 2 short hairs . . . Pachypappa rosettei
4 Body elongate. Antenna 5-segmented. R V very short . . . . . . . . . . . . . 5
Body oval. Antenna 6-segmented. R V elongate . . . . . . . . . . . . . . . . . . 6
5 Second tarsal segments mostly dusky to dark. ANT PT at least
0.5 x diameter of rhinarium on ANT V. R IV+V 0.07-0.10 mm. Tarsal
claws with distinctly bifurcate apices (Fig. 73) . . . Essigella californica
Second tarsal segments mainly pale, sometimes dusky on distal third.
Primary rhinarium on ANT V exceptionally distal, so that length of PT
is greatly reduced, less than 0.5 x diameter of rhinarium. R IV+V
0.04-0.07 mm. Tarsal claws with apices double but not distinctly bifurcate
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Essigella wilsoni
6 SIPH pores on broad pigmented conical bases 0.4mm or more in
diameter. (Antennal hairs numerous, 80 µm or more long; hind tibial hairs
100-150 µm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara commatula*
SIPH cones very small, with pigmented area less than 0.2 mm in diameter
............................................................. 7
7 All tibiae pale except at apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
At least hind tibia with distal third or more dark . . . . . . . . . . . . . . . . 9
8 Longest hairs on dorsal side of hind tibia less than 65 µm long . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara splendens
Longest hairs on dorsal side of hind tibia 90-160 µm long . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pseudotsugae
9 Longest abdominal hairs more than 100 µm long . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara vagabunda*
Longest abdominal hairs less than 100µm long . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pseudotaxifoliae
PSYCHOTRIA Rubiaceae
PTEROCARPUS Leguminosae
PTEROSPERMUM Sterculiaceae
PTEROSTYRAX Styracaceae
Rhopalosiphum insertum
Sappaphis piri
Schizaphis punjabipyri, pyri
Toxoptera aurantii, citricidus,
odinae
P. granulosa Nippolachnus bengalensis
P. korshinskyi Dysaphis pyraria, reaumuri
P. kumaoni Melanaphis pahanensis
P. lanata [Dysaphis pavlovskyana]
P. montana Aphis spiraecola
[Brachysiphoniella montana]
Sappaphis piri
P. nivalis Dysaphis pyri
P. pashia Dysaphis sharmai
Melanaphis pahanensis
Nippolachnus bengalensis
Pyrolachnus pyri
P. pulcherrima Aphis gossypii
P. pyrifolia (incl. var. culta) Aphanostigma iaksuiense
(Dolbae, Chinese or Japanese Pear) Aphis gossypii, pomi, spiraecola
Aulacorthum magnoliae
Brachycaudus helichrysi
Longistigma caryae
Melanaphis siphonella
Myzus persicae
Nippolachnus piri
Ovatus crataegarius
Prociphilus kuwanai, oriens
Rhopalosiphum insertum,
nymphaeae, rufiabdominalis
Sappaphis piri
Schizaphis piricola
Toxoptera odinae
P. salicifolia Dysaphis reaumuri
P. syriaca Aphanostigma piri
Dysaphis reaumuri
P. ussuriensis Dysaphis pyri
Prociphilus kuwanai
Sappaphis piri
P. vestita Eurnyzus eastopi
Pyrus 383
8 ANT III similar in length to R IV+V. Wax pore plates with narrow, mostly
interconnecting central areas . . . . . . . . . . . . . . . . . . Eriosoma lanigerum
- ANT III less than 0.5 x R IV+V. Wax pore plates with separate, circular
or polygonal clear central areas . . . . . . . . . . . . . . Eriosoma americanum
9 BL of spring migrant alata from leaf-nest gall 4mm or more. R IV+V
more than 0.2 mm long and bearing 7 or more accessory hairs . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
- BL of spring migrant alata less than 4 mm. R IV+V less than 0.16 mm long
and bearing 2-5 accessory hairs . . . . . . . . . . . . . . . Prociphilus kuwanai
10 Body elongate, 2 or more x longer than wide. Antennae and legs pale
except for black tibial apices and hind tarsi. (Alata with a black quadrate
dorsal abdominal patch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Body broadly oval, less than 2 x longer than wide. Antennae and legs
mainly dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 ANT PT/BASE 0.25-0.35, without any long fine hairs distal to primary
rhinarium. R I V + V 0.14-0.18 mm long, 0.62-0.75 x HT II, and bearing
9-16 accessory hairs . . . . . . . . . . . . . . . . . . . . . Nippolachnus bengalensis
- ANT PT/BASE 0.45-0.75, with some long fine hairs distal to primary
rhinarium. R IV+V 0.17-0.22 mm long, 0.7-1.0 x HT II, and bearing
more than 20 accessory hairs . . . . . . . . . . . . . . . . . . . . Nippolachnus piri
12 Abdomen with paired, pigmented spinal tubercles on each tergite. Alata
with maculate forewings and a short pterostigma . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterochloroides persicae
- Abdomen without spinal tubercles. Forewings of alata not maculate, with
a long pterostigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 HT I (measured on ventral side) less than 2 x its basal diameter, less than
0.35 x HT II. Forewing of alata with pterostigma almost straight, not
extending around tip of wing . . . . . . . . . . . . . . . . . . . . Pyrolachnus pyri
- HT I more than 2 x its basal diameter, more than 0.35 x HT II. Forewing
of alata with pterostigma curved distally, extending around wing-tip
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 HT II 1.8-2.3 x HT I . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- HT II about 2.6 x HT I . . . . . . . . . . . . . . . . . Longistigma xizangensis*
15 SIPH shorter than CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
- SIPH as long as or longer than CAUDA . . . . . . . . . . . . . . . . . . . . . . . 20
16 Abdomen without any dorsal markings . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Abdomen with dark dorsal markings . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17 Stridulatory ridges present on abdominal sternites 5 and 6. CAUDA dark,
rounded at apex (Fig. 121O) . . . . . . . . . . . . . . . . . . . . . Toxoptera odinae
Pyrus 385
40 First tarsal segments all with only 2 hairs. Marginal tubercles well
developed on ABD TERG 1 and 7 . . . . . . . . . . . . . . . Aphis chetansapa
- First tarsal segments with 3 hairs at least on fore and mid-legs. Marginal
tubercles if present then on ABD TERG 2-5 (or 2-6) as well as on 1
and 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41 Antenna with numerous long very fine hairs, the longest more than
2 x basal diameter of ANT III. (Alata with separate transverse bars on
dorsal abdomen) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
- Longest hairs on ANT III less than 1.8 x basal diameter of segment.
SIPH without hairs. (Alata with a more-or-less solid dark patch on dorsal
abdomen) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
42 SIPH 2 or more x basal width, and sometimes bearing 1 or more hairs.
CAUDA rounded, much shorter than its basal width . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sappaphis piri
- SIPH less than 1.5 x basal width, without hairs. CAUDA bluntly
pointed, about as long as its basal width . . . . . Melanaphis pahanensis
43 Spiracular openings large and rounded . . . . . . . . . . . . . . . . . . . . . . . . . 44
- Spiracular openings small and reniform . . . . . . . . . . . . . . . . . . . . . . . . 46
44 SIPH less than 2 x CAUDA. Dorsal abdomen dusky or pale. Meso-
sternum without mammariform processes . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus almatinus/helichrysi
- SIPH more than 2 x CAUDA. Dorsal abdomen usually dark. Anterior
part of mesosternum with a pair of mammariform processes . . . . . 45
45 R IV+V more than 0.17 mm, more than 0.6 x SIPH. Dorsal abdomen and
SIPH pale or dark. (Alata with secondary rhinaria only on ANT III)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus cardui
- R IV+V less than 0.17mm, about 0.5 x SIPH or less. Dorsal abdomen
always extensively sclerotized, black, and SIPH black. (Alata with secon-
dary rhinaria on ANT III and IV, or III, IV and V) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus persicae
46 Antennae at least as long as distance from frons to SIPH bases (in alata
about as long as body). SIPH more than 0.3 mm long, black . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis plantaginea
- Antennae shorter than distance from frons to SIPH bases (in alata less
than BL). SIPH pale or dark, less than 0.3 mm long . . . . . . . . . . . . 47
47 R IV+V 0.18-0.21 mm, 1.4-1.5 x HT II. Marginal tubercles present on
ABD TERG 1-5 and 7. Spring migrant alata (s.m.a.) with 13-22 second-
ary rhinaria on ANT I I I , 2-5 on IV, 0 on V . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis tschildarensis
Quercus 389
QUALEA Vochysiaceae
About 225 aphid species in more than 40 genera feed on oaks. Some species,
notably of Hormaphidinae, seem restricted to subgenus Cyclobalanopsis, but
within the main subgenus Euquercus there is no clear pattern of host
specificity relating to the sections of the subgenus; e.g. the North American
aphid species complexes within Myzocallis, subgenera Neomyzocallis and
Lineomyzocallis, are thought to be mainly associated with red oaks (sect.
Erythrobalanus), yet many species in these groups are sometimes recorded
390 Quercus
Hoplochaitophorus quercicola
Myzocallis bella, discolor, mimica,
punctata, walshii
Neosymydobius albasiphus,
mimicus
Phylloxera querceti, rileyi, ?stellata
Stegophylla querci
Stomaphis quercus
Thelaxes californica
Tuberculatus pallidus,
tuberculatus
Q. aliena Diphyllaphis quercus
(Galcham, Korean Oak) Greenidea nipponica
Tuberculatus capitatus, indicus,
querciformosanus, stigmatus,
yokoyamai
Q. arizonica Tuberculatus pallidus
Q. bicolor Hoplochaitophorus spiniferus
(Swamp White Oak) Myzocallis discolor, multisetis,
punctatus, walshii
Phylloxera rileyi
Thelaxes dryophila
Q. boisieri = var. of Q. infectoria
Q. borealis = Q. rubra
Q. calliprinos Hoplocallis microsiphon, [pictus]
(Prickly-leaved Evergreen Oak) Lachnus roboris
Myzocallis cocciferina, ?tauricus
Phylloxera quercus
Siculaphis vittoriensis
Thelaxes suberi
Tuberculatus maculipennis,
pallescens
Q. canariensis Myzocallis mediterraneus
(Mirbeck's Oak) Tuberculatus africanus, annulatus
Q. castanaefolia Myzocallis boerneri, castanicola
(Boland Masu, Persian Oak) Thelaxes suberi
Tuberculatus annulatus, pallidus
Q. castanea Myzocallis pepperi ssp. iturbide
(Encino, Mexican Oak) Stegophylla mugnozae
Q. catesbei = Q. laevis
392 Quercus
Q. floribunda = Q. dilatata
Q. formosana Greenidea nigra
Mollitrichosiphum fasciatum
Q. frainetto Phylloxera texana
(Hungarian or Italian Oak) Tuberculatus eggleri, etruscus
Q. frutex? Mexicallis analiliae ssp. pumilus
Q. gambelii (incl. hybrids, e.g. Lachnus allegheniensis
'undulata') Myzocallis discolor
(Gambel Oak) Neosymydobius chrysolepis
Stegophylla quercicola, quercifoliae
Thelaxes californica
Tuberculatus kiowanicus
Q. garryana Myzocallis punctata
(Oregon White Oak) Stegophylla essigi
Thelaxes californica
Tuberculatus annulatus,
columbiae, pallidus
Q. gemelliflora [Eutrichosiphum sp. (Noordam, in
prep.)]
Q. gilva Lachnus tropicalis
(Ichii-gashi) Metathoracaphis isensis
Quernaphis tuberculata
Q. glandulifera = Q. serrata
Q. glauca Allothoracaphis piyananensis
(Ara-kashi, Barin, Japanese Oak) Allotrichosiphum kashicola
Dermaphis japonensis
Eutrichosiphum dubium, [sp.
(Noordam, in prep.)]
Greenidea kuwanai, quercifoliae,
[sp. (Noordam, in prep.)]
Kurisakia querciphila
Lachnus tropicalis
Mollitrichosiphum glaucae,
nigrofasciatum
Neothoracaphis glaucae,
saramaoensis
Nipponaphis distyliicola, monzeni
Parathoracaphis setigera
Xenothoracaphis kashifoliae
Q. griffithi Eutrichosiphum khasyanum
Tuberculatus indicus, quercicola
Q. grisea Neosymydobius ajuscanus
Q. gunnisonii Lachnus allegheniensis
Myzocallis punctatus
Thelaxes californica
Tuberculatus kiowanicus
Q, havardii Phylloxera tuberculifera
Myzocallis granovskyi,
longiunguis, melanocera,
meridionalis, spinosa
Neosymydobius luteus
Q. nuttallii Myzocallis frisoni
Q. obtusiloba = Q. stellata
Q. pagoda = Q. falcata var. pagodifolia
Q. palustris Longistigma caryae
(Pin Oak, Swamp Spanish Oak) Myzocallis discolor, exultans,
frisoni, granovskyi, longiunguis,
melanocera, multisetis,
punctata, spinosa, walshii
Neosymydobius albasiphus
Patchia winforii
Q. paucidentata Neothoracaphis elongata
Q. peduncularis Neosymydobius butzei 'var.
papillata'
Tuberculatus mexicanus
Q. pedunculata = Q. robur
Q. persica Hoplochaetaphis zachvatkini
(Manna Oak) Lachnus swirskii ssp. persicae,
Lachnus sp.
Myzocallis komareki, persica
Thelaxes suberi
Tuberculatus maximus
Q. petraea Diphyllaphis mordvilkoi
(Bergek, Durmast Oak) Hoplochaetaphis zachvatkini
Lachnus pallipes, roboris
Moritziella corticalis
Myzocallis castanicola
Phylloxera coccinea, confusa,
foaae, glabra, italica, quercus
Stomaphis quercus
Thelaxes dryophila, suberi
Tuberculatus annulatus, eggleri,
borealis. neglectus
Q. phellos Myzocallis castanicola, exultans
(Willow Oak) Stegophylla quercicola
Tuberculatus querceus
Q. phylliraeoides Atarsaphis agrifoliae
(Ubame-gashi) Diphyllaphis alba
Quercus 399
Eutrichosiphum tattakanum
Greenidea kuwanai, nipponica
Lachnus tropicalis
[Myzocallis punctata]
Tuberculatus pallidus, pilosus
Q. pontica Tuberculatus annulatus
Q. prinus Hoplochaitophorus heterotrichus,
(Chestnut Oak) quercicola
Myzocallis discolor, melanocera,
punctatus
Thelaxes californica
Q. pseudoturneri Tuberculatus borealis
Q. pubescens Diphyllaphis mordvilkoi
(Downy Oak) Hoplocallis microsiphon
Hoplochaelaphis zachvatkini
Lachnus iliciphila, roboris
Myzocallis castanicola,
mediterraneus, occidentalis
Phylloxera foaae, quercus
Thelaxes dryophila, suberi
Tuberculatus annulatus, eggleri,
inferus, maculipennis
Q. pyrenaica Diphyllaphis mordvilkoi
(Rebollo, Pyrenean Oak) Lachnus roboris
Myzocallis annulatus, castanicola,
eggleri, komareki, occidentalis,
schreiben
Tuberculatus remaudierei
Q. robur (incl. var. fastigiata) Acanthochermes quercus
(English Oak) Foaiella danesii
Hoplocallis microsiphon, ruperti
Hoplochaetaphis zachvatkini
Lachnus pallipes, roboris
Myzocallis bella, castanicola
Phylloxera coccinea, confusa,
foaae, glabra, italica, quercus
Stomaphis quercus
Thelaxes dryophila, suberi
Tuberculatus annulatus, borealis,
moerickei, neglectus, querceus
Q. rubra Diphyllaphis microtrema
(Northern Red Oak) Lachnochaitophorus querceus
Myzocallis bella, castanicola,
400 Quercus
exultans, granovskyi,
longirostris, longiunguis,
melanocera, multisetis, occulta,
punctata, spinosa, walshii
Stegophylla quercifoliae, quercina
Q. rugosa Mexicallis analiliae, areolatus,
(Netleaf Oak) spinifer
Tuberculatus mexicanus, spiculatus,
spiculatus ssp. rebecae
Q. salicifolia Myzocallis spinosa
Q. semicarpifolia Eutrichosiphum khasyanum, pyri
Myzocallis polychaeta
Q. serrata (incl. var. donarium) Cervaphis quercus
(Ko-nara) Diphyllaphis konarae, quercus
Eutrichosiphum davidi,
[narafoliae], pasaniae, taoi,
tattakanum
Greenidea kuwanai, quercifoliae
Kurisakia querciphila
Lachnus tropicalis
Mollitrichosiphum nigrofasciatum
Myzocallis castanicola, kuricola
Neothoracaphis garwhalensis,
quercicola, yanonis
Stomaphis japonicus
Tuberculatus capitatus, higuchii,
kashiwae, konaracola,
quercicola, stigmatus, yokoyamai
[ Tuberculatus fulviabdominalis,
kunugi, naganoe (Shinji, 1941)]
Q. sessiliflora = Q. petraea
Q. sessilifolia Neothoracaphis elongata
Q. shumardii Myzocallis elliotti, granovskyi,
(Shumard Red Oak) longiunguis, melanocera,
meridionalis, spinosa
Q. sinuata = Q. durandii
Q. spinosa Neothoracaphis tarakoensis
Q. stellata Hoplochaitophorus quercicola
(Iron Oak, Post Oak) Neosymydobius albasiphus,
quercihabitus
Phylloxera rileyi
Stegophylla quercifoliae
Quercus 401
1 In galls on leaves, with only two generations per year and no alatae;
fundatrices produce sexuales. Primary rhinarium on ANT III protruding
404 Quercus
Fig. 90. A, ANT III of fundatrix of Acanthochermes quercus', B, stellate process of immature fundatrix of
A. quercus (after Grassi, 1912); C, stellate process of aptera of Phylloxera stellata (after Duncan,
1922); D, ANT III of aptera of Ph. glabra; dorsal processes of E, Foaiella danesii (after Grassi, 1912),
F-H, Phylloxera quercus group, I, Ph. rileyi (after Duncan, 1922), J, Ph. reticulata (after Duncan, 1922),
K, Ph. confusa (fundatrix, after Grassi, 1912), L, Ph. confusa (later generation aptera, after Grassi,
1912), M, Ph. kunugi (after Miyazaki and Teramoto, 1991), N, Ph. quercina, 0-P, Moritziella corticalis.
- Longest dorsal processes less than 3 x their basal width, with apical hair
long, 5-6 x its apical diameter (Fig. 90J). ANT III rather stout, broadest
on distal half. Dorsal cuticle finely reticulate . . . . Phylloxera reticulata*
10 Dorsal processes with broad, bract-like spicules (Figs90K, L). Apical
hairs on spinal processes long, at least 4 x their apical diameter ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera confusa*
- Dorsal processes with narrower, pointed or denticulate spicules, and with
short apical hairs which are usually less than 3 x their apical diameter
(except when the processes are very short) . . . . . . . . . . . . . . . . . . . . . . 11
11 All thoracic tergites and ABD TERG 1 with pleural as well as marginal
processes. Dorsal cuticle with nodulose ornamentation. Spiracles evident
on ABD TERG 2-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- Pleural processes absent from hind margin of pronotum and from ABD
TERG 1. Dorsal cuticle variously ornamented. Abdominal spiracles
present only on ABD TERG 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
406 Quercus
12 BL 2.0-2.5 mm. Dorsal processes conical, all less than 2 x their basal
diameters (Fig. 90M) . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera kunugi
- BL 0.9-1.5 mm. Longest dorsal processes have attenuated, cylindrical
apices and are more than 3 x longer than their basal diameters (Fig. 90N)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxerina quercina
13 Rostrum short; when directed backwards not reaching to hind coxae.
Longest dorsal processes shorter than HT II. Dorsal cuticle with minute,
short ridges or wrinkles . . . . . . . . . . . . . . . . . . . . . . Phylloxera similans*
- Rostrum reaching well beyond hind coxae. Longest dorsal processes
usually longer than HT II; if a little shorter then the processes are pig-
mented and the dorsal cuticle is very rugose . . . . . . . . . . . . . . . . . . . . 14
14 Thoracic and anterior abdominal tergites with very rugose, denticulate
sculpturing. Dorsal processes pigmented, the longest usually not more
than 2 x HT II and often much shorter (Figs 90O, P) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moritziella corticalis
- Dorsal cuticle less rugose, variously ornamented. Longest dorsal processes
often more than 2 x HT II, degree of pigmentation unknown . . . 15
15 Longest dorsal processes more than 0.8 x hind femora, which are rather
stout. Tibiae without annular ridges . . . . . . . . . . . Phylloxera querceti*
- Longest dorsal processes less than 0.7 x hind femora, which are relatively
slender. Tibiae with annular ridges . . . . . . . . . . . Phylloxera davidsoni*
1 SIPH present as small pores or shallow cones on the posterior part of the
'abdominal plate' (= fused ABD TERG 2-7). Tarsi always 2-segmented
with claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- SIPH completely absent. Tarsi variably developed, reduced in some
species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Quercus 407
Fig. 91. A, aptera of Nipponaphis monzeni with detail of central part of prosoma; detail of similar area
of prosoma of B, N. distyliicola, C, Paranipponaphis takaoensis, D, Allothoracaphis piyananensis,
E, Pseudothoracaphis himachali', dorsal (right) and ventral views of aptera with detail of spines on ABD
TERG 7 of F, Parathoracaphis setigera, 6, P. kayashimai; H, end of abdomen of aptera of
Parathoracaphis gooti; I, dorsal view of Atarsaphis agrifoliae; J, dorsal view of Quernaphis tuberculata',
K, dorsal (right) and ventral views of Neothoracaphis yanonis; L, marginal hairs of Dermaphis sp.
Quercus 411
1 SIPH with short hairs. Body with very large, branched, hair-bearing pro-
cesses (Fig. 92A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cervaphis quercus
- SIPH with long hairs. Body without large branched hair-bearing processes
............................................................. 2
2 CAUDA usually with an apical papilla (Fig. 92B). SIPH always with pale
reticulation, at least at base often snapped off at base in preserved
specimens). Body often brown, very broadly pear-shaped and rather flat
(genus Greenidea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- CAUDA never with an apical papilla. SIPH usually without any reticula-
tion. Body broadly pear-shaped, oval or elongate . . . . . . . . . . . . . . . . 8
412 Quercus
3 SIPH with pale reticulation extending over most of length (except at apex)
............................................................. 4
- SIPH reticulated only at base (subgenus Trichosiphum) . . . . . . . . . . . 5
4 ANT III reticulated over most of length except at extreme base, and also
distinctly imbricated distally . . . . . . . . . . . . . . . . Greenidea querciphaga
- ANT III neither reticulated nor imbricated . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea quercifoliae/(mushana)
5 SIPH very short and stout (Fig. 92D), less than 0.18 x BL and less than
0.75 x head width across (and including) eyes . . . . Greenidea haldari
- SIPH longer, more than 0.2 x BL, more than 0.8 x head width across
eyes (e.g. Fig. 92E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 R IV+V 2.4-3.0 x HT II . . . . . . . . . . . . . . . . . . . . Greenidea quercicola
- R IV+V 1.2-2.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Venter of abdomen strongly and densely spinulose. SIPH 0.29-0.36 x BL
(Fig. 92E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea kuwanai
- Venter of abdomen smooth or only locally or weakly spinulose, at least
on anterior sternites. SIPH 0.20-0.30 x BL . . . . Greenidea nipponica
8 Hind tibia with a series of narrow (stridulatory?) tranverse ridges spaced
out on basal 0.5-0.9 of length (e.g. Fig. 57D; genus Mollitrichosiphum)
............................................................. 9
- Hind tibia without transverse ridges (but often imbricated distally) . . . 15
9 Antenna 5- or 6-segmented. Dorsum pale. Hind tibia with over 50 trans-
verse, very narrow ridges extending about 0.9 of its length . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum niitakaensis
- Antenna 6-segmented. Dorsum pale or dark, or with a dark patch. Hind
tibia with only 14-30 transverse ridges on basal 0.5-0.6 . . . . . . . . . . 10
10 R IV+V 2.2-2.6 x HT I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- R IV+V 1.4-2.1 X HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Body pear-shaped, BL less than 2 x maximum width of abdomen . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum luchuanum
- Body elongate, BL more than 2 x maximum width of abdomen . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum godavariense
12 Body elongate, BL 2.3-2.5 x maximum width of abdomen. SIPH pale,
0.5-0.6 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Abdomen broader, BL about 2 x maximum width of abdomen or less.
SIPH dark, 0.3-0.4 x BL. (Dorsal abdomen with dark lateral areas linked
by a broad cross-band) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Quercus 413
Fig. 92. A, hair-bearing marginal process of aptera of Cervaphis quercus; B, CAUDA of Greenidea
quercifoliae; C, body hairs of aptera of Allotrichosiphum kashicola; SIPH of aptera of D, Greenidea
haldari, E, G. kuwanai, F, Eutrichosiphum tattakanum, G, Eu. heterotrichum (from type series); spinules
on dorsum of aptera of H, Eu. davidi and I, Eu. pasaniae; J, ANT III of aptera of Eu. dubium.
414 Quercus
- SIPH pores on conical bases that are usually large, pigmented and clearly
demarcated, but sometimes small and pale; flange small. Hairs numerous,
but not forming a dense pile; hair bases more than 20 µm apart, at least
on dorsum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 R IV+V 1.20-1.31 x HT II . . . . . . . . . . . . . . . . . . . Lachnus takahashii
- R IV+V 0.65-1.12 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 R IV+V 0.65-0.82 x HT II, but if 0.79-0.82 x HT II then less than 0.2
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- R IV+V 0.80-1.12 x HT II; if less than 0.85 x HT II then more than
0.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 Maximal basal diameter of SIPH cone 0.16-0.24 mm. ANT PT/BASE a
little less than 0.5. R IV+V 0.74-0.82 x HT II. Hairs on ABD TERG
1-3 all shorter than basal diameter of ANT III . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus acutihirsutus
- Maximal basal diameter of SIPH cone 0.3-0.7 mm. ANT/PT BASE 0.5
or more. R IV+V 0.65-0.70 x HT I I . Longest hairs on ABD TERG 1-3
more than 1.5 x basal diameter of ANT III . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus allegheniensis
9 Hind tibia with some of more distal hairs on ventral side very long and
fine-pointed, at least 2-3 x longer than neighbouring short hairs
(Fig. 93D). Mesosternal processes absent or only weakly developed. SIPH
cones small and often pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Hind tibia with hairs on ventral side often longer and finer distally, but
not with some hairs 2-3 times longer than others. Mesosternum of summer
aptera with a pair of mammariform or papilliform processes. SIPH cones
usually large and pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 Dorsal body hairs long and fine-pointed . . . . . . . . . . . Lachnus pallipes
- Dorsal body hairs very short (less than 30 µm) and blunt . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Lachnus sp. ex Quercus aegilops, Turkey
11 SIPH cones small and pale. Hairs on ABD TERG 1-6 all short and blunt,
maximally about 20 µm . . . . . Lachnus sp. ex Quercus persica, Turkey
- SIPH cones usually large (basal diameter more than 0.3 mm) and well-
pigmented. Hairs on ABD TERG 1-6 variable, but the longest always
more than 20 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 ANT PT/BASE 0.5 or more. Mesosternal processes mammariform
(Fig. 93B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus tropicalis
- ANT PT/BASE clearly less than 0.5. Mesosternal processes papilliform
(Fig. 93C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 ANT PT shorter than diameter of primary rhinarium on VI (Fig. 93E).
Hairs on ABD TERG 1-3 mostly thick, blunt and less than 30 µm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus swirskii
418 Quercus
Fig. 93. K, SIPH of aptera of Lachnus crassicornis, dorsal and side views; mesosternal processes of
apterae of B, L. tropicalis and C, L. roboris; D, distal part of hind tibia of L. pallipes; ANT VI of
E, L. swirskii and F, L roboris.
9 ANT III with 7-14 secondary rhinaria. ABD TERG 1-4 with spinal pro-
cesses all of similar size . . . . . . . . . . . . . . . . . . . Tuberculatus kiowanicus
- ANT III with 1-7 secondary rhinaria. ABD TERG 1 and 2 with spinal
processes about twice as tall as those on more posterior tergites . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 R IV+V 0.11-0.13 mm with usually 2 pairs of lateral accessory hairs as
well as some ventral hairs. Head with anteriodorsal hairs slightly but
distinctly capitate . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus quercifolii
- R IV+V 0.095-0.11 mm, usually with 1 pair of lateral accessory hairs as
well as some ventral ones. Head with anteriodorsal hairs blunt but not
distinctly capitate . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus pallidus
11 First tarsal segments all with 5 ventral hairs (as well as 2 dorsal ones).
Immatures with spiculose dorsal hairs (Fig. 94B; spicules also discernible
on the longest dorsal hairs of embryos within adult alata) (subgenus
Toltecallis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- First tarsal segments all or mostly with 6 (or 7) ventral hairs. Dorsal hairs
of immatures and embryos without spicules . . . . . . . . . . . . . . . . . . . . . 14
12 ANT PT/BASE more than 1.5, and PT more than 1.6 x R IV+V. ANT
II pigmented. (ABD TERG 1-4 with spinal processes) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus mexicanus
- ANT PT/BASE less than 1.3, and PT less than 1.4 x R IV+V. ANT II
pale. (ABD TERG 1-2 or 1-4 with spinal processes) . . . . . . . . . . . . . 13
13 R IV+V 0.092-0.101 mm. ABD TERG 1-2 (and also mesonotum) with
spinal processes; height of those on mesonotum 0.3-0.5 x HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus garciamartelli
- R IV+V 0.112-0.162 mm. Mesonotum, ABD TERG 1-4, and often 6 also,
with spinal processes; those on mesonotum 0.7-1.1 x HT II . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus spiculatus*
14 Frontal hairs mostly shorter than, and hairs on ANT III all much shorter
than, basal diameter of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- Frontal hairs usually 2 or more x basal diameter of ANT III, and longest
hairs on ANT III often as long as basal diameter or longer . . . . . . 26
15 Forewings with a pattern of infuscation (use hand lens). ABD TERG 1
with a pair of large pigmented spinal processes, those on more posterior
tergites being much smaller . . . . . . . . . . . . . . Tuberculatus tuberculatus
- Forewings mainly pale. Spinal processes on ABD TERG 1 often unpig-
mented, and never larger than those on ABD TERG 2-3 (subgenus Tuber-
culoides) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 ABD TERG 1-4 each with a pair of pale spinal processes (e.g. Fig. 94C;
although those on 4 may be small) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Quercus 421
Fig. 94. A, spinal process of Tuberculatus querceus; B, spiculose hairs of immature T. mexicanus;
spinal processes of C, T. borealis and D, T. neglectus (after Krzywiec, 1965).
- Only ABD TERG 1-3 with spinal processes (e.g. Fig. 94D) . . . . . . . 20
17 R IV+V 0.13-0.19 mm, 1.15-1.50 x HT II, with 3 pairs of lateral
accessory hairs (as well as 3-6 ventral ones) . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus remaudierei
- R IV+V 0.08-0.12 mm, 0.80-1.05 x HT II, with 2 pairs of lateral acces-
sory hairs (and 3-5 ventral ones) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 Spinal hairs of embryos (seen through abdominal cuticle of alate vivipara)
all similar in length to the long marginal hairs on the same tergites
(Fig. 95A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus eggleri
- Spinal hairs on at least ABD TERG 2-3 of embryos only half as long as
marginal hairs on same segments or shorter . . . . . . . . . . . . . . . . . . . . 19
19 Embryos with spinal hairs on ABD TERG 1-6 all about 5-8 µm long, thin,
with blunt apices; marginal hairs 15-45 µm long, with those on at least
ABD TERG 2-3 not overlapping those of next tergite (Fig. 95B) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus borealis
- Embryos with distinctly capitate spinal hairs, those on ABD TERG 1
being 6-10 µm long, those on more posterior tergites 26-75 µm; marginal
hairs 26-75 µm long, all except those on ABD TERG 1 overlapping those
on next tergite (Fig. 95C) . . . . . . . . . . . . . . . . . . . Tuberculatus africanus
20 Spinal hairs of embryos mostly very long and overlapping between
segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- Spinal hairs on ABD TERG 1-5 of embryos all very short (less than 5 µm)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
21 ANT PT/BASE 2.1-2.8. R IV+V 1.20-1.35 X HT II and bearing 12-15
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus etruscus
- ANT PT/BASE 0.9-1.8. R IV+V 0.90-1.15 X HT II and bearing 7-10
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
422 Quercus
- Longest hairs on ANT III 1.3-1.5 x basal diameter of segment. ANT III
with 4-7 secondary rhinaria restricted to basal 0.5 of segment even in
spring populations. Cu la , Cu lb and media of forewing all of similar
thickness and degree of pigmentation . . . . . . . . . . . . Tuberculatus paiki
34 Front of head, ANT I-III and dorsal thorax all with strongly capitate
hairs. Dorsal hairs on basal half of hind tibia rather stiff, about 1.0-1.3
x diameter of tibia, with blunt or slightly capitate apices. Embryos with
spinal hairs on ABD TERG 2-6 very thickly capitate . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus capitatus
- Front of head, ANT I-III and dorsal thorax with acute, blunt and/or
slightly capitate hairs. Dorsal hairs on basal half of hind tibia thin with
very fine apices, up to 2 x diameter of tibia. Embryos with spinal hairs
on ABD TERG 2-6 blunt or only slightly capitate . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus fangi
35 Longest hair on ANT II 2 or more x longer than longest hair on ANT
III. Pronotum without any spinal processes (subgenus Camelaphis) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
- Longest hair on ANT II usually similar in length to or shorter than longest
hair on ANT III, or not more than 1.5 x longer. Pronotum with or
without spinal processes (subgenus Acanthocallis) . . . . . . . . . . . . . . . 38
36 Mesonotum pale. Abdomen without dark marginal processes. ABD
TERG 1-3 with mainly pale, sometimes black-tipped, spinal processes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus pallescens
- Mesonotum dark. Abdomen with large, dark, spiculose marginal pro-
cesses, those on ABD TERG 3 being about as long as the siphunculi.
Spinal processes also dark, at least on ABD TERG 2 and 3 . . . . . . 37
37 Forewings maculate. Spinal processes on ABD TERG 1-4 pigmented, and
shorter, pale spinal processes also present on more posterior tergites.
SIPH only dark basally . . . . . . . . . . . . . . . . . Tuberculatus maculipennis
- Forewings not maculate. Abdomen with 2 pairs of dark spinal processes
(very long and basally joined on ABD TERG 3). SIPH wholly dark
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus cornutus
38 Forewings with many fine hairs, not only on posterior margin of ptero-
stigma but also scattered over most of wing membrane distal to Cu lb
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
- Forewings without hairs on wing membrane . . . . . . . . . . . . . . . . . . . . 40
39 All forewing veins evenly and broadly bordered with fuscous over their
entire length. Long hairs on inner side of ANT III very finely pointed.
Spinal processes on ABD TERG 1-3, especially those on 3, much larger
and darker than those on ABD TERG 4-5 . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus quercicola
Quercus 425
Fig. 96. A, dorsal body markings of Hoplocallis pieta; B, ANT III of Myzocallis kuricola; C, CAUDA and
anal plate of M. granovskyi; D, body and wing markings of M. bella; E, head and prothorax of
M. melanocera; spinal sclerites on ABD TERG 4 of F, M. komareki and G, M. cocciferina; pterostigma
of H, M. schreiberi and I, M. komareki.
- ANT III-V usually wholly dusky, whereas ANT I and II are usually pale.
Prothoracic stripes 0.09-0.14 mm wide, with pale area between them at
least as wide as long (Fig. 96E) . . . . . . . . . . . . . . Myzocallis melanocera
68 PT 1.5 or more x ANT III; ANT PT/BASE 2.5-3.0. ANT III with 1-3
(usually 2) secondary rhinaria near base . . . . . . . . Myzocallis tenochca
- PT shorter than ANT III; ANT PT/BASE 1.2-3.2. ANT III with 2-12
(rarely 2) secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
69 Rs dark, with a black spot at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
- R s very pale, especially at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
70 R IV+V less than 0.11 mm, 0.80-0.95 x HT II . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis punctata (pale form)
- R IV+V 0.11 or more, 1.0-1.2 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis rostropunctata
71 Pterostigma of normal length, i.e. length measured from proximal end of
black patch 3.8-4.7 x maximum width (Fig. 96H) . . . . . . . . . . . . . . 72
- Pterostigma short, trapezoid, the length 2.5-3.8 x maximum width
(Fig. 96I) (subgenus Pasekia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
72 R IV+V stiletto-shaped, 0.15-0.22 mm long, 1.6-2.0 x HT II . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis occidentalis
- R IV+V 0.10-0.16 mm long, 0.8-1.5 x HT II . . . . . . . . . . . . . . . . . . 73
73 R IV+V 1.2-1.5 X HT II, with 4-7 accessory hairs . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis schreiberi
- R IV+V 0.84-1.19 x HT I I , with 4-12 accessory hairs . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis boerneri
74 ANT PT/BASE 2.4-3.2. (Spinal sclerites on ABD TERG 1.5 oval, with
longest hairs 28-50 µm long, having blunt or slightly capitate apices;
Fig. 96G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis cocciferina
- ANT PT/BASE 1.2-2.1. (Spinal sclerites on ABD TERG 1-5 and the
hairs upon them of various shapes and forms, but if longest hairs are more
than 40 µm long then they usually have strongly expanded or capitate
apices) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
75 Longest hairs on vertex very long, about 2.5 x basal diameter of ANT
III, and those on ANT III about equal to basal diameter of ANT III or a
little longer, and distinctly capitate. Primary rhinarium on ANT VI not
ciliated, elongate, 41-50µm long. ANT III with 8-12 secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis persica
- Longest hairs on vertex 0.5-1.2 x basal diameter of ANT III, and those
on ANT III about 0.5 x basal diameter. Primary rhinarium on ANT VI
ciliated, less than 40 µm long. ANT III with 2-8 secondary rhinaria . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
430 Quercus
76 Empodial hairs setiform. Spinal hairs on ABD TERG 1-5 mostly pointed,
and all less than 40 µm long. Sclerites on ABD TERG 4 and 5 often fused
intersegmentally to form a pair of large dark patches . . . . . . . . . . . . . . .
.............................................. Myzocallis taurica
- Empodial hairs spatulate. Longest spinal hairs on ABD TERG 1-5 often
more than 40 µm long, with expanded or capitate apices. Sclerites on ABD
TERG 4 and 5 never fused intersegmentally . . . . . . . . . . . . . . . . . . . . 77
77 Hairs on vertex arising from pigmented sclerites (pale in spring forms).
ABD TERG 1-5 each with a pair of oval to quadrate sclerites that have
a dark margin around a large pale central area (Fig. 96F). Primary
rhinarium on ANT VI 26-33 µm long . . . . . . . . . . Myzocallis komareki
- Vertex without pigmented sclerites at hair-bases, but with paired longi-
tudinal bands of brown pigment running through lateral ocelli (indistinct
in spring forms). Paired spinal sclerites on ABD TERG 1-5 oval or
irregular in shape, often paler in centre but without a clearly defined dark
margin. Primary rhinarium on ANT VI 19-25 µm long . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis mediterraneus
Fig. 97. Spinal hairs on ABD TERG 5 of apterae of A, Thelaxes valtadorosi, B, Th. dryophila,
C, Th. suberi, D, Siculaphis vittoriensis, E, Hoplochaitophorus heterotrichus, f, H. quercicola; G, CAUDA
of Globulicaudaphis pakistanica (ventral view).
8 Antennae 4- or 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Antennae 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
9 Dorsal cuticle strongly and densely nodulose. Frons with 2 long forwardly-
directed hair-bearing processes, and thorax and abdomen with large
lateral processes bearing backwardly-directed hairs . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mexicallis areolatus
- Dorsal cuticle spiculose, wrinkled or very weakly nodulose. Frons without
processes, and thorax and abdomen with marginal hairs on tuberculate
bases not much longer than their basal diameters . . . . . . . . . . . . . . . 10
10 ANT PT/BASE 0.6-0.7. Dorsal hairs on head and pronotum less than
20 µm long. Dorsal cuticle with faint nodulose sculpturing . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mexicallis calvus
- ANT PT/BASE 0.7-1.4. Dorsal hairs on head and pronotum longer than
25 µm. Dorsal cuticle spiculose or wrinkled . . . . . . . . . . . . . . . . . . . . . 11
11 Dorsal cuticle spiculose. Antennae of 4 or 5 segments, with ANT III only
1-2 x SIPH. SIPH 0.046-0.166 mm long, 2-6 x their minimum diameter
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mexicallis analiliae
Dorsal cuticle not spiculose. Antennae always 4-segmented, with ANT III
3.8-6.8 x SIPH. S1PH 0.027-0.044 mm long, less than twice their mini-
mum diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mexicallis spinifer
12 CAUDA bluntly conical, without a constriction, anal plate only weakly
indented. Dorsal spine-like hairs bearing many minute spinules . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Serralocallis takahashii
- CAUDA knobbed, anal plate bilobed. Dorsal spine-like hairs not bearing
minute spicules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Spine-like dorsal hairs outnumbered by fine hairs with finely-pointed
apices (Fig. 97E; sometimes spine-like hairs are few in number, or even
absent in weakly-pigmented midsummer individuals). Longest hairs on
ANT III long and fine, at least twice basal diameter of segment . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplochaitophorus heterotrichus
- Most dorsal body hairs spine-like and on tuberculate bases (e.g. Fig. 97F)
Longest hairs on ANT III 1.0-1.25 x basal diameter of segment . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 SIPH flared apically and almost as long as R IV+V. ANT PT/BASE more
than 0.65 . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplochaitophorus quercicola
- SIPH small, not much widened apically, about 0.5 x R IV+V. ANT
PT/BASE less than 0.60 . . . . . . . . . . . . . Hoplochaitophorus spiniferus
15 Anal plate bilobed, CAUDA with or without a constriction. Wax pore
plates not evident. ANT II shorter than ANT I . . . . . . . . . . . . . . . . . 16
Quercus 433
- Anal plate entire, CAUDA broadly rounded. Wax pore plates evident.
ANT II much longer than ANT I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
16 ANT PT/BASE 1.0 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- ANT PT/BASE more than 1.2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
17 Many of dorsal body hairs very long, up to or exceeding length of ANT
III. CAUDA knobbed . . . . . . . . . . . . . . . . Hoplochaetaphis zachvatkini
- Dorsal body hairs all very much shorter than ANT III. CAUDA with or
without a constriction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 Antennae densely hairy; ANT VI BASE with 7-15 hairs. (Alata with
forewing veins thickly bordered with fuscous) . . . . . . . . . . . . . . . . . . . 19
- Antennae sparsely hairy; ANT VI BASE with 1-3 hairs. (Alata with
forewing veins not or only weakly bordered) . . . . . . . . . . . . . . . . . . . . 21
19 ANT PT/BASE about 0.6 . . . . . . . . . . . . . . . . . . . . . . . Patchia winforii*
- ANT PT/BASE less than 0.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Alata with 4-8 (most commonly 5 or 6) secondary rhinaria on ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnochaitophorus querceus
- Alata with 6-11 (most commonly 8 or 9) secondary rhinaria on ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnochaitophorus obscurus
21 Mid-femora, hind femora and hind tibiae mainly dark . . . . . . . . . . . 22
- Mid-femora, hind femora and tibiae mainly pale . . . . . . . . . . . . . . . . 25
22 ABD TERG 8 with more than 25 hairs . . . . . . . . . . . . . . . . . . . . . . . . 23
- ABD TERG 8 with less than 16 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 24
23 ANT PT/BASE less than 0.4. ABD TERG 1-5 without transverse bars,
but with small sclerites at bases of the very short dorsal hairs. R IV+V
with 8-11 accessory hairs . . . . . . . . . . . . . . . . Neosymydobius agrifoliae
- ANT PT/BASE more than 0.5. ABD TERG 1-5 with transverse bars and
long fine hairs (more than 50 µm). R IV+V with 4-6 accessory hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosymydobius albasiphus/canadensis
24 Hairs on ABD TERG 1-6 pointed, blunt or very slightly capitate. Hairs
on ANT III pointed, more than 0.5 x basal diameter of segment . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosymydobius chrysolepis
- Hairs on ABD TERG 1-6 flared apically. Hairs on ANT III blunt, less
than 0.5 x basal diameter of segment . . . . . . . Neosymydobius butzei
25 ABD TERG 1-7 each with 1-3 (usually 2) marginal hairs on each side
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
- ABD TERG 1-7 each with 3-7 marginal hairs on each side . . . . . . 27
434 Quercus
- SIPH pore only 1-2 x diameter of nearest wax pores. Fore femur 2.5-3.5
x its maximum width. R IV+V with slightly concave sides, either blunt
apically or stiletto-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
34 ABD TERG 8 with 2 hairs. ANT PT/BASE 0.2 or less . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diphyllaphis mordvilkoi
- ABD TERG 8 with 4 hairs. ANT PT/BASE more than 0.3 . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diphyllaphis quercus
35 ABD TERG 8 with 8-10 hairs, CAUDA with 2 hairs. R IV+V blunt
apically (Fig. 98C), with primary (subapical) hairs not displaced proxi-
mally (all on distal third) . . . . . . . . . . . . . . . . . Diphyllaphis microtrema
- ABD TERG 8 with 4 hairs, CAUDA with 3-4 hairs. R IV+V pointed or
stiletto-shaped, with primary hairs displaced proximally (Fig. 98B) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 ANT II shorter than ANT III. R IV+V shorter than HT II. ABD TERG
6-8 fused . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diphyllaphis konarae
- ANT II longer than ANT III. R IV+V longer than HT II. ABD TERG
6-8 with separate sclerites . . . . . . . . . . . . . . . . . . . . . . . Diphyllaphis alba
37 Second tarsal segments with lateral and often also ventral hairs at about
midlength, in addition to the subapical hairs . . . . . . . . . . . . . . . . . . . . 38
- Second tarsal segments with only subapical hairs . . . . . . . . . . . . . . . . 39
38 R IV+V 0.7-0.85 x HT II, with one pair of primary hairs displaced
proximally (in addition to a pair of accessory hairs; Fig. 98D) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegophylla quercicola
- R IV+V 0,95-1.05 x HT II, with subapical hairs not displaced . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegophylla mugnozae
39 R IV+V 0.13-0.15 mm long, 1.2 or more x HT II . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegophylla quercifoliae
- R IV+V 0.07-0.10 mm long, 0.7-1.1 x HT II . . . . . . . . . . . . . . . . . . 40
436 Radermachera
RADERMACHERA Bignoniaceae
Myzus persicae
RANDIA Rubiaceae
RAPANEA Myrsinaceae
RAPHIA Palmae
Cerataphis variabilis
RAUWOLFIA Apocynaceae
RHAMNUS Rhamnaceae
Macchiatiella rhamni
Toxoptera aurantii
Rh. alnifoliae Aphis nasturtii
Rh. alpina Macchiaüella rhamni
Rh. californica Sitobion rhamni
Rh. cathartica Aphis commensalis, frangulae
group, gossypii, mammulata,
nasturtii, spiraecola
Macchiatiella rhamni, rhamni ssp.
tarani
Rh. dahurica Aphis glycines, gossypii
Rh. frangula Aphis frangulae group, gossypii,
nasturtii
Macchiatiella rhamni
Rh. japonica Aphis gossypii
Macchiatiella itadori
Rh. koraiensis Aphis gossypii
Rh. lanceolata Macchiatiella rhamni
Rh. libanoticus Aphis craccivora, frangulae group,
spiraecola
Rh. nepalensis Myzus ornatus
Rh. palaestinus Aphis craccivora, frangulae group,
spiraecola
Rh. purshiana Aphis gossypii
Macchiatiella itadori, rhamni
[Myzus lythri]
Sitobion rhamni
Rh. utilis Aphis utilis
Rh. virgatus Aphis rhamnifila
Rhamnus sp. Aphis fabae group
Myzus persicae
RHAPIS Palmaceae
A revised and extended version of the key given in Blackman and Eastop
(1984) follows the host plant list. Subgenus Toxicodendron, which does not
contain any trees, is not included.
[Macrosiphum euphorbiae]
Nurudea ibofushi, shiraii,
yanoniella
[Prociphilus oriens]
Schlechtendalia chinensis
Toxoptera odinae
Rh. succedanea Carolinaia japonica
(Japanese Sumac, Wax-tree) Toxoptera odinae
Rh. trichocarpa Carolinaia japonica, nigra
Rh. typhina Carolinaia rhois
(Staghorn Sumac) Melaphis rhois
Rh. vernicifera Carolinaia rhois
(Varnish Tree) Stomaphis rhusivermiciflua
Rh. viminalis Aphis gossypii
Myzus persicae
Rhus spp. indet. Aphis gossypii
Aulacorthum magnoliae
RICINUS Euphorbiaceae
ROBINIA Leguminosae
Fig. 99. Forewing of A, Schlechtendalia chinensis and B, Kaburagia rhusicola; antenna of alata from gall
of C, Schlechtendalia elongallis (from Tsai and Tang, 1946), D, S. chinensis, E, Kaburagia rhusicola
(from Tsai and Tang, 1946, as Macrorhinarium ovogallis), F, Melaphis rhois (6-segmented), G, Nurudea
meitanensis (from Tsai and Tang, 1946, as Floraphis), H, Melaphis rhois (5-segmented), I, Nurudea
ibofushi, J, N. shiraii, K, N. yanoniella.
446 Rothmannia
Appendiseta robiniae
Myzus persicae
[Protrama orientalis Narzikulov,
1962]
[ Tinocallis zelkowae]
ROTHMANNIA Rubiaceae
SALACCA Palmae
Over 120 aphid species feed on Salix, the main groups belonging to the genus
Chaitophorus (48 species), the Pterocommatini (29 species in 4 genera) and
the genus Cavariella (20 species), the latter having host alternation between
Salix and Umbellifereae or Araliaceae. Although almost all willow-feeding
aphids restrict themselves exclusively to the genus Salix there are no clear
patterns of host specificity within the genus. Therefore, following the host
plant list, the aphids are keyed for the genus as a whole; the three main groups
noted above are taken out at couplets 16-17 and keyed separately in sup-
plementary keys A, B and C respectively.
Stomaphis longirostris
Tuberolachnus salignus
S. amygdaloides Chaitophorus pusillus
(Peachleaf Willow) Macrosiphum californicum
S. apennina Chaitophorus salicti
S. arbuscula Cavariella intermedia
S. arbustifolia Cavariella salicicola
S. arctica Cavariella borealis
S. aurita Aphis farinosa
Cavariella intermedia
Chaitophorus capreae, ramicola,
salicti
Plocamaphis flocculosa
Pterocomma jacksoni, konoi,
pilosum
S. babylonica Aphis farinosa, gossypii
(Weeping Willow) Cavariella aegopodii, aquatica,
bonica, indica, konoi,
pastinacae, saliapterus,
salicicola, salicis
Chaitophorus beuthani, niger,
pakistanicus, salijaponicus,
saliniger, truncatus, viminalis,
vitellinae
Lachnus salicis
Phylloxerina salicis
Plocamaphis salijaponica
Pterocomrna bicolor, pilosum,
salicis, [salijaponica],
sanguiceps
Toxoptera aurantii
Tuberolachnus salignus
S. bakko Chaitophorus matsumurai
Prociphilus oriens
S. bebbiana Chaitophorus horii, saliapterus
Pterocomma konoi, rufipes
S. brachypoda Chaitophorus niger
S. caesia Chaitophorus truncatus
S. canariensis Tuberolachnus salignus
S. caprea Aphis farinosa
(Goat Willow, Sallow) Cavariella aegopodii,
Salix 449
archangelicae, nipponica,
pastinacae, theobaldi
Chaitophorus capreae, flavissimus,
furcatus, horii, niger,
quinquemaculatus, ramicola,
salicti, salijaponicus, vitellinae
[Lachnus distinguendus Dahlbom,
1851]
Pachypappa warschavensis
Phylloxerina capreae
Plocamaphis flocculosa (sspp.
brachysiphon, goernitzi)
Pterocomma jacksoni, konoi,
pilosum, rufipes, salicis
S. cardiophylla Pterocomma rufipes
S. caroliniana Chaitophorus longipes, minutus
Phylloxerina sp.
Tuberolachnus salignus
S. chaenomeloides Phylloxerina capreae
Tuberolachnus salignus
S. chrysostella Cavariella aegopodii
S. cinerea Aphis farinosa
(Grey Willow) Cavariella aegopodii, japonica,
theobaldi
Chaitophorus capreae, niger,
ramicola, salicti
Plocamaphis flocculosa (sspp.
brachysiphon, goernitzi)
Pterocomma jacksoni, konoi,
pilosum, rufipes, salicis
Tuberolachnus salignus
S. cordata Chaitophorus nigrae, viminalis
S. crassijulis Plocamaphis coreana
S. daphnoides Aphis farinosa (incl. ssp.
(Violet Willow) yanagicola)
Cavariella aegopodii
Chaitophorus salicti
Phylloxerina daphnoides
Pterocomma pilosum
Tuberolachnus salignus
S. dasyclados = S. smithiana
S. decipiens Cavariella konoi
450 Salix
S. denticulata = S. elegans
S. dichroa Cavariella archangelicae
S. discolor Chaitophorus pallipes, pusillus
(American Pussy Willow) Phylloxerina salicicola
Plocamaphis flocculosa
Tuberolachnus salignus
S. dshugdshurica Chaitophorus saliapterus
Pterocomma rufipes
S. eleagnos = S. incana
S. elegans Cavariella aegopodii, biswasi,
nigrae, salicicola
Neoacyrthosiphon dubium
S. elegantissima Chaitophorus niger
Tuberolachnus salignus
S. eriocarpa Cavariella salicicola
Chaitophorus salijaponicus
S. exigua Chaitophorus macrostachyae,
(Sandbar Willow) viminalis
Pterocomma sanguiceps
S. fluviatilis Chaitophorus nigrae, populicola
(River Willow) Tuberolachnus salignus
S. fragilis Aphis farinosa
(Crack Willow) Cavariella aegopodii,
archangelicae, konoi,
pastinacae, quinquemaculalus,
theobaldi
Chaitophorus beulhani, niger,
truncatus, vitellinae
Lachnus longirostrum
Neopterocomma asiphum
Pemphigus saliciradicis
Phylloxerina salicis
Pterocomma chaetosiphon, konoi,
pilosum, salicis, smithiae
Tuberolachnus salignus
S. fulvopubescens Cavariella ?aegopodii, japonica
S. fuscescens Pterocomma rufipes
S. gilgiana Cavariella salicicola
S. glandulosa (var. glabra) Aphis farinosa
Cavariella salicicola
Salix 451
Chaitophorus saliciniger
Plocamaphis coreana
Tuberolachnus salignus
S. glauca Cavariella konoi
Chaitophorus lapponum
Pterocomma smithiae
S. graciliglans Chaitophorus saliciniger
Plocamaphis coreana
S. gracilistyla Aphis farinosa
Cavariella archangelicae,
nipponica
Chaitophorus niger, saliciniger
Pterocomma rufipes, [salijaponica]
Tuberolachnus salignus
S. hastata Aphis farinosa
S. herbacea Cavariella aegopodii
Pemphigus saliciradicis
S. humilis Phylloxerina salicicola
S. incana Chaitophorus salicti, similis
Phylloxerina daphnoides
Tuberolachnus salignus
S. integra Cavariella aegopodii, nipponica
Chaitophorus matsumurai,
saliapterus, salijaponicus
S. irrorata Cavariella archangelicae
Chaitophorus nigrae
S. kinuyanagi Cavariella nipponica
Chaitophorus horii, saliapterus
S. koreensis Cavariella salicicola
Tuberolachnus salignus
S. koriyanagi Aphis farinosa
Cavariella aegopodii, konoi,
nipponica
Chaitophorus saliapterus,
salijaponicus
Plocamaphis coreana
[Pterocomma salijaponica]
Tuberolachnus salignus
S. laevigata Cavariella pastinacae
Chaitophorus macrostachyae,
452 Salix
Chaitophorus beuthani,
diversisetosus ssp. austriacus,
mordvilkoi, niger, salicti,
truncatus, vitellinae
Plocamaphis amerinae
Pterocomma italica, konoi,
pilosum, salicis
Tuberolachnus salignus
S. pyrenaica Aphis farinosa
S. repens Aphis farinosa
Cavariella aegopodii
Chaitophorus hypogaeus, niger,
parvus, salicti
Neopterocomma verhoeveni
Plocamaphis flocculosa ssp.
goernitzi
Pterocomma jacksoni, konoi,
pilosum
S. reticulata [Pemphigus groenlandicus]
S. rorida Aphis farinosa ssp. yanagicola
Cavariella konoi, nipponica,
salicicola
Chaitophorus horii, saliapterus
S. russica Chaitophorus brunealineatus
S. sachalinensis Aspidophorodon salicis
Cavariella nipponica
Chaitophorus horii
Prociphilus oriens
S. salicifolium Pterocomma tuberculatum
S. schwerinii Cavariella nipponica
Chaitophorus horii, niger
Pterocomma konoi, rufipes
S. scouleriana Aphis farinosa
Macrosiphum californicum
Pterocomma salicis
S. sieboldiana Cavariella nipponica
Chaitophorus matsumurai
S. silesiaca Pachypappa warschavensis
S. sitchensis Aphis farinosa
(Sitka Willow) Pterocomma smithiae
Tuberolachnus salignus
Salix 455
Fig. 100. Head and prothorax, with detail of structure of wax pore plates, of aptera of A, Phylloxerina
salicis and B, Ph. capreae.
Fig. 101. A, SIPH of Aspidophorodon harvensis; front of head of B, A. salicis and C, A. harvensis.
hairs; if hairs are rather short and sparse mid-dorsally then the antennae,
legs and CAUDA are all densely hairy . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Body hairs sparse and usually rather short, at least dorsally . . . . . . 18
17 Small to medium-sized (BL 1.5-2.6 mm). SIPH stump-shaped, usually with
reticulate sculpturing (see Fig. 103). CAUDA either knobbed, rounded or
bluntly triangular, with 6-14 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus spp. - KEY A (p. 460)
- Medium to large (BL 2.2-4.5 mm). SIPH almost cylindrical or swollen,
usually without reticulate sculpturing (see Fig. 106). CAUDA either
rounded or bluntly triangular with about 20-60 hairs, or tongue-shaped
with a constriction and fewer hairs (in which case SIPH are dark and
strongly swollen and antennae are 4-segmented) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocommatini spp. - KEY B (p. 467)
18 ABD TERG 8 with a backwardly-directed supracaudal process bearing a
pair of hairs near apex (Fig. 107) . . . Cavariella spp. - KEY C (p. 472)
- ABD TERG 8 without a supracaudal process . . . . . . . . . . . . . . . . . . . 19
19 Antennal tubercles with forwardly-directed processes on either side of a
well-developed median tubercle, and ANT I also with projecting inner
face. Antenna 4- or 5-segmented. SIPH thin and flangeless, spoon-
shaped, obliquely truncated at apex (Fig. 101A) . . . . . . . . . . . . . . . . . 20
- Antennal tubercles developed or undeveloped, without processes. Antenna
usually 6-segmented. SIPH not spoon-shaped, with a flange . . . . . . . 21
20 Processes on antennal tubercles not much longer than their basal widths,
only slightly longer than median tubercle, which is bilobed (Fig. 101B).
Hairs on frontal processes, and on ABD TERG 8, blunt or pointed
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aspidophorodon salicis
- Processes on antennal tubercles more than 2 x longer than median
tubercle, which is not bilobed (Fig. 101C). Hairs on frontal processes and
ABD TERG 8 slightly capitate . . . . . . . . . . . Aspidophorodon harvensis
21 SIPH usually dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
460 Salix
- SIPH pale at least at base, if rather dark then with subapical polygonal
reticulation ..................................................23
22 ANT PT/BASE 3.5 or more. CAUDA dark with 10-26 hairs. Stridulatory
apparatus present (Fig. 122) . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
- ANT PT/BASE 2.0-3.2. CAUDA paler than SIPH, with 4-9 hairs. No
Stridulatory apparatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
23 Antennal tubercles undeveloped. ANT PT/BASE 1.5-2.3. CAUDA dark
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
- Antennal tubercles moderately to well developed, broadly divergent (e.g.
Fig. 42K). ANT PT/BASE 2.1-7.5. CAUDA pale . . . . . . . . . . . . . . . . 25
24 R IV+V 1.03-1.25 x HT II. ABD TERG 2-6 each with 2-3 marginal hairs
on each side. ABD TERG 8 with 2, or rarely 3-4, hairs . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis farinosa
- R IV+V 0.93-1.07 x HT II. ABD TERG 2-6 each with 5-11 marginal hairs
on each side. ABD TERG 8 with 4-7 hairs . . . . . . . . . Aphis roumanica
25 ANT PT/BASE 6.5-7.5. SIPH darkened distally with a subapical zone of
polygonal reticulation . . . . . . . . . . . . . . . . . . . Macrosiphum californicum
- ANT PT/BASE 2.1-4.5. SIPH wholly pale with no distinct zone of
polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 ANT III with 1-5 secondary rhinaria. ANT PT/BASE 3.5-4.5. SIPH
1.00-1.32 x CAUDA . . . . . . . . . . . . . . . . . Acyrthosiphon aurlandicum*
- ANT III without rhinaria. ANT PT/BASE 2.1-2.6. SIPH 2.5-3.5 x
CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
27 Antenna more than 0.75 x BE, with ANT III longer than head width
across (and including) eyes . . . . . . . . . . . . . . . Neoacyrthosiphon dubium
- Antenna less than 0.7 x BL, with ANT III about 0.5 x head width across
eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elatobium hidaense
Fig. 102. CAUDA of aptera of A, Chaitophorus saliciniger, B, Ch. viminalis, C, Ch. nigrae, D, Ch. vitellinae;
ANT III of aptera of E, Ch. crucis, F, Ch. saliciniger and G, Ch. capreae.
Fig. 103. Hairs on ABD TERG 4-6 of aptera of A, Chaitophorus capreae and B, Ch. horii; C, basal part
of hind tibia of apterous vivipara of Ch. saliniger; ABD TERG 5 and 6 of aptera of D, Ch. nigrae and
E, Ch. pentandrinus.
8 BL 1.8-2.1 x maximum body width. ABD TERG 1-5 each with only 0-12
smaller accessory hairs between the long spinal, pleural and marginal
pairs, and mostly forming a single row with them (Fig. 103A) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . capreae
- Body narrower; BL 2.1-2.5 x maximum body width. ABD TERG 1-5
each with 12-30 smaller accessory hairs, some in an additional row
anterior to the large primary hairs (Fig. 103B) . . . . . . . . . . . . . . . . horii
9 Abdominal tergum membranous and mainly pale except for dusky
scleroites at bases of spinal and pleural hairs, lateral sclerites and trans-
verse bands on ABD TERG 7 and 8. Dorsal body hairs very long and
pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hypogeus
- Abdominal tergum sclerotic, pale or pigmented, with usually at least ABD
TERG 2-6 fused together. Dorsal body hairs variable . . . . . . . . . . . 10
10 Hind tibia slightly swollen ventrally near base, with a group of 2-17
small circular ?scent plaques (Fig. 103C). Tergum dark, with nodulose
ornamentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . saliniger
Salix 463
- Hind tibia without a basal swollen part bearing ?scent plaques. Tergum
pale or dark, ornamentation variable . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 CAUDA bluntly conical, often with an indentation or constriction
partially delimiting a tongue-shaped or rounded apical part, but with the
width at the constriction not usually less than 0.8 x maximum diameter
of apical part (e.g. Figs 102C, D; examine several specimens) . . . . . 12
- CAUDA always with a distinct constriction or heck dividing it into a
triangular basal part and a globular apical knob, with the width at the
constriction less than 0.8 x maximal diameter of the knob (Fig. 102B)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
12 ANT BASE VI with 2-4 hairs, but usually 3 hairs (examine several
specimens). Tergum ornamented with separate small nodules. (R IV+V
1.05-1.20 x HT II; ANT PT/BASE 1.5-2.2; dorsal body hairs long,
100-160 µm; dorsum pale in spring and early summer, with dark pleural
bands in late summer) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vitellinae
- ANT BASE VI with 2 hairs, or rarely 3 hairs, and without the above com-
bination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Tergum pale. Dorsal body hairs mostly with furcate apices . . . . . . . 14
- Tergum usually dark, either t o t a l l y or with a paler spinal region. Dorsal
body hairs usually with pointed apices . . . . . . . . . . . . . . . . . . . . . . . . . 16
14 R IV+V less than 0.8 x HT I I . . . . . . . . . . . . . . . . . . . . . . . . . . . monelli
- R IV+V at least 0.9 x HT I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 ANT PT/BASE 1.0-1.3, R IV+V 1.1-1.3 x HT II . . . . shaposhnikovi
- ANT PT/BASE 2.1-2.5, R IV+V 0.9-1.1 x HT II . . . . . . . . furcatus
16 ABD TERG 1-6 all fused . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- ABD TERG 1 separate, 2-6 fused or separate . . . . . . . . . . . . . . . . . . 18
17 SIPH unpigmented, with a clear surrounding area. ABD TERG 1-8 all
with pale spinal areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pallipes
- SIPH dusky, without any clear surrounding area (Fig. 103D). ABD TERG
1-5 often with clear spinal areas in heavily pigmented specimens, but then
ABD TERG 7 and 8 are always evenly pigmented . . . . . . nigrae group
18 ABD TERG 2 separate from 3, and ABD TERG 3-6 often with separate
or only partially fused bands. R IV+V 1.0-1.3 x HT II . . . . . ramicola
- ABD TERG 2-6 usually wholly or partially fused; if separate in some
specimens then R IV+V 0.8-1.0 x HT II . . . . . . . . . . . . . . . . . . . . . . 19
19 R IV with 3-6 accessory hairs (most commonly 4). First tarsal segments
most commonly with 5 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . parvus
- R IV with 2 accessory hairs. First tarsal segments most commonly with
7 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . mordvilkoi
464 Salix
Fig. 104. SIPH and surrounding cuticle of aptera of A, Chaitophorus salijaponicus and B, Ch. niger.
Fig. 105. Empodial hairs of A, Plocamaphis flocculosa and B, Fullawaya saliciradicis; prothoracic
marginal tubercle (right side) of aptera of C, Fullawaya bulbosa (after Richards, 1966b), and
D, Pterocomma bicolor; E, ANT VI of Pterocomma chaetosiphon.
18 ANT II with 8-20 hairs. ANT III almost always with a few (1-13) second-
ary rhinaria. SIPH slightly to markedly swollen . . . . . . . . . . . . . . . . . 19
- ANT II with 3-6 hairs. ANT III never with secondary rhinaria. SIPH
swollen or cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
19 ANT PT/BASE 0.7-1.0; R IV+V 0.65-0.90 X HT II. SIPH strongly
swollen, to more than 1.5 x their basal width (Fig. 106J) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma salicis
- ANT PT/BASE 0.8-1.9 (only below 1.0 in spring); R IV+V 0.85-1.30 X
HT II. SIPH slightly to moderately swollen, up to 1.5 x basal width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 SIPH moderately swollen, with maximal swelling distal to midlength, and
with imbrication reduced on swollen part, the imbrication throughout
showing no trace of spinules (Fig. 106K). ANT PT/BASE 0.8-1.3 in
spring and 1.25-1.65 in summer/autumn . . . . . . Pterocomma jacksoni
- SIPH slightly swollen, with maximal swelling at or proximal to midlength,
and with rather evenly distributed spinulose imbrication (Fig. 106L). ANT
PT/BASE 1.2-1.8 in spring and 1.5-1.9 in summer/autumn . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma italica
21 SIPH short and stout, with maximal swelling near base (Fig. 84A). ANT
PT/BASE 1.0-1.2 . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma smithiae
- SIPH cylindrical or swollen distally. ANT PT/BASE 1.0-2.2 . . . . . 22
22 ANT BASE VI with 5-10 long hairs exceeding diameter of segment, plus
2-4 short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
- ANT BASE VI with only 1-3 long hairs, plus 2-4 short hairs . . . . 25
23 SIPH (Fig. 106M) a little shorter than HT II . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma sanguiceps
- SIPH (Fig. 106N) longer than HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Prothorax and ABD TERG 1-7 usually all with well-developed marginal
tubercles, rarely missing from some segments. ABD TERG 7 with 13-34
spinopleural hairs (i.e. not including marginal hairs near the marginal
tubercle and spiracle on each side). Alata with 22-38 secondary rhinaria
on ANT III, 0-3 on ANT IV . . . . . . . . . . Pterocomma pilosum konoi
- Prothorax and ABD TERG 1-7 without marginal tubercles, or rarely with
1-2 small ones. ABD TERG 7 with 10-20 spinopleural hairs. Alata with
33-44 secondary rhinaria on ANT III and 0-5 on IV . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma pilosum pilosum
25 Prothoracic and abdominal marginal tubercles small (not much larger
than a hair-base) or absent. ANT PT/BASE 1.4-1.8; SIPH 0.23-0.31 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma groenlandica
- Marginal tubercles present and well developed on prothorax and most of
ABD TERG 1-7; variably shaped but often conical and broad-based,
472 Salix
much larger than adjacent hair-bases (e.g. Fig. 105D). ANT PT/BASE
1.3-2.2; SIPH 0.22-0.45 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Longest hairs on ANT III 85-115 µm, 2.2-3.3 x basal diameter of seg-
ment. SIPH usually markedly swollen distally, up to 1.5-1.9 x their basal
diameter (Fig. 106O). Alata with 12-26 secondary rhinaria on ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma bicolor
- Longest hairs on ANT III 120-150 µm, 3-4 x basal diameter of segment.
SIPH varying in shape, almost cylindrical or swollen to not more than 1.5
x their basal diameter (e.g. Fig. 106P). Alata with usually 25-30 secon-
dary rhinaria on ANT III . . . . . . . . . . . . . . . . . . . . . Pterocomma rufipes
Fig. 107. Supracaudal process of aptera (from Salix) of A, Cavariella digitata, B, C. theobaldi,
C, C. araliae, D, C. japonica, E, C. konoi, F, C. borealis, G, C. rutila (paratype), H, C. aegopodii,
I, C. aspidaphoides.
Salix 473
0.1 mm
Fig. 108. SIPH (left side) of aptera (from Salix) of A, Cavariella digitata, B, C. theobaldi,
C, C. pastinacae, D, C. araliae, E, C. japonica, F, C. salicicola, G, C. konoi, H, C. aquatica, I, C. salicis,
J, C. aegopodii.
basad (Fig. 107A). SIPH strongly imbricated over entire length (Fig. 108A)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . digitata
- ANT PT/BASE more than 2.0. Process on ABD TERG 8 usually
quadrate, with 2 hairs at apical corners and none more basad (Fig. 107B).
SIPH more weakly imbricated, often almost smooth on basal half
(Fig.108B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . theobaldi
4 ANT PT/BASE more than 2.4. SIPH long, clavate, smooth or weakly
imbricated (Fig. 108C), process on ABD TERG 8 quadrate . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pastinacae
- ANT PT/BASE less than 2.4. SIPH various . . . . . . . . . . . . . . . . . . . . . 5
5 R IV+V more than 1.3 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- R IV+V less than 1.3 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
6 ANT PT/BASE less than 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . nipponica
474 Salix
Most Sambucus species are shrubs; nevertheless a full list of elder aphids and
their recorded hosts is given for the sake of completeness.
SAPINDUS Sapindaceae
SAPIUM Euphorbiaceae
SARCOCEPHALUS Rubiaceae
Compared with Bambusa and Arundinaria, the aphid fauna of Sasa seems
more limited. Use key to aphids on Arundinaria. Ceratovacuna japonica will
come out to C. sylvestris in that key, apart from the shorter frontal horns;
separation of Ceratovacuna spp. on bamboos is provided in the key to aphids
on Bambusa, couplets 27-29. Yezaphis sasicola Matsumura cannot be keyed
as no specimens are available (but see information from original description
summarized on p. 925).
SASSAFRAS Lauraceae
SCHEFFLERA Araliaceae
SCHIMA Theaceae
SCHOTIA Leguminosae
SCHOUTENIA Tiliaceae
SCLEROCARYA Anacardiaceae
SCOLOPIA Flacourtiaceae
SECURINEGA Euphorbiaceae
SEQUOIA Taxodiaceae
- ANT PT/BASE less than 0.5. SIPH as large dark hairy cones . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara cupressi
- ANT PT/BASE 3.7-4.4. SIPH tubular and swollen on distal half .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia morrisoni
SEQUOIADENDRON Taxodiaceae
SESBANIA Leguminosae
SHIIA = CASTANOPSIS
SINDORA Leguminosae
SINOJACKIA Styracaceae
SMODINGIUM Anacardiaceae
SOPHORA Leguminosae
4 Antennal tubercles with inner faces convergent, scabrous (Fig. 42I). SIPH
slightly swollen on distal half . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
Antennal tubercles divergent, smooth. SIPH tapering . . . . . . . . . . . . . 5
5 SIPH 0.33-0.50 x BL . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon gossypii
SIPH about 0.3 x BL or less . . . . . . . . . . . . . Acyrthosiphon sophorae*
6 Abdominal dorsum with an extensive black patch . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
Abdominal dorsum with separate irregular spinal and small marginal
sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis sophoricola*
Fig. 109. A, wax pore plate of Eriosoma sorbiradicis (aptera); section of dorsal cuticle (ABD TERG 3) of
B, Muscaphis drepanosiphoides (aptera), C, Nearctaphis yohoensis (aptera); D, SIPH of Dysaphis pavlovskyana.
4 BL more than 1.5mm. Total antennal length 0.29-0.46 mm. Wax pore
plates with central areas of irregular shape, often subdivided (Fig. 60C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma lanigerum
- BL less than 1.4 mm, with total antennal length only 0.12-0.18 mm. Wax
pore plates with always undivided, approximately circular, central areas
(Fig. 109A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma sorbiradicis
5 R IV+V (of fundatrix) more than 0.2mm long and bearing 7 or more
accessory hairs. BL of spring migrant alata more than 4mm . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
- R IV+V (of fundatrix) less than 0.2 mm long with 2-4 accessory hairs. BL
of spring migrant alata less than 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus corrugatans
6 CAUDA short; helmet-shaped, triangular or semicircular, shorter than or
about as long as its basal width in dorsal view . . . . . . . . . . . . . . . . . . . 7
- CAUDA tongue- or finger-shaped, clearly longer than its basal width
in dorsal view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
7 Dorsal cuticle of aptera (fundatrix) very rough with numerous small
denticulate projections, and with fine, long hairs (Fig. 109B). Alata with
transverse bars of dorsal abdominal pigment, often broken, never fused
between tergites . . . . . . . . . . . . . . . . . . . . . . Muscaphis drepanosiphoides
- Dorsal cuticle of aptera (fundatrix or subsequent generation) smooth
486 Sorbus
or weakly reticulate, with long or short hairs. Alata with a solid black
dorsal abdominal patch, or with extensive bars fused between some
tergites ...................................................... 8
8 SIPH more than 4 x CAUDA and bearing a number of long hairs (in both
fundatrix and spring migrant alata) . . . . . . . . . Sorbaphis chaetosiphon
- SIPH less than 4 x CAUDA and without hairs . . . . . . . . . . . . . . . . . . 9
9 SIPH with closely-spaced rows of imbrications bearing small nodules or
spinules (e.g. Fig. 42M) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- SIPH smooth or normally imbricated, the imbrications sometimes spinu-
lose but not in regular closely-spaced rows . . . . . . . . . . . . . . . . . . . . . 11
10 ABD TERG 2-4 each with incomplete double rows of 6-10 spinopleural
hairs 50-70 µm long with fine apices. Tergum rough, with numerous
spinules arranged on a distinct reticulation (Fig. 109C), and with an
extensive solid black abdominal patch . . . . . . . . Nearctaphis yohoensis
- ABD TERG 2-4 each with a single row of 4 spinopleural hairs that are
blunt or acuminate. Tergum with faint reticulation and few spinules, and
broad transverse sclerotic bars that are not or only partially fused between
tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis californica
11 SIPH smooth, less than 2 x width at base. Spiracular apertures rounded
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus helichrysi
- SIPH imbricated, more than 2 x width at base. Spiracular apertures small
and reniform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 SIPH pale, about 2 x CAUDA or less, and distinctly swollen on distal
half (Fig. 109D) . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis pavlovskyana
- SIPH pale or dark, more than 2 x CAUDA, cylindrical or tapering or
only very slightly swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 SIPH pale. Marginal tubercles present on ABD TERG 1-7 (sometimes
absent from 6). CAUDA with 6-12 hairs . . . . . . . . . . . . Dysaphis sorbi
- SIPH dark, at least on distal part. Marginal tubercles present on ABD
TERG 1-4 or 1-5, absent from 6 and 7. CAUDA with 5, or rarely 6,
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 Hairs on ANT III pointed, maximally 0.9-1.5 x basal diameter of
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis aucupariae
- Hairs on ANT III blunt, maximally 0.3-0.8 x basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Hairs on ABD TERG 1-3 consistently short and blunt, maximally
15-20 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis ariae
- Hairs on ABD TERG 1-3 of variable length, often short and blunt
but usually with at least some pointed hairs 20-60 µm long, especially
marginally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis plantaginea
Sorbus 487
23 SIPH slightly swollen on distal half (Fig. 121H). ANT PT/BASE less
than 5. (Alata with secondary rhinaria only on ANT III) . . . . . . . . . . .
................................................. Myzus persicae
- SIPH tapering or cylindrical on distal half. ANT PT/BASE more than
5. (Alata with secondary rhinaria distributed ANT III 60-83, IV 36-52,
V 13-22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus insitus
24 Body spindle-shaped. SIPH with a subapical zone of polygonal reticula-
tion (Fig. 121C). ANT III usually with several secondary rhinaria on basal
half, and with longest hairs more than half basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae/pyrifoliae
- Body oval. SIPH without any polygonal reticulation (Fig. 42Q). ANT
III with one small rhinarium near base, and with minute hairs of length
less than half basal diameter of segment . . . . . . . Fimbriaphis gentneri
SPATHODEA Bignoniaceae
SPONDIAS Anacardiaceae
STAPHYLEA Staphyleaceae
STEGANOTAENIA Umbelliferae
STEWARTIA = STUARTIA
STREBLUS Moraceae
STRYCHNOS Loganiaceae
STUARTIA Theaceae
STYRAX Styracaceae
The life cycles of most Hormaphidinae described from galls on Styrax are
not yet known, although some have recently been elucidated. Most species
were originally described in Astegopteryx, but as the secondary host forms
became known some have been transferred to other genera (e.g. Ceratovacuna
Styrax 491
Fig. 111. A, fore tarsus of Tuberaphis vandermeermohri (alata from gall); B, antenna of alata from gall
of Astegopteryx leeuweni; C, ANT V of alata from gall of Cerataphis fransseni; D, ANT V of alata from
gall of Ceratoglyphina styracicola.
- Front of head with less than 20 hairs, none of which are immediately
anteriodorsal to median ocellus. ANT V with 0-8 secondary rhinaria
............................................................. 9
9 ANT V with 0-4 secondary rhinaria, and with primary rhinarium extend-
ing across more than half width of segment (Fig. 111C) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerataphis fransseni
- ANT V with 4-8 secondary rhinaria, and with a very small primary
rhinarium extending across less than half width of segment (Fig. 111D)
....................................... Ceratoglyphina styracicola
10 ANT V usually without secondary rhinaria, rarely with 1-2 . . . . . . . . .
.......................................... Tuberaphis takenouchii
- ANT V with 3-16 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 ANT V with 10-16 secondary rhinaria. SIPH without any clearly asso-
ciated hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratovacuna nekoashi
- ANT V with 3-11 secondary rhinaria. SIPH with 2-7 hairs around it
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 ABD TERG 7-8 distinctly sclerotized, thickened and dusky. (Secondary
494 Swietenia
SYMPLOCOS Symplocaceae
SYRINGA Oleaceae
SYZYGIUM Myrtaceae
TABEBUIA Bignoniaceae
Fig. 112. End of abdomen of fourth instar nymph of A, Taiwanaphis pseudocaudata, B, T. atrovirens
and C, T. montanicola (all after Noordam and Hille Ris Lambers, 1985).
TABERNAEMONTANA Apocynaceae
TAMARINDUS Leguminosae
TAMARIX Tamaricaceae
TAXODIUM Taxodiaceae
TECLEA Rutaceae
TECOMA Bignoniaceae
TECTONA Verbenaceae
TERMINALIA Combretaceae
Fig. 114. A, CAUDA of Lizerius brasiliensis (aptera); B, examples of tapering and wart-like spinal
processes of Paoliella.
Tilia 503
TETRACLINIS Cupressaceae
THESPESIA Malvaceae
THUJA Cupressaceae
Key to aphids on Tilia (a revised and expanded version of the key in B & E,
1984)
1 SIPH absent or present only as inconspicuous pores (forming leaf-nest
galls) ........................................................ 2
- SIPH present either as short truncate cones, large pores on broad hairy
cones, or long tubes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Alata from gall with numerous elongate secondary rhinaria on ANT
III-V, not in a single row, protruding as sharp transverse ridges so that
the outline is serrate (Fig. 115A) . . . . . . . . . . . . Tiliphagus lycoposugus
- Alata from gall with secondary rhinaria on ANT III-V arranged in a row
and not protruding as sharp ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 BL of alata from gall more than 4 mm. Media of forewing unbranched.
Antenna with sparse hairs and at least 4 secondary rhinaria on ANT V
.............................................. Prociphilus oriens
- BL of alata from gall less than 3 mm. Media of forewing once-branched.
Antenna with numerous hairs and 0-3 secondary rhinaria on ANT V
(Fig. 115B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patchiella reaumuri
Tilia 505
Fig. 115. Antenna of spring migrant alata of A, Tiliphagus lycoposugus and B, Patchiella reaumuri.
4 BL more than 5 mm. Body and appendages covered in long hairs. SIPH
as large pores on broad hairy cones. (Alata with pterostigma extending
to tip of wing) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- BL less than 4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 SIPH tubular, much longer than their basal width. CAUDA finger-like
or triangular, anal plate entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- SIPH as truncate cones, hardly longer than their basal width. CAUDA
knobbed, anal plate bilobed (all viviparae alate) . . . . . . . . . . . . . . . . . . 8
6 SIPH black, markedly swollen over distal 0.7 of length . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphoninus tiliae
- SIPH pale or dark, not at all swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Aptera with SIPH mainly dark, about 3 x CAUDA, with a subapical
zone of polygonal reticulation; cuticle of head smooth; ANT III with 3-10
secondary rhinaria in a row . . . . . . . . . . . . . . . . . . . . Macrosiphum tiliae
- Aptera with SIPH pale except at apex, about 2 x CAUDA, without
polygonal reticulation; cuticle of head spiculose; ANT III with one small
secondary rhinarium near base . . . . . . . . . . . . . . . . . Aulacorthum solani
8 Secondary rhinaria on ANT III round to oval, with ciliated borders.
Forewing with Rs absent or indistinct and costal margin not heavily
pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Shivaphis tilisucta
- Secondary rhinaria on ANT III transversely elongate, not ciliated. Costal
margin of forewing heavily pigmented and Rs present . . . . . . . . . . . . . 9
9 Forewing with separate fuscous patches at ends of veins. Hind femur
black except at base. ABD TERG 1-7 with black marginal as well as
pleural pairs of sclerites, and dark SIPH . . . . . . . . . Eucallipterus tiliae
- Forewing with a zig-zag black band joining distal ends of media, Cu la
and Cu lb . Hind femur mainly pale. ABD TERG 1-7 with or without
dark markings. SIPH pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
506 Tocoyena
TOCOYENA Rubiaceae
TODDALIA Rutaceae
TOONA Meliaceae
TORRUBIA = PISONIA
Trochodendron 507
TOURNEFORTIA Boraginaceae
TREMA Ulmaceae
SIPH tubular with distal ring of about 4 hairs and 1-2 hairs on the
basal half. ABD TERG 7 with 4 finger-like processes. ANT PT/BASE
0.25-0.30 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sumatraphis celti
Without this combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
TRISTANIA Myrtaceae
TROCHODENDRON Trochodendraceae
TURPINIA Staphyleaceae
UAPACA Euphorbiaceae
In all, 68 aphid species are recorded from elms, which are primary hosts for
most of the Eriosomatini (44 species). For gall formers, Eriosomatini show
remarkably little evidence of host specificity, the recorded host range reflect-
ing geographical distribution rather than host affinities, with the more widely-
distributed species such as Eriosoma ulmi recorded from Ulmus species native
to different parts of the world. Elm-feeding aphids are therefore all included
in a single key as far as possible. However, at the present time it is not possible
to key together all species of the large genus Eriosoma and its close relatives
(Georgiaphis, Schizoneurella), so these are taken out of the main key and
included in separate keys under particular host species, within the host plant
list that follows the main key.
- ANT PT/BASE 0.6 or more. Tibiae thin and mainly pale with mostly
short hairs. Secondary rhinaria very narrow, slit-like. Spinal hairs on ABD
TERG 1 and 2 on prominent finger-like or conical tubercles (pleural hairs
absent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Media of forewing once-branched, with a patch of fuscous distally
between the branches, and another at the distal end of Cu lb (Fig. 116C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis (Sappocallis) ulmicola
- Forewing with media twice-branched, and with or without a different
pattern of markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Head with 3 pairs of dorsal tubercles, bearing apical hairs; the most
posterior pair longest, finger-like (Fig. 116A) . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis ulmiparvifoliae
- Head without paired dorsal tubercles or with only one very small pair
............................................................. 8
8 Forewing with media and CU1b thickly and rather evenly bordered with
fuscous basally as well as distally, and Cu la also often partially bordered.
Abdomen robust, with more-or-less extensive dark dorsal markings. SIPH
dark, with a dark basal sclerite . . . . . . . . . . . . . . . . . . Tinocallis platani
- Forewing with veins mostly not bordered or with fuscous restricted to
branches of media distally, or to spots or patches at distal ends of veins
............................................................. 9
9 Pronotum with 1 (posterior) or 2 pairs of finger-like spinal tubercles
(e.g. Fig. 116B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Pronotum without spinal tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
10 Head and pronotum wholly black (dark brown in mounted specimens),
and dorsal abdomen without dark markings . . . . . . . . . . . . . . . . . . . . 11
- Head and pronotum, or at least pronotum, pale or dusky or with restricted
dark markings. ABD TERG 3-5 (or 3-7) with dark sclerites at bases of
spinal hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Forewings with distal branches of media bordered with fuscous and with
more-or-less extensive fuscous patches at distal ends of Cula and Cu lb
(Fig. 51B). ANT III with 14-24 secondary rhinaria. R IV+V with 6-10
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis takachihoensis
- Forewings without fuscous markings (or, at least, none discernible in the
mounted specimens available). ANT III with 10-12 secondary rhinaria.
R IV+V with 4-6 accesory hairs . . . . . . . . . . . . Tinocallis sapporoensis
12 Distal branches of media of forewing bordered with fuscous, and a
patch of fuscous also on the hind margin at the end of Cu1b (Fig. 116D).
R IV+V 0.76-0.90 x HT II. ANT III with 6-17 secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis saltans
512 Ulmus
- Forewing veins not bordered with fuscous, and hind margin without a
patch of fuscous at end of Cu lb . R IV+V 0.90-1.03 x HT II. ANT III
with 11-28 secondary rhinaria . . . . . . . . . . . . . . . . . . . Tinocallis nevskyi
13 Head with a single narrow dark dorsal mesial stripe. ANT III with 14-22
secondary rhinaria. Spinal tubercles on ABD TERG 1 and 2 shorter than
their basal widths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis zelkowae
- Head with variable pigmentation; sometimes with paired longitudinal
stripes but never with a single narrow mesial stripe. ANT III with 5-12
secondary rhinaria. Spinal tubercles on ABD TERG 1 and 2 large and
prominent, usually larger than their basal widths, sometimes united at
bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis ulmifolii
14 Body of aptera broadly pear-shaped, broadest anteriorly, without distinct
head and prothorax. Oviparous. (Alata undescribed) . . . . . . . . . . . . . . .
............................................... Olegia ulmifoliae
- Body of aptera not broadest anteriorly, head and prothorax distinct
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Rostrum much longer than body. SIPH as large pores on broad pigmented
hairy cones. Abdomen with a mesial row of 6 dark patches ventrally
............................................. Stomaphis ulmicola
- Rostrum much shorter than body. SIPH variously shaped . . . . . . . . 16
Ulmus 513
16 SIPH tubular, much longer than basal width. ANT PT/BASE more than
1. Wax glands absent . . . . . . go to key to polyphagous aphids, p. 532
- SIPH absent, ring- or pore-like, or raised on small shallow cones. ANT
PT/BASE less than 0.6. Wax glands often present . . . . . . . . . . . . . . 17
17 Apterae in brown microrrhizal cysts on roots, with distal hairs on tibiae
thick and spine-like or dagger-like, and tarsi 1-segmented. SIPH absent
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mimeuria ulmiphila
- Apterae in galls or leaf nests, except for one species (Eriosoma rileyi)
forming free-living colonies on bark; tibial hairs normal, tarsi 1- or
2-segmented. SIPH present as pores, sometimes raised on small shallow
cones with surrounding hairs, or absent . . . . . . . . . . . . . . . . . . . . . . . . 18
18 (All remaining couplets refer only to alatae produced in or emerging from
galls or leaf nests.) BL more than 4mm. Hindwing with 2 oblique veins
arising very close together . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
- BL less than 4 mm. Hindwing if with 2 oblique veins then their bases are
separated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 Hindwing with 2 oblique veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
- Hindwing with 1 oblique vein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
20 SIPH present as fairly large, often raised pores on ABD TERG 5, with
partially sclerotized rims and surrounding hairs. R IV+V with 4 or more
accessory hairs. ANT III elongate (Fig. 118), usually more than 0.8 x
head width across (and including) eyes (except in Eriosoma mimicum)
............................... go to keys to species of Eriosoma,
Georgiaphis and Schizoneurella under individual Ulmus spp.
- SIPH absent or present as small or large pores on ABD TERG 6, with
wholly sclerotized rims and no clearly associated hairs. R IV+V with 2-3
accessory hairs. ANT III less than 0.8 x head width across eyes (e.g.
Figs 117A-F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 Dorsal abdomen with dark intersegmental transverse bands, broadest in
midline. ANT BASE VI much shorter than R IV+V. Secondary rhinaria
distributed ANT III 12-18, IV 2-5, V 2-4, VI 0-3 (Fig. 117A) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphidounguis mali
- Dorsal abdomen without transverse bands. ANT BASE VI much longer
than R IV+V. Secondary rhinaria distributed ANT III 9-24, IV 2-7,
V 4-12, VI 4-10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 Embryos without mouthparts (sexupara) . . . . Kaltenbachiella japonica
- Embryos with mouthparts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 SIPH absent. Secondary rhinaria distributed ANT III 17-25, IV 4-7,
V 5-8, VI 7-10 (Fig. 117B) . . . . . . . . . . . . . . . . . Kaltenbachiella pallida
514 Ulmus
Fig. 117. Antennae of alatae from galls of A, Aphidounguis mali, B, Kaltenbachiella pallida,
C, K. spinosa, D, Colopha compressa, E, Tetraneura ulmi, F, T. nigriabdominalis; G, SIPH of
T. yezoensis, lateral and dorsal views; H, hind claws of embryo of T. javensis (seen through cuticle of alata).
- SIPH usually present, but if sometimes absent then ANT III has fewer
than 17 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Secondary rhinaria distributed ANT III 17-25, IV 5-9, V 7-9, VI 6-10
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kaltenbachiella ulmifusa
- Secondary rhinaria distributed ANT III 9-17, IV 2-6, V 4-12, VI 4-10
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 Media of forewing unbranched. Secondary rhinaria distributed ANT III
11-17, IV 4-6, V 8-12, VI 6-10 (Fig. 117C) . . . . . . . . . . . . . . . . . . . . . . .
.......................................... Kaltenbachiella spinosa
- Media of forewing usually once-branched. Secondary rhinaria distributed
ANT III 9-12, IV 2-4, V 4-6, VI 4-6 . . . . . . Kaltenbachiella nirecola
Ulmus 515
Fig. 118. Antennae of alatae from galls of A, Georgiaphis gillettei, B, Schizoneurella indica, C, Eriosoma
americanum, D, E. grossulariae (ANT VI only), E, E. ulmi (ANT VI only), F, E. moriokense, G, E. harunire.
U. campestris = U. procera
U. canescens Aphis spiraecola
Toxoptera aurantii
U. carpinifolia Colopha compressa
(Smooth-Leaved Elm) Eriosoma anncharlotteae, flavum,
Ulmus 519
grossulariae, lanuginosum,
patchiae, pyricola, ulmi
Tetraneura caerulescens,
nigriabdominalis, ulmi
Tinocallis platani
(For separation of Eriosoma spp. use key under Ulmus procera, p. 532)
U. davidiana Chromocallis nirecola
Tinocallis ulmicola
Tetraneura nigriabdominalis
U. davidiana var. japonica = U. japonica
U. densa Aphis craccivora, [sogdiana
Nevsky, 1929b]
Eriosoma phaenax
Tetraneura ulmi
U. effusa Colopha compressa
(Russian White Elm) Eriosoma ulmi
Tetraneura ulmi
Tinocallis platani
U. foliacea = U. carpinifolia
U. fulva = U. rubra
U. glabra Aulacorthum solani
(Wych Elm) Colopha compressa
Eriosoma flavum, grossulariae,
lanuginosum, patchiae,
phaenax, ulmi, ?ulmosedens
Kaltenbachiella japonica, pallida
Tetraneura caerulescens, ulmi
Tinocallis nevskyi, platani,
takachihoensis
Toxoptera aurantii
(For separation of Eriosoma spp. use key under Ulmus procera, p. 532)
U. japonica Chromocallis nirecola
(Harunire, Japanese Elm) Colopha nirecola
Eriosoma alabastrum, auratum,
eligulatum, harunire, japonica,
longicornutum, mediocornutum,
moriokense, ulmi, yangi ssp.
parasiticum
Kaltenbachiella japonica, nirecola,
spinosa
Olegia ulmifoliae
Prociphilus kuwanai, oriens
Sinochaitophorus mordvilkoi
520 Ulmus
Stomaphis ulmicola
Tetraneura fusiformis,
nigriabdominalis, radicicola,
sorini, [triangula], ulmi,
yezoensis
Tinocallis saltans, takachihoensis,
ulmicola, zelkowae
U. montana = U. glabra
U. parvifolia Aphidounguis mali
(Akinire, Chinese Elm) Eriosoma yangi
Tetraneura nigriabdominalis,
sorini, yezoensis
Tinocallis ulmiparvifoliae,
zelkowae
U. pedunculata Colopha compressa
U. pinnatoramosa Eriosoma ulmi
Tetraneura ulmi
Tinocallis platani
U. procera Aphis craccivora, [sogdiana
(English Elm) Nevsky, 1929b]
Chromocallis nirecola
Colopha nirecola
Eriosoma anncharlotteae, flavum,
grossulariae, japonicum,
lanuginosum, patchiae,
phaenax, pyricola, ulmi,
ulmosedens
Kaltenbachiella japonica, pallida
Mimeuria ulmiphila
Tetraneura africana, caerulescens,
nigriabdominalis, ulmi,
yezoensis
Tinocallis nevskyi, platani,
saltans, ulmicola, ulmifolii,
ulmiparvifoliae
Key to alatae of Eriosoma spp. from galls on Ulmus procera (including also
species on U. carpinifolia and U. glabra)
1 Antenna with only 14-28 secondary rhinaria in total (distributed III
12-20, IV 2-5, V 0-4, VI 0) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Antenna with at least 29 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . 3
2 ANT III shorter than ANT IV+V+VI. Secondary rhinaria on III mostly
not extending more than half-way around segment . . . . . . . japonicum
ANT III longer than ANT IV+V+VI. Secondary rhinaria on III mostly
extending more than half-way around segment . . . . . . . . . . . . . . flavum
3 ANT V without secondary rhinaria. (Secondary rhinaria distributed III
26-46, IV 3-8, V 0, VI 0) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
ANT V with secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Ulmus 523
4 Total antennal length less than 1 mm, with ANT III less than 0.6 mm.
HT 0.15-0.18 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . anncharlotteae
Total antennal length more than 1 mm, with ANT III more than 0.6 mm.
HT 0.17-0.21 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 ANT PT long and thin; PT/BASE 0.46-0.67 (Fig. 118D). Many hairs on
ABD TERG 1-5 standing on small scleroites. HT II with 22-25 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . grossulariae
- ANT PT shorter and thicker; PT/BASE 0.29-0.49 (Fig. 118E). Few hairs
on ABD TERG 1-5 with basal scleroites. HT II with 15-22 hairs . . . .
.......................................................... ulmi
6 ANT V longer than IV. R IV+V 1.20-1.35 x HT II, and bearing 18-23
accessory hairs. Apices of tibiae and first tarsal segments strongly spiculose.
(Secondary rhinaria distributed III 18-35, IV 2-7, V 1-7, VI 0) .... 7
- ANT V shorter than or as short as IV. R IV+V 0.8-1.2 x HT II and
bearing 8-14 accessory hairs. Apices of tibiae and first tarsal segments
with only a few minute spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 Alatae are sexuparae, having embryos without mouthparts . . . . . . . . . .
.................................................... ulmosedens
Alatae have embryos with mouthparts (coiled stylets visible through
maternal cuticle) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . patchiae
8 Secondary rhinaria distributed ANT III 33-48, IV 10-14, V 10-12,
VI 2-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . phaenax
- Secondary rhinaria distributed ANT III 21-35, IV 7-11, V 5-16, VI 0-1
............................................................. 9
9 BL 2.1-3.1 mm. R IV+V 0.19-0.23 mm. CAUDA with only 2 or rarely
3 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lanuginosum
- BL 1.3-2.4 mm. R IV+V 0.10-0.19 mm. CAUDA with 3-10 hairs . . . .
....................................................... pyricola
U. propinqua = U. japonica
U. pubescens = U. fulva
U. pumila Chromocallis nirecola, [pumili
(Asiatic Dwarf Elm) Zhang, 1982 (in Zhang and
Zhong, 1982c)], [similinirecola
Zhang, 1982 (in Zhang and
Zhong, 1982e)],
Eriosoma dilanigiosum,
eligulatum, mediocornutum,
ulmipumilae
Sinochaitophorus maoi
Tetraneura asymmachia, chinensis,
nigriabdominalis
Tinocallis saltans
524 Ulmus
U. racemosa = U. thomasii
U. rubra Colopha graminis, ulmicola
(Slippery Elm) Eriosoma ?crataegi, mimicum,
rileyi
Georgiaphis gillettei
Kaltenbachiella ulmifusa
(For separation of Eriosoma and Georgiaphis spp. use key under Ulmus
americana, p. 517)
U. scabra = U. glabra
U. suberosa = U. procera
U. thomasii Colopha compressa, ulmicola
Eriosoma ulmi
Tinocallis platani
U. uyematsui Eriosoma lishanense
Kaltenbachiella glabra
U. wallichiana Eriosoma kashmiricum, phaenax
Indiochaitophorus furcatus
Tetraneura javensis
Tinocallis saltans
UMBELLULARIA Lauraceae
UVARIA Annonaceae
VEPRIS Rutaceae
VERNONIA Compositae
VIRGILIA Leguminosae
VITEX Verbenaceae
WIDDRINGTONIA Cupressaceae
XANTHOPHYLLUM Polygalaceae
XYLOPIA Annonaceae
ZANTHOXYLUM Rutaceae
ZELKOVA Ulmaceae
The small but probably ancient aphid fauna of Zelkova is poorly studied. The
gall makers span 4 or 5 genera of Eriosomatini, all distinct from those on
Ulmus ('Colopha` caucasica probably does not belong in that genus), and are
probably relicts of a much larger fauna in the past. Tinocallis on Zelkova
include some that also feed on Ulmus (nirecola, ulmiparvifoliae), plus several
others that are included in the key rather tentatively as they have not been
seen by the authors. There are certainly other Tinocallis still to be described
from Zelkova. Note that Tinocallis zelkovae and T. zelkowae are separate
species.
Fig. 119. A, head of alata of Tinocallis viridis; dorsal abdominal markings of B, T. zelkovae and
C, T. zelkowae.
- CAUDA rounded, anal plate entire. SIPH absent or if present then not
in form of short truncate cones without hairs. (Adult viviparae apterous
and alate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2 Head with all 4 pairs of dorsal cephalic hairs borne on conical tubercles
(Fig. 119A). (ANT III with 15-26 secondary rhinaria) . . . . . . . . . . . . . . .
................................................ Tinocallis viridis
- Head without dorsal tubercles, or with only one very small pair . . . . . 3
3 ANT III about as long as IV+V+VI together and bearing about 40
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis mushensis*
- ANT III 0.5-0.7 x ANT IV+V+VI together and bearing 4-25 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Forewing with media and Cu lb thickly bordered with fuscous, especially
distally, and Cula with a distal fuscous patch. Dorsal abdomen with
extensive dark markings (Fig. 119B). ANT III with 4-13 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis zelkovae
- Forewing veins not thickly bordered with fuscous; sometimes with small
spots at distal ends. Dorsal abdomen without extensive dark markings;
sometimes with spots at bases of hairs (e.g. Fig. 119C). ANT III with
10-25 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Thorax without spinal tubercles, and ABD TERG 1-2 with short conical
spinal tubercles, shorter than their basal widths (Fig. 119C). (Head with
a single, narrow medial longitudinal dark stripe; ANT III with 13-22
secondary rhinaria) . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis zelkowae
- Thorax with spinal tubercles, and spinal tubercles on ABD TERG 1-2
finger-like, longer than their basal widths . . . . . . . . . . . . . . . . . . . . . . . 6
6 Spinal hairs on ABD TERG 3-7 without pigmented bases. ANT III with
20-25 secondary rhinaria . . . . . . . . . . . . . . . . . . . Tinocallis suzhouensis*
- Bases of spinal hairs on ABD TERG 3-7 encircled by pigment. ANT III
with 10-18 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
530 Zelkova
7 BL less than 2 mm. ANT III about as long as ANT IV+V together and
bearing 10-18 secondary rhinaria extending over 0.7 or more of length
............................................ Tinocallis sophorae*
- BL more than 2 mm. ANT III much shorter than ANT IV+V together and
bearing 12-13 secondary rhinaria confined to basal half ............
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis allozelkowae*
8 BL more than 4 mm. Rostrum much longer than body. Abdomen with a
medial ventral row of 5 dark patches . . . . . . . . . . . . Stomaphis yanonis
- BL less than 3 mm. Rostrum much shorter than body. Abdomen without
ventral dark markings except posteriorly . . . . . . . . . . . . . . . . . . . . . . . . 9
9 (Remaining couplets refer to alatae emerging from leaf-roll or pouch
galls.) Media of forewing once- or twice-branched. Hindwing with 1 or
2 oblique veins. SIPH pores present . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Media of forewing unbranched. Hindwing always with only 1 oblique
vein. SIPH pores present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
10 ANT III about as long as IV+V+VI together and bearing 28-43 secondary
rhinaria often nearly encircling segment, and on thickened ridges
giving a serrate profile (Fig. 120A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- ANT III (of 6-segmented antenna) shorter or longer than IV+V+VI
together but bearing only 10-22 narrow secondary rhinaria, mostly not
encircling more than 0.7 of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Total length of antennal flagellum (III-VI) 1.2-1.4 mm. ANT PT
0.01-0.02 mm long, with quadrate apex (Fig. 120A). Secondary rhinaria
distributed ANT III 31-43, IV 9-14, V 10-16, VI 8-11 . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colophina clematis
- Total length of antennal flagellum 0.9-1.1 mm. ANT PT very short and
indistinct, less than 0.01 mm long, with rounded apex. Secondary rhinaria
distributed ANT III 28-37, IV 8-13, V 9-13, VI 5-8 . . . . . . . . . . . . . . .
................................................ Colophina arma
12 R IV+V 0.15-0.17 mm long, more than 1.3 x HT II. Secondary rhinaria
distributed ANT III 12-22, IV 2-4, V 2-4, VI 0-1 . . . . . . . . . . . . . . . . .
......................................... Hemipodaphis persimilis
- R IV+V 0.07-0.10 mm long, less than 0.8 x HT II. Secondary rhinaria
distributed ANT III 10-13, IV 2-3, V 2-3, VI 0 . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colopha caucasica
13 ANT VI BASE (or V in 5-segmented antennae) with 3-7 secondary
rhinaria. ANT PT very short and broad, about 0.01 mm or less, with
rounded apex (Fig. 120B). SIPH pores absent. Secondary rhinaria dis-
tributed ANT III 7-16, IV 4-6, V 3-8, VI 3-7 . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracolopha morrisoni
Ziziphus 531
Fig. 120. Antennae of alatae from galls of A, Colophina clematis, B, Paracolopha morrisoni,
C, Byrsocryptoides zelkovaecola, D, B. zelkovae.
ZIZIPHUS Rhamnaceae
1 ANT PT/BASE much less than 1. SIPH on flat dark hairy cones
(Fig. 121A). Very large aphid (BL more than 5 mm) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- ANT PT/BASE much more than 1, and usually more than 2. SIPH
tubular, of varying shape. BL less than 4 mm . . . . . . . . . . . . . . . . . . . . 2
2 SIPH pale at least on basal third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- SIPH wholly and uniformly dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
3 CAUDA long, dark and pointed, as long as or slightly longer than SIPH,
which are dark on about distal two thirds (Fig. 121B) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinomegoura citricola
Polyphagous Aphids 533
- CAUDA pale or dusky, much shorter than SIPH, which are pale or dark
only at apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 SIPH long, with a subapical zone of polygonal reticulation extending
over about distal 0.15 of length; CAUDA long and pale (Fig. 121C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
- SIPH without any subapical polygonal reticulation . . . . . . . . . . . . . . . 5
5 SIPH slightly to moderately swollen over distal 0.5-0.7 of length (Figs
121D-H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- SIPH tapering from base to apex, without any trace of swelling (Figs
121I-K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
6 SIPH markedly inflated over distal two thirds, the swollen part being
smooth and of maximum diameter more than 1.5 x the minimum dia-
meter of the stem; CAUDA triangular, usually with 5 hairs (Fig. 121D)
.................................... Rhopalosiphoninus staphyleae
- SIPH slightly to moderately inflated, the swollen part being weakly or
strongly imbricated and of maximum diameter less than 1.5 x minimum
diameter of stem; CAUDA triangular or finger-like with 6-8 hairs
(Figs121E-H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Head and ANT I and II dark, ANT III with 1-2 secondary rhinaria near
base. Legs mostly dark, and SIPH conspicuously dark-tipped (Fig. 121E)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorihum magnoliae
- Head and ANT I and II pale, ANT III without secondary rhinaria. Legs
mostly pale, SIPH not conspicuously dark-tipped . . . . . . . . . . . . . . . . 8
8 SIPH less than 0.8 x ANT III, and coarsely imbricated even on swollen
part; CAUDA triangular, about 1.5 x its basal width (Fig. 121F) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus (Sciamyzus) cymbalariae
- SIPH more than 0.9 x ANT III, and only weakly to moderately imbri-
cated. CAUDA finger-like, about 2 x its basal width . . . . . . . . . . . . . 9
9 Usually dark green in life. SIPH usually slightly dusky overall, with
maximum width of swollen part usually more than 0.11 x SIPH length
(Fig. 121G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus antirrhinii
- Usually pale green, yellow-green or straw-coloured in life (but darker
green in cold conditions). SIPH usually quite pale except at apices, with
maximum width of swollen part usually less than 0.11 x SIPH length
(Fig. 121H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
10 Dorsal abdomen with a large dark roughly horseshoe-shaped patch
(Fig. 121I) . . . . . . . . . . . . . . . . . Aulacorthum (Neomyzus) circumflexum
- Dorsal abdmen without a large horseshoe-shaped patch . . . . . . . . . . 11
11 Dorsal abdomen with an intersegmental pattern of dark ornamentation
(Fig. 121J). ANT III without any secondary rhinaria. ANT PT/BASE less
534 Polyphagous Aphids
Fig. 121. End of abdomen of apterous vivipara of A, Pterochloroides persicae, B, Sinomegoura citricola,
C, Macrosiphum euphorbiae, D, Rhopalosiphoninus staphyleae, E, Aulacorthum magnoliae, F, Sciamyzus
cymbalariae, 6, Myzus antirrhinii, H, M. persicae, I, Aulacorthum circumflexum, J, Myzus ornatus,
K, Aulacorthum solani, L, Uroleucon compositae, M, U. ambrosiae, N, Aphis craccivora, 0, Toxoptera
odinae, P, T. aurantii, Q, T. citricidus, R, Aphis fabae group, S, A. nerii, T, A. spiraecola, U, A. gossypii.
Polyphagous Aphids 535
than 2.5. SIPH tapering, with a shallow S-curve, and coarsely imbricated.
BL less than 2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus ornatus
- Dorsal abdomen without any dark markings. ANT III with a single
rhinarium near base. ANT PT/BASE more than 3.5. SIPH fairly straight,
weakly imbricated (Fig. 121K). BL usually more than 2mm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
12 SIPH with a distal zone of polygonal reticulation . . . . . . . . . . . . . . . 13
- SIPH without any polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . . 14
13 CAUDA black (Fig. 121L). Coxae dark. Tibiae mainly pale . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleucon compositae
- CAUDA pale (Fig. 121M). Coxae pale. Tibiae mainly dark . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleucon ambrosiae
14 Dorsal abdomen almost wholly covered by an extensive solid black
sclerite; CAUDA black and usually bearing 4-7 hairs (Fig. 121N) ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
- Dorsal abdomen without an extensive solid black sclerite. CAUDA if
black then usually with more hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Stridulatory mechanism present, consisting of a conspicuous pattern of
ridges on ventrolateral areas of abdominal sternites 5 and 6, and a row
of short peg-like hairs on the hind tibia (Fig. 122) . . . . . . . . . . . . . . . 16
- Stridulatory mechanism not present, although peg-like hairs are some-
times present along part of hind tibia . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
16 SIPH much shorter than (0.4-0.6 x) CAUDA (Fig. 121O). ANT PT/
BASE 2.5-3.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T o x o p t e r a odinae
- SIPH usually longer than (0.9-1.5 x) CAUDA (Figs 121P, Q). ANT
PT/BASE 3.5-5.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 Longest hairs on ANT III 12-27 µm long, 0.5-1.0 x basal diameter of
segment. Longest hairs on hind tibia up to 60 µm long, less than 0.6 x HT
II. CAUDA with 10-26 hairs (rarely more than 20). BL often less than
2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
- Longest hairs on ANT III 29-64 µm long, 1.5-2.0 x basal diameter
of segment. Longest hairs on hind tibia 80-110 µm, 0.7-1.0 x HT II.
CAUDA with 19-54 hairs (rarely less than 25). BL usually more than
2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera citricidus
18 Dorsum with variable sclerotic pigmentation, usually consisting of at least
cross-bands on pronotum, mesonotum and ABD TERG 7 and 8, and
small dark marginal sclerites (Fig. 121R); often also with small dark
sclerites scattered over other abdominal tergites. CAUDA with 12-19
hairs ....................................................... 19
536 Polyphagous Aphids
- Femoral hairs all rather short, not exceeding the diameter of the femur at
its base. CAUDA pale or dark, but usually clearly paler than siphunculi,
tongue-shaped without a constriction, and bearing 4-7 hairs (Fig. 121U)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
THE APHIDS
INTRODUCTION
In this part of the book we provide information on the aphids listed and keyed
in Section II, treating them in alphabetical order of genera, and of species
within genera. If a species listed in Section II is not found in the alphabetical
sequence then it is probably mentioned in the discussion of some other species;
the index to aphid species names (Section IV) will indicate where to look.
The classification of Aphidoidea follows that used in Blackman and
Eastop (1984), rather than that of Heie (1980) which elevates most of the
subfamilies of Aphididae to family level. Table 3 compares these two classi-
fications and shows the equivalent categories.
The information given comprises appearance in life, body length of one
or more morphs, host plants, distribution, life cycle including time of appear-
ance of sexual morphs where known, parasitoids, selected references to the
biology and economic importance, and chromosome number where known.
The range of body length (BL) gives an indication of size, but should be
treated warily for lesser-known species where it may be based on very few
specimens. References are selected mainly to provide a lead into the literature
on a particular species, so more recent publications are sometimes included
in preference to earlier more comprehensive or significant pieces of work.
Chromosome number is usually given as 2n (female), with a reference for
those chromosome numbers not included in published lists (e.g. Kuznetsova
and Shaposhnikov, 1973; Blackman, 1980); previously unpublished chromo-
some numbers are indicated by an asterisk*.
538
Acanthochermes 539
Table 3. Classification of Aphidoidea used in this book, compared with Heie (1980)
PHYLLOXEROIDEA APHIDOIDEA
Adelgidae Adelgidae
Phylloxeridae Phylloxeridae
APHIDOIDEA Aphididae
Pemphigidae Pemphiginae
Eriosomatinae Eriosomatini
Pemphiginae Pemphigini
Fordinae Fordini
Hormaphididae Hormaphidinae
Oregminae Cerataphidini
Hormaphidinae Hormaphidini
(Nipponaphidini) Nipponaphidini
Phloeomyzidae Phloeomyzinae
Thelaxidae Thelaxinae
Anoeciidae Anoeciinae
Mindaridae Mindarinae
Drepanosiphidae Drepanosiphinae
Drepanosiphinae Drepanosiphini
Phyllaphidinae Phyllaphidini
Chaitophorinae Chaitophorinae
Greenideidae Greenideinae
Greenideinae Greenideini
Cervaphidinae Cervaphidini
Aphididae Aphidinae
Pterocommatinae Pterocommatini
Aphidinae: Aphidini Aphidini
Aphidinae: Macrosiphini Macrosiphini
Lachnidae Lachninae
Lachninae Lachnini
Cinarinae Cinarini
Traminae Tramini
these tubercles, the spinal pairs being shorter than their basal width and the
lateral pairs being a little longer.
plant preferences and life cycle are discussed by Müller and El Tigani (1986).
2n = 6.
Acyrthosiphon pisum (Harris) Pea Aphid (see B & E, 1984, p. 212) Apterae
rather large (2.5-4.4 mm), pale green or pink, usually in colonies on young
growth or pods of Leguminosae; only tree recorded from is Robinia pseu-
doacacia. 2n = 8.
Fig. 123. Galls of Adelges on Picea. A, A. abietis on P. glauca (after Patch, 1910a); B, A. cooleyi on
P. sitchensis (after Carter, 1976); C, A. nordmannianae on P. orientalis; D, A. lands on P. sitchensis
(after Carter, 1976); E, A. pectinatae (after Cholodkovsky, 1907).
Adeiges 543
first instars have fewer wax pores on posterior abdominal segments than
A. viridana.
Aiceona is not as closely related to Anoecia as might be thought from its name
and placement in Anoeciinae. There are about 14 recognized species, all in
East and Southeast Asia, mostly associated with Lauraceae. Several species
are described from unidentified hosts. Unusually, oviparae as well as males
are alate, and they may be found in colonies at almost any time of year.
Siphunculi are well developed as hairy cones in all female morphs, but
strangely absent in males. Accounts of Aiceona are available for India (M.R.
Ghosh and Raychaudhuri, 1973), northeast India (Raychaudhuri et al.,
1980d) and Japan (Takahashi, 1960a). The genus is clearly in need of revision.
550 Aiceona
Aiceona pseudosugii David, Sekhon and Bindra Apterae are 2-3 mm, pale
yellow with dark hind tibiae, occurring in small colonies on undersides of
young leaves, with attendant ants. Alatae have unpigmented wings. The
only identified host is Machilus odoratissima (David et al., 1970). Aiceona
longisetosa M.R. Ghosh and Raychaudhuri, 1973 seems to be a synonym.
Recorded from Himachal Pradesh and West Bengal, India. Sexuales and
life cycle are unknown.
A little-known genus with four or five nominal species in eastern Asia. It has
aleyrodiform apterae with a complete crenulate margin of wax glands, as in
Cerataphis, but distinguished from that genus by absence of frontal horns and
a more evident division between prothorax and mesothorax. Life cycles are
unknown, but an undescribed species has been collected (by G. Zhang and
VFE) from boat-shaped galls on Sinojackia xylocarpa at Hangzhou, China.
Others are described from Compositae (three species), Bambusa and Salix.
Two of the species on Compositae in Japan are monoecious and anholocyclic
(Takahashi and Sorin, 1958).
India (Chakrabarti and Maity, 1982). The last rostral segment (R IV+V) is
shorter than in other Aleurodaphis (1.1-1.5 x HT II). Appearance in life is
unknown, presumably dark with marginal wax.
A genus for the one species, A. quaestionis (Börner), a rather large green aphid
rolling apple leaves in spring and migrating for the summer to Senecio doro-
nicum. In Central Europe, with a boreo-alpine distribution (Shaposhnikov,
1951; Hille Ris Lambers and Wildbolz, 1958; B & E, 1984, p. 215).
ALLOTHORACAPHIS Takahashi
Hormaphidinae: Nipponaphidini
The apterae of the only known species have long marginal prosomal hairs like
Thoracaphis, but lack siphuncular pores and marginal hairs on the reduced
abdominal tergites 2-7. They have a less sclerotic dorsum than Thoracaphis,
with an irregular mosaic-like ornamentation.
A genus for two oak-feeding species, narrow-bodied and with long siphunculi
like Eutrichosiphum, but differing in the presence of distinctly capitate or
spatulate hairs.
Anoecia 553
A genus with only one known species, probably of African origin (Hille Ris
Lambers, 1954a).
species utilize the native Cornus in North America, Europe and eastern Asia.
This is a difficult group in need of revision; in the BMNH collection there
are undescribed species from Cornus mas in Italy, and from Cornus spp. in
Korea. See also B & E, 1984, pp. 217-218.
Anoecia major Börner Not yet found on Cornus in nature, but Börner
(1950) transferred sexuparae to C. sanguinea and obtained two-three genera-
tions of fundatrigeniae. Occurs widely in Europe on roots of Phalaris arun-
dinacea, and also recorded from Brachypodium pinnatum. BL 2.3-3.2 mm.
See also Heie (1980). 2n = 8.
A genus for two Australian species living on Myrtaceae (one on Agonis and
the other on Leptospermum). The antennae of both apterae and alatae are
5-segmented, and abdominal tergites 7 and 8 each bear slender hair-bearing
processes. The siphunculi are cylindrical or tapering, with a ring of 3-8 hairs
on the distal half, and the CAUDA is broadly rounded.
Anomalaphis casimiri Carver Apterae are small (BL 1.3-1.7 mm) and
brown. Distinguished from the only other described species (A. comperei
Pergande) by the presence of polygonal reticulation on the siphunculi distal
to the ring of hairs, and by dorsal hairs of aptera mostly arising from
tubercles. Apterae and alatae occur together with intermediate forms on
Leptospermum sp. var. Red Damask in New South Wales, Australia (Carver,
1971). Appearance of colonies in life and biology are not recorded. (Ano-
malaphis comperei on Agonis has apterous oviparae and is presumably
monoecious and holocyclic.)
in Korea. The Ulmus galls consist of several rolled leaves on one shoot;
initially only one of these is occupied by the fundatrix, but later its progeny
migrate to fill the empty leaf-rolls (Akimoto, 1983). Alate emigrants, BL
1.5-1.7 mm, have dark brown head, thorax, legs and antennae, and dark
bands between the abdominal tergites. Apterae on roots of secondary hosts
are small white aphids secreting wax wool.
Of the more than 400 species of Aphis, only about 50 are recorded from trees.
Some of these are monoecious on particular tree species, some host-alternate
between trees and herbs, and some are polyphagous. For an introduction to
the genus see B & E, 1984, p. 220. There are now revisions of Aphis for Britain
(Stroyan, 1984) and Fennoscandia/Denmark (Heie, 1986), and Brown (1989)
has keyed the alatae of the northwestern European species. A key to Aphis
of Manitoba, Canada, is also available (Rojanavongse and Robinson, 1977).
Aphis caliginosa Hottes and Frison Small, brownish aphids with dark
legs and antennae, in ant-attended colonies on ends of branches of Cornus
racemosa in eastern North America (Illinois, Pennsylvania, North Carolina,
New Jersey). Life cycle unknown (Robinson and Chen, 1969).
Aphis celastrii Matsumura Apterae yellow or green with dark legs (but dark
greenish-brown according to original description), recorded from Celastrus
orbiculatus, Deutzia sp., Ilex serrata and Polygonum spp. in Japan (Higuchi
and Miyazaki, 1969). Closely related to A. spiraecola. Life cycle unstudied,
but oviparae have been collected on Deutzia (Takahashi, 1966). 2n = 8.
Aphis cornifoliae Fitch Apterae are small (1.0-1.4 mm), dark brown to dull
greenish-black, on both upper and undersides of leaves of Cornus spp., living
without host alternation; males are apterous (Palmer, 1952). Widely distri-
buted in USA and Canada. Robinson and Chen (1969) discussed differences
from A. helianthi.
Aphis craccivora Koch Apterae 1.4-2.0 mm, dark brown to shining black,
without any wax dusting when adult, on young growth and usually ant-
attended. Polyphagous, but with a marked preference for Leguminosae.
Cosmopolitan. (See B & E, 1984, p. 223.) 2n = 8.
Aphis cytisorum Hartig Apterae 1.5-2.8 mm, very dark green to black and
covered with grey wax powder, ant-attended on young shoots and petioles
and later on inflorescences of Laburnum anagyroides; also recorded from
Spartium junceum. Monoecious holocyclic, with alate males. Widely distri-
buted in Europe eastward to Bulgaria and Russia; introduced and widespread
in North America. See Stroyan (1984), Heie (1986). 2n = 8.
Aphis fabae Scopoli Apterae 1.5-3.1 mm, dull black, sometimes with white
wax markings, usually ant-attended. Curling leaves of Euonymus europaeus
(also Philadelphus coronarius and Viburnum opulus) in spring and migrating
560 Aphis
Aphis farinosa Gmelin Apterae 1.6-2.5 mm, dull green with CAUDA dis-
tinctly darker than the long pale siphunculi, in dense colonies on young shoots
of Salix spp. in spring and early summer, attended by ants. Monoecious, with
green oviparae and orange males appearing from July onwards, although
occasionally populations of viviparae persist until August or September
(Stroyan, 1984; Heie, 1986). Occurs throughout northern temperate parts
of the world, and in South America (Argentina). A nominal subspecies,
yamagicola Matsumura, occurs commonly on Salix spp. (daphoides, rorida)
in Japan and is distinguished by having long, fine hairs on the third antennal
segment (about twice the middle diameter of the segment); it produces sexuales
in the autumn (Takahashi, 1966). Populations on Salix gracilistyla in
Siberia have short antennal hairs like European farinosa, but a longer last
rostral segment; R IV+V 1.27-1.45 x HT II, as compared with 1.07-1.25 x
HT II (Holman, 1987). 2n = 6.
Aphis feminea Hottes Apterae red with black head, pro- and mesothorax,
antennae, legs, siphunculi and CAUDA, lightly dusted with white wax. It
forms dense colonies on young twigs and small branches of Prunus serotina in
northeastern USA (Hottes and Frison, 1931).
hybridization may tend to confuse secondary host associations and life cycle
categories, and there are also probably geographical differences. 2n = 8.
Aphis utilis Zhang is a closely-related species in China.
Aphis lantanae Koch (= setacea Hille Ris Lambers) Apterae about 1.6mm,
colour in life probably black, resembling A. fabae. In Western Europe,
normally monoecious on Viburnum lantana (see Stroyan, 1984). Specimens
collected on Sorbus aria in Bavaria, Germany, in 1933 were described as new
under the name Doralis setacea, but this name was synonymized with A.
lantanae by Eastop and Hille Ris Lambers (1976).
Aphis mammulata Gimingham and Hille Ris Lambers Apterae 1.3-1.9 mm,
greenish-grey, found in autumn in dense, ant-attended colonies on petioles of
berries and undersides of leaves of Rhamnus cathartica. Monoecious holo-
cyclic; the colonies include oviparae and orange-brown males. The alata is
described by Heie (1986). Appearance of the colonies earlier in the season is
unrecorded. Recorded from England, Sweden, Germany, Poland, Czechoslo-
vakia and Russia (as A. rhamnicola Mamontova).
Aphis nerii Boyer de Fonscolombe Apterae 1.5-2.6 mm, bright yellow with
black siphunculi and CAUDA; antennae and legs also predominantly dark.
Alatae have dark wing veins. Mainly colonizing Asclepiadaceae and Apo-
cynaceae, often in large, dense colonies on young stems, and occasionally
on plants (including trees) of other families (e.g. Catalpa, Citrus). Widely
distributed throughout the tropics including many Pacific islands. Apparently
mainly anholocyclic, but sexual morphs were recently described from Japan
(Takada and Miyazaki, 1992). (See also B & E, 1984, p. 230; Stroyan, 1984.)
2n = 8.
Aphis pomi De Geer Apterae 1.3-2.2 mm, bright apple-green with black
siphunculi and dark CAUDA, in dense colonies on young growth of several
genera of woody Rosaceae (Pomoidea); ant-attended, and causing slight leaf
curl (B & E, 1984, p. 231). Later generations occur on the undersides of leaves
and are yellow-green. Monoecious holocyclic, with apterous males. In Europe,
southwest Asia and North America (where there is some confusion in the
literature with A. spiraecola on the same hosts). Alatae have darker wing veins
than A. spiraecola; Halbert and Voegtlin (1992) gave some other discriminants
based on North American populations of the two species. Hogmire et al.
(1992) compared population development and insecticidal susceptibility of
A. pomi and A. spiraecola. Oriental records of A. pomi all seem to apply
to other species, although it is surprising that it has not spread worldwide. (See
also Heie, 1986.) 2n = 8.
Aphis rhoicola Hille Ris Lambers Apterae 1.7-2.1mm, brown with pale
antennae and legs (except black tarsi) and black siphunculi. On leaves and
branches of Rhus abyssinica in Eritrea (Hille Ris Lambers, 1954a). Resembling
A. gossypii, but with a consistently shorter antennal terminal process. Life
cycle is unknown; however, very similar aphids occur on Rumex in India
(BMNH colln), so this is a possible secondary host.
Aphis salicariae Koch (= corniella Hille Ris Lambers) Apterae 1.6-2.4 mm,
reddish-brown with a light dusting of wax; legs pale (except tarsi), siphunculi
Aphis 565
and CAUDA black. Causing leaf curl to Cornus spp. (alba, sanguinea,
stolonifera) in spring and also feeding in developing flower umbels; host-
alternating to Epilobium angustifolium. Often ant-attended. Widely distri-
buted in Europe (except Spain and Portugal), eastward to Central Asia; and
introduced into and widespread in North America. 2n = 8.
Aphis sambuci Linnaeus Apterae 1.9-3.5 mm, dark green or dark brown to
yellowish-brown, in dense colonies around young stems of Sambucus spp. in
spring. Host alternation occurs to roots of various plants, especially Caryo-
phyllaceae and Rumex (Stroyan, 1984; Heie, 1986). Colonies may persist
through summer on Sambucus and may produce oviparae, but males only
seem to be produced on secondary hosts. In mild winters in England apterae
may hibernate on subterranean parts of Sambucus. Occurs throughout Europe
and temperate Asia (except that in Japan it seems to be replaced by a long-
haired relative, A. horii) and North and South America. It is often recorded
in North American literature as sambucifoliae Fitch. 2n = 8.
Aphis sophoricola Zhang Apterae blackish with whitish wax bloom, col-
lected on Sophora japonica at Beijing, China (Zhang and Zhong, 1981a).
Closely related to, if not synonymous with, A. cytisorum.
Aphis spiraecola Patch (= citricola van der Goot of recent authors) Apterae
1.2-2.2 mm, bright greenish-yellow to apple green, with brownish head and
blackish siphunculi and CAUDA; in colonies often distorting growing points,
usually ant-attended. Very polyphagous, particularly on plants of shrubby
habit, but also often on trees. Anholocyclic through most of the world, but
a holocycle involving Spiraea spp. as primary hosts is recorded from North
America, Brazil and Japan; in the latter country, Citrus is also a primary host
(B & E, 1984, p. 222, as A. citricola). 2n = 8.
Aphis verbasci Schrank Apterae 1.7-2.5 mm, bright golden yellow to pale
green with black siphunculi, ant-attended, on undersides of leaves of Buddleja
spp. (farreri, japonica, madagascariensis, paniculata), Verbascum spp., and
also recorded from Scrophularia. Monoecious holocyclic on these hosts;
sexuales are undescribed, but there are oviparae and apterous males in the
BMNH collection. In Europe (except Fennoscandia), Middle East, North
Africa, India and Pakistan. 2n = 8.
Aphis versicolor (Börner) Apterae 1.1-1.6 mm, body and siphunculi dark
green, on leaves of Rhamnus cathartica in Austria and Bulgaria (Börner,
1950). This species has similar very large marginal tubercles to A. mammulata
on this host from other parts of Europe, but much shorter hairs on antennae
and marginal areas of abdomen. Possibly it should be regarded as a geo-
graphical variant of that species. Monoecious holocyclic; males are apterous
and orange-brown, as in mammulata.
Aphis viticis Ferrari Apterae 0.8-1.7 mm, grass green with brown siphunculi
in spring, pale yellow in midsummer. Monecious holocyclic on Vitex agnus-
castus, with alate males; recorded from France, Iran, Israel, Italy, Mallorca,
Spain and Turkey. Barbagallo and Stroyan (1982) redescribed the apterous
and alate viviparae, and Nieto Nafria et al. (1986) described the sexuales.
whereas in Slavum the wings are held roof-like. The life cycle of only one
species is known and that has a specific primary host association with Pistacia
lentiscus.
A genus with one North American species related to Pterocallis, but with two
pairs of anterior prothoracic marginal hairs and a small hair arising from the
siphunculus, on the ventral side near its base (Richards, 1965).
A genus for one species in Italy related to Myzocallis, but with one pair of
spinal hairs on each abdominal tergite in all morphs and with other differences
in larval chaetotaxy.
Apulicallis trojanae Barbagallo and Patti Alatae lemon yellow with head
and thorax very slightly brownish; BL 1.3-2.3 mm. Holocyclic and mono-
568 Asiphonaphis
A small North American genus with only two currently recognized species,
related to Aphis but completely lacking siphunculi. Both alatae and apterae
have prominent marginal tubercles on all abdominal segments.
Asiphonaphis pruni Wilson and Davis Apterae are pale yellow to whitish-
green, sometimes banded with darker green, with darker head and pronotum,
in colonies on leaves and distorting the new growth of Prunus spp. (armeniaca,
serotina, virginiana) in Canada and USA. Monoecious holocyclic, with
apterous males. Robinson (1964) gives an account of this species on P. virgi-
niana in Manitoba.
The two species of this genus form large, cockscomb-like galls on Pistacia and
migrate to roots of grasses, especially Cynodon dactylon, with a holocycle
taking two years. Accounts are given by Remaudière and Tao (1957) and
A.K. Ghosh (1984b).
A small genus associated with Salix in East Asia. The antennae are 4- or
5-segmented with terminal process 0.6-1.0 times base of last segment and
the siphimculi have clavate ends with the aperture small and subapical.
The head of the aptera bears anteriorly-directed medial and lateral frontal
projections.
Astegopteryx minuta (van der Goot) The name most commonly applied to
populations of bambusae-like aphids with marginal wax pore plates reduced
(so that they are small, circular and discontiguous) or totally absent. In life
such apterae are generally yellowish-white to pale green with two irregular
dark green longitudinal markings and scanty wax; BL 1.1-1.8 mm (Noordam,
1991). On undersides of leaves of bamboos (Bambusa, Dendrocalamus).
Populations in which the apterae have no visible marginal wax pore plates and
are not clearly distinguishable from A. minuta have been described from India
as new species: A. neelagiriensis David, 1959a, on Bambusa sp. in Madras;
and Pseudastegopteryx himalayensis M.R. Ghosh, Pal and Raychaudhuri,
1977b, on unidentified bamboo in West Bengal. Alatae occur (in Java) from
May to mid-October (Noordam, 1991). 2n = 12 (in India; Kar et al., 1990).
Astegopteryx roepkei Hille Ris Lambers The gall of this species, found on
Styrax paralleloneurus in Sumatra, is remarkable; it is composed of thin,
spirally-twisted tubular subgalls, 15-20 cm long, their bases radiating from a
circular attachment site on one side of the stem near an axil-bud (Fig. 124E).
Alatae, BL about 1.8 mm, are adult in September. Secondary host is unknown.
A genus with nominally two species. The type species is described from
Lonicera, and has first tarsal segments with 3 hairs, elongate siphunculi and
alatae without rhinaria on antennal segment IV. The Indian Prunus-feeding
species placed here has 5 hairs on first tarsal segments, rather short siphunculi
and numerous rhinaria on antennal segment IV of alatae, suggesting that it
may not belong in this genus.
A genus for one or two species related to Aploneura and Slavum but with more
normal distribution of rhinaria on the antennae of the alatae, which fold their
wings roof-like in repose. The large characteristic horn-like galls (Fig. 126C)
can be formed on several species of Pistacia.
Six species are known, all from East and Southeast Asia. All viviparae are
alate, active insects (behaving like Euceraphis), and have a characteristic
transverse dark brown to black bar across the ventral side of the head between
the eyes. Most of the species are recorded from Betulaceae, which are also the
hosts of several related genera. The Indian species are reviewed by Chakrabarti
(1988) and A.K. Ghosh and Quednau (1990), and the Japanese species by
Higuchi (1972).
Betacallis alnicolens Matsumura Alatae are pale green with dark tibiae,
large (about 3.5mm), very active, on leaves of Alnus spp. (hirsuta, japonica,
matsumurae) in Japan, Korea and China, and also collected from Corylus
mandshurica in Korea (BMNH colln). 2n = 22.
Betacallis odaiensis Takahashi Alatae are pale lemon yellow to apricot, with
black basal, sensoriated, part of third antennal segment, black tibiae, two
black bars on the dorsal abdomen, and black siphunculi; BL 2.4-3.1 mm. On
leaves of Betula spp. (ermannii, maximowicziana, platyphylla var. japonica)
in Japan. (Indian records of odaiensis refer to other species.) 2n = 22.
Betacallis prunicola R.C. Basu, A.K. Ghosh and Raychaudhuri Alatae light
yellowish-green with dark tibiae, dark transverse bars on abdominal tergites
4 and 5 and dark siphunculi; BL 1.9-2.2 mm (A.K. Ghosh and Quednau,
1990). Described from adults and immatures on Prunus cerasus in Meghalaya,
India (R.C. Basu et al., 1975), but this is unlikely to be the true host. This
species is also recorded from Arunachal Pradesh, Manipur and West Bengal,
and from China and the Philippines. It can be distinguished from other
Betacallis species by the long ultimate rostral segment (R IV+V 1.5-1.7 x
HT II).
colln) indicate that the normal host association is probably with Betula.
Parasitized by Trioxys (Betuloxys) assamensis in India (D. Raychaudhuri,
1990).
Betulaphis pelei Hille Ris Lambers Apterae are yellow with dark tarsi
and tips of antennae, BL 1.4-1.7 mm. Alatae sometimes have a dark dorsal
abdominal patch. On Betula nana with a boreo-alpine distribution in northern
Europe, Iceland and Greenland (Heie, 1982). Aphids occurring on Betula
glandulosa, described from Baffin Island as Betulaphis arctosetis Richards,
1961, are probably also this species. 2n = 20.
Boernerina variabilis Richards Apterae are pale to bright yellow with black
on distal parts of antennae and legs, flattened, elongate oval, BL 1.9-2.6 mm.
Alatae vary in pigmentation; when well pigmented they have broad dark bars
dorsally and ventrally on the abdomen. On undersides of leaves of Alnus spp.
(sinuata, rugosa, alnobetula) in east and west Canada (British Columbia,
Northwest Territory, Ontario, Quebec, Labrador). Oviparae and small,
apterous males occur in August in Northwest Territory (BMNH colln, leg. R.
O'Doherty), but oviparae were collected in October in northern Ontario
(BMNH, leg. J. Sÿpkens). 2n = 16*.
Brachycaudus 583
One species. Brachysiphoniella montana (van der Goot), with short siphunculi
and a long CAUDA, mostly known from wax-covered colonies on aquatic
Gramineae (Takahashi, 1923) in East and Southeast Asia, and Australia. In
Korea there appears to be host alternation, with Pyrus montana as the primary
host, spring colonies causing leaf curl (Paik, 1972); as Semiaphis montana).
However, this needs confirmation, as it is possible that Paik's account con-
fuses B. montana with Melanaphis siphonella.
Related to Aphis, but with antennal processus terminalis clearly shorter than
base of last segment. The siphunculi are pale and much shorter than the
CAUDA, and the body is usually covered with wax meal or dust. There are
about 25 nominal species, mostly associated with xerophytes in Central and
586 Brachyunguis
Brachyunguis calligoni (Nevsky) Apterae are light green covered with wax
meal, BL 1.3-1.7 mm, with very small inconspicuous siphunculi, living on
young shoots of Calligonum spp. (eriopodium, setosum, ?comosum) in
Uzbekistan (Nevsky, 1928b). Biology is unknown.
Three species, two described from galls on the primary host (Zelkova) and
one only known from Cyperaceae. Alate spring migrants, emerging from
pseudogalls on Zelkova, have an unbranched media, only one oblique vein in
the hind wing and inconspicuous ring-like siphunculi. Resembling Kalten-
bachiella in many characters, but in Byrsocryptoides the secondary rhinaria
are less annular and the processus terminalis is relatively longer. Dzhibladze
588 Calaphis
Delicately built aphids with rather long thin legs, generally pale-coloured but
sometimes with distinctive markings and/or dark wing veins. Other characters
of the genus are the well-developed antennal tubercles and the absence or
indistinctness of the radial sector in the forewing. Some species have all
viviparae alate, others have both apterous and alate viviparae. As treated here,
Calaphis is holarctic and includes Kallistaphis and Cepegillettea. There are
about 15 species, 11 of which are in North America, two are European and
one is East Asian. All except one (myricae) live on Betulaceae. Parasitized by
Aphidius aquilus, which also attacks other related Betula-feeding aphids
(Stary, 1973).
Calaphis alni Baker Apterae and alatae of BL about 2.3-2.4 mm, yellowish
with distinctive black markings; alatae usually have a large black dorsal
abdominal patch and apterae have two bands across the thorax and one across
the posterior abdomen. Feeding mainly on the stems and foliage of tender
growing terminals of Alnus rugosa (Pepper, 1950), in eastern USA and
Canada (New Brunswick). The sexuales are undescribed.
with dark antennae and legs. Sexuales occur in October; males are alate
(Pepper, 1950).
Calaphis betulella Walsh, Plate 9e Viviparae all alate, yellowish with orange
and black markings, black stripes along the femora and wholly black tibiae
and tarsi; BL 2.0-2.5 mm. Feeds only on Betula nigra, on undersides of leaves.
Apterous oviparae and alate males occur in September-October. In eastern
USA. 2n = 18.
Calaphis (Cepegillettea) myricae (Patch) Apterae and alatae are both green
with dusky or dark distal antennal segments, tibiae and tarsi and siphunculi
either wholly or apically black; BL about 2.5 mm. On leaves of Myricacae
(Myrica asplenifoliae, M. gale, Comptonia peregrina) in northeastern USA
and eastern Canada. Apterous males and oviparae occur in September
(BMNH colln.; leg. E. MacGillivray).
Calaphis similis Quednau All viviparae alate, yellow with reddish head
and two red spots on abdomen, with antennal flagellum, tibiae and tarsi
mainly black, siphunculi pale, wing veins bordered with black; BL 3.0-
3.5 mm. On undersides of leaves of Betula spp. (costata, davurica, mand-
shurica, platyphylla var. japonica) in China, Korea, Siberia and Japan
(Pashchenko, 1988b; Quednau, 1979). Oviparae and males were collected in
October (BMNH colln, and Paik, 1972; as Kallistaphis sp.). Higuchi's (1972)
account of C. betulaecolens applies to this species,
About 17 species typically associated with Rhus, or with host alternation bet-
ween Rhus and Cyperaceae. Remaudière and Muñoz Viveros (1992) recently
expanded the concept of the genus to include Glabromyzus (four North
American spp.) and Juncomyzus (five East Asian spp.). Carolinaia sensu
stricto and all except one species of Glabromyzus are associated only with
Rhus subgenus Toxicodendron, so they do not concern us here.
Three species, all recorded only from Castanea bungeana in China. All
viviparae are alate. The genus resembles Myzocallis (Nippocallis) in wing
pigmentation, presence of spinal and marginal tubercles and a rather short
processus terminalis, but the species of Castanocallis are much more robust.
See Zhang and Zhong (1981c) for further information. Castanocallis
castanocallis has 2n = 14*.
Cavariella intermedia Hille Ris Lambers Apterae are evenly light green,
rather broadly spindle-shaped, BL 1.4-2.3 mm. Alatae differ from aegopodii
by having only a rather indistinct smoky dorsal abdominal patch due to pale,
partly coalescent sclerotic bars on ABD TERG 3-6 (becoming somewhat
darker in alatae developing at lower temperatures). Alatae also differ from
aegopodii by having 2-7 secondary rhinaria on the fourth antennal segment.
Recorded from Salix aurita, S. arbuscula and S. phylicifolia in Belgium,
Netherlands, Switzerland and UK. Life cycle is not completely known; Hille
Ris Lambers (1970a) found apterae producing oviparae and alatoid nymphs
(presumed males) on Salix in September, which seems to rule out host alterna-
tion. 2n = 6.
Cavariella nigra Basu Apterae in spring colonies on Salix are dusky black
including antennae, legs, siphunculi and CAUDA, pear-shaped, somewhat
flattened; BL 1.9-2.5 mm. Alatae have a dark brown central dorsal abdominal
patch. The black coloration of the aptera is unusual for a Cavariella; the dark
form of japonica reported from Taiwan is possibly this species, although
japonica from Japan has a longer last rostral segment (R IV+V 1.4-1.6 x HT
II, as opposed to R IV+V 1.0-1.2 x HT II in nigra). Cavariella nigra is
Cavariella 599
also unusual in having 0-1 secondary rhinaria near the middle of ANT V of
the aptera (none on III or IV). On undersides of leaves of Salix elegans in West
Bengal, India (A.N. Basu, 1964). The life cycle is unknown.
Cavariella takahashii Hille Ris Lambers Only the alata of this species is
described, so it could not be included in the key to Cavariella on Salix. BL
of alata about 2.2-2.3 mm; dorsal abdominal markings are indistinct and the
femora have numerous long wavy hairs as well as a number of much shorter
hairs on distal half similar to those on tibiae. The antennae have a short pro-
cessus terminalis and numerous secondary rhinaria; 51-52 on ANT III, 10-11
on IV and 3-5 on V. On Salix spp. in April in Japan (Hille Ris Lambers,
1965). Appearance in life and biology are unknown.
About six Southeast Asian species, some of which may host-alternate between
galls on Styrax and monocotyledonous secondary hosts (Araceae, bamboos,
Ceratoglyphina 601
Aphids similar to Astegopteryx but distinguishable from that genus and most
other Hormaphidinae by the rounded CAUDA and anal plate. The primary
host has recently been shown to be Styrax as in other Cerataphidini, but many
populations are probably anholocyclic on bamboos. Noordam (1991) reviewed
the species in Java.
602 Ceratovacuna
Ceratoglyphina bengalensis L.K. Ghosh Apterae are pale brown with two
rather indistinct longitudinal green stripes and a narrow, flat, wax fringe;
BL 2.1-2.5 mm. On undetermined bamboos in India (West Bengal, Assam;
M.R. Ghosh et al., 1977b) and also recorded from Java, where the aphids were
found living on the border of the sheath and at the leaf-bases of young shoots
(Noordam, 1991).
A genus in East and Southeast Asia, closely related to Astegopteryx and with
a similar alternation between Styrax and Gramineae, although this has only
been confirmed for one species. An indefinite number of species form dense
colonies on the undersides of bamboo leaves. The apterae are small to
medium-sized, yellow to brown in colour, and secrete much wax. Alatae of
the bamboo feeders are rare or unknown. As in the case of Astegopteryx,
characters used to separate species are frequently those known to vary accord-
ing to the environment, such as the degree of development of the wax glands,
and the genus is in need of further taxonomie revision. Accounts are available
Ceratovacuna 603
for Japan (Takahashi, 1958c), Java (Noordam, 1991), Korea (Paik, 1965),
India (M.R. Ghosh et al., 1977b) and Taiwan (Liao, 1976).
and less sclerotization of ABD TERG 6 (Takahashi, 1958c). They deposit first
instar sexuales on the leaves; these move to the branches without moulting and
develop to maturity without feeding, the eggs being laid on bark (Kurosu and
Aoki, 1990c). In Japan, Taiwan, Korea and Uttar Pradesh, India (Chakrabarti
and Maity, 1982). 2n = 12.
A genus with five species in India and Southeast Asia, in which the apterae
have remarkable branched marginal hair-bearing processes. In alatae these
processes are reduced to low, flat hair-bearing tubercles, except on the head.
The siphunculi are long, cylindrical, slightly curved outwards and a little
swollen subapically, where there is a ring of small hairs. For revisions of
Cervaphis see Hille Ris Lambers (1956a) and A.K. Ghosh (1982a).
A genus of three species, two of which are only known from grass roots in
India. They are related to Geoica but the emigrant alatae have ciliated secon-
dary rhinaria and their embryos have fewer dorsal hairs.
Chaitophorus gomesi Ilharco Apterae pale green mottled with dark green;
BL 1.1-1.4 mm. Alatae are undescribed. In large colonies on undersides
of leaves of Salix vitellina, attended by ants. Only known from Portugal.
Apterous males and oviparae were collected in November (Ilharco, 1968).
although Higuchi (1972) may have been correct in synonymizing it with horii,
as the differences in form of the dorsal hairs, etc. could be ascribed to
geographical variation. Apterous males and oviparae of the European form
occur in October. Colonies are not ant-attended (see also Heie, 1982).
Chaitophorus hypogaeus Hille Ris Lambers Apterae broadly oval, dark
dirty green, with paler greyish central area and pale antennae and legs; BL
1.4-2.0 mm. Alatae with narrow, usually interrupted, dusky to dark dorsal
abdominal cross-bands. In often large, ant-attended colonies on the subterra-
nean parts of one-year-old branches of Salix repens (= S. rosmarinifolia),
apparently restricted to plants growing in dry sandy soil (Hille Ris Lambers,
1947b; Pintera, 1987). Recorded from Belgium, The Netherlands and Poland.
Apterous males and oviparae occur in October.
Chaitophorus israeliticus Hille Ris Lambers Apterae oval, colour in life not
known, probably pale; BL 1.0-1.2 mm. Alatae and other morphs unknown.
Originally described from an unidentified Salix sp. in Israel (see Pintera,
1987); also collected from Salix alba in Turkey (BMNH colln).
Ris Lambers, 1966c). Oviparae and both apterous and alate males are recorded
by Saha and Chakrabarti (1988b), who also record predators and attendant
ants.
level, but colonies may also form at tips of twigs (Essig, 1912a). Four other
species have been described from this microhabitat and resembling Ch. macro-
stachyae in most characters with the exception of the degree of sclerotization
and pigmentation of the dorsum, some differences in shape and length of
hairs, and in chaetotaxy of ABD TERG 8; they are Ch. abditus (Hottes),
Ch. canens Richards, Ch. knowltoni Hille Ris Lambers and Ch. salicicorticis
(Essig). None of these differences give reliable separation of paratype and
other material in the BMNH collection, so they are all treated here as
synonyms of macrostachyae. Chaitophorus utahensis, however, synonymized
with macrostachyae by Richards (1972a), is a distinct species (q.v.).
Chaitophorus pakistanicus Hille Ris Lambers Apterae are oval, pale green;
BL 1.2-1.7 mm. Alatae are undescribed; two specimens in the BMNH collec-
tion have no dorsal abdominal markings. In dense colonies along the veins
on both sides of leaves of Salix spp. in Central and East Asia (Afghanistan,
Iran, India, Pakistan, Thailand). Oviparae were collected on S. acmophylla
in December (Hille Ris Lambers, 1966c; Pintera, 1987). Bhagat (1982b)
described a new species of parasitoid from Ch. pakistanicus in Kashmir.
middle legs; BL 1.8-2.4 mm. Alatae have black dorsal abdominal cross-bands.
First instars are yellow (cf. nigrae). On young growth of Salix discolor (BMNH
colln; RLB) and possibly other Salix spp. in Ontario, Canada; also recorded
from British Columbia, Prince Edward Island and North Carolina (Richards,
1972a). Ant-attended. The synonymy of pallipes with saliciniger by Eastop
and Hille Ris Lambers (1976) was in error.
Chaitophorus parvus Hille Ris Lambers Apterae are black with pale anten-
nae and legs; BL 1.2-1.8 mm. Alatae with narrow dorsal abdominal cross-
bands. On undersides of leaves of Salix repens (= rosmarinifolia), attended
by ants. Recorded from Poland, Denmark, Norway and Sweden (see Heie,
1982). Pintera (1987) synonymized this species with niger, but it appears
distinct.
Chaitophorus ramicola (Börner) Apterae are broadly oval, mainly very dull
greyish-olive to black with a more-or-less distinct paler spinal stripe and dark
antennae and legs; BL 1.4-2.6 mm. Alatae have narrow dorsal abdominal
cross-bands. In ant-attended colonies on bark of twigs of mainly broad-leaved
Salix spp., in northern and Central Europe. Apterous males and oviparae
occur in October (Heie, 1982). Chaitophorus brunealineatus Juchnevitch,
1970, described from S. russica in Kazakhstan, is probably a synonym.
Three species of small pale yellow aphids, related to Panaphis and asso-
ciated with Juglans. Quednau (1973) and Chakrabarti (1988) revised the two
palaearctic species, which are also both recorded from Aleurites moluccana
(Euphorbiaceae), presumably acting as a 'reserve host'. Trioxys pallidus
and Praon flavinode specialize on Chromaphis and the related genera
Eucallipterus, Myzocallis, Tinocallis and Tuberculatus (Stary, 1976). Aphe-
linus perpallidus attacks Melanocallis and Monellia as well as Chromaphis.
Chromaphis californica (Essig) Adult viviparae all alate, yellow with dark
antennae and tibiae, and with a short narrow black marginal stripe on each
side of prothorax; BL 1.7-2.0 mm. Immatures and oviparae (collected in
November) have banded antennae. Main host is Juglans californica in
California (Richards, 1968b) and Utah (BMNH colln); there are also records
(perhaps 'casual' occurrences) from Carya illinoensis and Pterocarya spp.
(Walker et al., 1978).
One or more species on Ulmus in East Asia. They are plump-bodied aphids
with rather short antennae, thick dark hairy tibiae and a deeply cleft anal
plate.
Chromocallis nirecola (Shinji) Adult viviparae all alate, green, with black
tibiae and forewing veins bordered with fuscous; BL 3.0-3.2mm. On Ulmus
spp. (japonica, pumila) in Japan, Korea, China and Siberia (Higuchi, 1972;
Pashchenko, 1988b). Biology is little known; alate males were collected in
September in Japan (BMNH colln, leg. M. Sorin), and Paik (1972) recorded
oviparae and males in Korea. Chromocallis pumili Zhang in Zhang and
Zhong, 1982c and Ch. similinirecola Zhang in Zhang and Zhong, 1982c,
described from China, do not seem clearly distinguishable from nirecola.
2n = 18*.
A genus for one East Asian species with a well-defined median suture on the
head, a very long third antennal segment with regularly-spaced, slit-like secon-
dary rhinaria and a rather short processus terminalis.
Chuansicallis chengtuensis Tao All adult viviparae are alate, green, secret-
ing profuse cottony wax threads, with dark hind legs and wings held hori-
zontally over the abdomen when at rest; BL 1.6-2.0 mm. The life cycle and
host relationships are unclear; numerous alate males and apterous oviparae
were described from Rhus semialata ( = javanica) in December in Taiwan
(Tao, 1963), but alate viviparae have been collected from Dalbergia hupeana
620 Cinara
A genus for one East Asian bamboo-feeding species with very long dorsal
abdominal tubercles.
1985); India (A.K. Ghosh, 1982b); Siberia (Pashchenko, 1988b); Korea (Paik,
1972); Japan (Inouye, 1941, 1956, 1970); New Zealand (Baker, 1986);
eastern Canada (Bradley, 1951); eastern USA Pinus-feeders (Pepper and
Tissot, 1973); and US Rocky Mountain region (Palmer, 1952). Danielsson and
Carter (1993a) keyed the subgenus Cupressobium. Voegtlin and Bridges (1988)
provide a catalogue and bibliography of the North American species.
Cinara acutirostris Hille Ris Lambers Dark brown to bronze, with a pattern
of wax dust similar to C. pini; BL of aptera 2.6-3.6 mm. On twigs of Pinus
nigra and P. pinea in Western, southern and Central Europe (England,
France, The Netherlands, Bulgaria, Czechoslovakia, Italy, Spain, Portugal).
Alatae are produced in May-June (Italy) or June-July (England). Apterous
males and oviparae occur in October-November, and eggs are laid on the
needles (Eastop, 1972; Binazzi, 1973).
Cinara alacra Hottes and Essig Colour in life not known; BL of aptera
4.6 mm. Described from a single specimen on Abies concolor in Nevada, USA
(Hottes and Essig, 1953b). There is also a record from California (Smith and
Parron, 1978).
Cinara apacheca Hottes and Butler Probably uniformly dark brown and
shining; BL of aptera about 2.9-3.0 mm (Hottes and Butler, 1955). On Pinus
edulis in Arizona, USA. Not collected since original description, biology and
sexuales unknown.
Cinara apini (Gillette and Palmer) Apterae dark brown mottled with wax
powder on spinal and marginal areas and intersegmentally, with appendages
mainly yellowish to dusky, siphunculi and distal two thirds of tibia black; BL
3.2-4.0 mm. In large colonies on bark of twigs and small branches of Pinus
flexilis (Palmer, 1952), and perhaps sometimes on P. albicaulis (one aptera
622 Cinara
in BMNH colln), in western North America. (The record of this species from
P. banksiana in Quebec (Quednau, 1966) is a misidentification of C. bank-
siana.) Oviparae and alate males occur in September-October, and eggs with
a light covering of wax-wool are laid on the needles.
Cinara atra (Gillette and Palmer) Apterae are rather long-bodied with
narrow head and prothorax, shining black or dark brown without any wax;
BL 2.0-3.0 mm. Immatures are orange-yellow to olive-brown or reddish. In
small colonies on smooth bark of small branches of young pinyon pines, Pinus
edulis and P. monophylla, in Colorado and Utah, USA. Apterous males and
oviparae occur in October-November (Palmer, 1952).
Cinara atripes Hottes Colour of apterae in life not recorded, probably dark
brown or black, with wholly dark legs; BL 1.6-2.3 mm (Hottes, 1958a). On
Picea glauca in Maine, USA. Biology and other morphs unknown, and
apparently not collected since the original description.
collection (leg. A.N. Chowdhuri) was identified as Picea smithiana, and this is
probably the true host. Biology and sexual morphs unknown.
Cinara atrotibialis David and Rajasingh Apterae are brown with darker
antennae and legs, the tibiae and tarsi almost black; BL 3.3-4.3 mm. On young
shoots at needle bases of Pinus kesiya (= insularis), and also recorded from
P. roxburghii (A.K. Ghosh, 1982b). In India (Himachal Pradesh, Meghalaya),
and probably as geographic variants in Thailand (as C. khasyae Robinson) and
the Philippines (Eastop, 1976). Agarwala (1989) studied population trends
in the field, Samanta and Raychaudhuri (1984) reported the effects of a
parasitoid (Pauesia laricis) and Agarwala (1988) studied development in the
laboratory. Sexuales and life cycle are unknown; active stages are present in
December, so it is presumably at least partially anholocyclic. 2n = 10 (Kurl,
1988).
Cinara banksiana Pepper and Tissot Apterae have dark head and thorax
and paler brown abdomen with paired dark markings and large, prominent
shiny black siphuncular cones; BL 2.4-3.6 mm. On previous year's growth or
in dense colonies on new shoots, producing numerous alatae in spring and
early summer; on Pinus banksiana, and apparently specific to this host, in
Canada and northern USA. Oviparae and alate males occur in October and
eggs are laid on the needles. Bradley and Hinks (1968) studied the relationship
with ants in Manitoba.
Cinara bonica Hottes Colour in life not recorded, probably brownish with
some wax dust; BL 3.4-3.6 mm. Collected in Alaska on an unknown host, pro-
bably either Picea glauca or P. mariana and not found since (Hottes, 1956g).
Cinara bonita Hottes Colour in life not recorded, probably brownish with
some wax dust; BL about 5.7 mm. Collected in Alaska on an unknown host,
probably either Picea glauca or P. mariana, and not found since (Hottes,
1956g, 1964).
Cinara braggii Gillette Apterae in life covered in white wax powder, with
legs pale; BL 3.2-3.8 mm. Feeding solitarily on bark of twigs and on new
624 Cinara
Cinara brevispinosa (Gillette and Palmer) Apterae are brown lightly dusted
with wax powder medially and marginally; BL 3.0-3.5 mm. In colonies on
bark of twigs and small branches of Pinus contorta (incl. var. murrayana) in
western North America. Oviparae and alate males in late August-October
(Gillette and Palmer, 1931). Cinara sclerosa Richards is a synonym.
Cinara caliente Hottes Apterae are dusky grey, variably dusted with wax
powder, with black sclerotization of thorax and ABD TERG 1 often in form
of an inverted 'V' (Hottes, 1955c); BL 1.6-2.2 mm. On bark of twigs, espe-
cially on needle-free sections, of Pinus edulis and P. monophylla in western
USA (Hottes, 1960b). Oviparae and alate males in September.
Cinara canatra Hottes and Bradley Shining black, without wax; BL 2.3-
3.5 mm. In spring in dense colonies on bark of smaller branches some distance
from the tips, later moving to form clusters on the bark of the trunk (Hottes
and Bradley, 1953). Probably specific to Pinus banksiana, where it may occur
in mixed colonies with, and be confused with, C. nigra (Pepper and Tissot,
1973); records from other hosts are referable to C. gracilis or C. nigra. In
Michigan, USA, and Ontario, Manitoba and Newfoundland, Canada.
Oviparae in October, males unrecorded.
Cinara caudelli (Wilson) Appearance in life unknown and not found since
original collection, on an unidentified Picea sp. in British Columbia (Wilson,
1919b). BL of aptera about 2.4mm. Life cycle and sexuales unrecorded.
Cinara chibi Inouye Apterae have brown head and thorax and shiny dark
brown to black abdomen, with a pale brown spinal stripe (Inouye, 1962); BL
2.4-3.1 mm. On stems of young trees, or on three- to four-year-old parts of
branches, of Larix spp. in Japan. Oviparae and alate males in October
(Inouye, 1970). Very similar to C. cuneomaculata.
Cinara cronartii Tissot and Pepper Apterae are dark brown with mainly
dark legs; BL 2.6-4.2 mm. This aphid seems normally to restrict its feeding
site to lesions and cankers of the rust Cronartium fusiforme on trunks and
branches of Pinus spp. (taeda, elliotti, clausa, serotina) in eastern USA (Pep-
per and Tissot, 1973). Introduced into South Africa, where severe infestations
cause economic damage to P. taeda (Raubenheimer and Shaw, 1987). Biology
was studied in the laboratory in South Africa by van Rensburg (1989), and
biocontrol with a parasitoid (Pauesia sp.) has been attempted (Kfir et al., 1985;
Kfir and Kirsten, 1991). Sexuales are unrecorded; in Florida the aphids remain
in rust lesions throughout the year. 2n = 10*.
Cinara curtihirsuta Hottes Apterae have orange-red head and thorax and
shiny black abdomen (Voegtlin, 1976); BL 3.0-3.9 mm. On Abies concolor in
California and Oregon, USA, on branches, or on trunks of young trees,
Oviparae in September-December (Voegtlin, 1976), males unrecorded.
Cinara curvipes (Patch) Apterae are entirely shiny or dull black, in early
summer sometimes developing a covering of pale grey wax (Voegtlin, 1976);
BL 3.4-5.5 mm. On trunk or branches of Abies spp. and also feeding on
Cedrus deodora; records from Picea spp. are questionable, although Voegtlin
(1976) found a small colony on Pinus contorta. Widely distributed in USA,
Canada and Mexico (Ortiz, 1982). Oviparae and alate males in September-
October.
Cinara dubia Hottes and Essig Appearance in life unknown, probably dark-
coloured; BL 3.0-3.5 mm (Voegtlin, 1976). Only the aptera is described
(Hottes and Essig, 1954b), from Pseudotsuga taxifolia in Oregon, USA. Also
recorded from California (Voegtlin and Bridges, 1988). Probably a synonym
of C. pseudotaxifoliae (q.v.).
Cinara ferrisi (Swain) Apterae rather elongate oval, brown mottled with
black, slightly pruinose; BL 4.3-6.5 mm. On Pinus albicaulis and P. monticola
in western USA and British Columbia. The only recorded feeding site is the
roots (Bradley, 1961). Life cycle and sexuales unknown. Records from Abies
concolor should probably be referred to other species, e.g. C. curvipes.
Cinara glabra (Gillette and Palmer) Apterae dusky olive green, olive-brown
or black, without dorsal wax; BL 2.5-3.0 mm. Singly or in small groups on
twigs at bases of needles of Pinus ponderosa in western USA; Colorado,
Arizona and Wisconsin (BMNH colln; leg. F.C. Hottes). Oviparae and alate
males in October (Gillette and Palmer, 1924, 1931).
Cinara gracilis (Wilson) Apterae are shiny black, sometimes with a whitish
wax spinal stripe on thorax and ABD TERG 1-2; BL 2.7-3.5 mm. It lives in
dense colonies on bark among needles, often near the bases of twigs and small
branches and is easily disturbed when approached (Pepper and Tissot, 1973).
The normal host is Pinus virginiana, and it is also found on P. taeda (Patti
and Fox, 1981), but the more northerly records from P. banksiana (e.g.
Bradley and Hinks, 1968) seem to be misidentifications of canatra and/or
nigra. In eastern USA.
since the original description (Hottes, 1956f), although the earlier records
of Lachnus abietis on A. concolor in California by Davidson, referred to
C. ferrisi by Swain (1918), possibly apply to this species. Other morphs and
life cycle unknown.
Cinara guadarramae Mimeur Apterae dark brown, often with two dorsal
longitudinal rows of small black sclerites; BL 2.3-3.7 mm. On young branches
of Pinus sylvestris and P. pinaster in Spain. Very similar to C. pini; a detailed
morphometric comparison is made by Garcia Sanchez and Nieto Nafria
(1978). Sexuales and life cycle are not known.
Cinara hattorii Kono and Inouye Apterae dark brown to black, rather
elongate oval; BL 3.2-4.4mm. Alatae have forewing membrane pigmented
between Cula and Cu1b. On Abies sachalinensis and A. mariesii in Japan,
forming colonies on bark of trunk or old twigs l m or less above ground.
Oviparae and alate males in October-November. Cinara konoi is a synonym
(Inouye, 1956).
Cinara horii Inouye Apterae rather elongate oval, with yellow head, light
green thorax and green abdomen (Inouye, 1956); BL 4.4-5.3 mm. Resembling
C. pilicornis, but without a dark cross-band on ABD TERG 8. On twigs
and young shoots of Picea jezoensis in Japan and Sakhalin (Inouye, 1970).
Eggs and fundatrices have been observed in April-May, but sexuales are
unrecorded.
Cinara hottesi (Gillette and Palmer) Apterae dull bluish-black with mainly
yellow appendages (Gillette and Palmer, 1924); BL 3.0-3.5 mm. In large dense
colonies on bark of twigs of Picea spp., moving to larger branches or trunk
as season progresses (Bradley, 1961). Widespread in North America. Oviparae
and apterous males in September-October in Colorado (Hottes, 1955a).
Cinara alaskana, described from Picea sp. in Alaska (Hottes, 1964), is prob-
ably a synonym; its BL is recorded as 2.2 mm, but this is possibly a misprint.
Cinara indica Verma Described from one aptera from Cedrus deodora at
Jammu, India (Verma, 1970b), possibly a small aberrant specimen of C. con-
finis. The much larger alatae described under this name by A.K. Ghosh
(1982b) are probably confinis.
Cinara inscripta Hottes and Essig Fundatrices have dark brown head and
thorax and shiny amber-brown abdomen with large prominent dark brown
siphuncular cones (Bradley, 1961), appearance of later generations in life
unrecorded; BL 3.7-4.2 mm. On Pinus albicaulis in Oregon, USA and British
Columbia, Canada, colonizing young branches in spring; otherwise biology
unknown, and sexuales are undescribed.
Cinara intermedia (Pašek) Apterae mainly pale brown, with two brownish
or greenish-black dorsal longitudinal stripes, converging on ABD TERG 5-8
(Heinze, 1962); BL 2.5-2.7 mm. On bark of trunk of young Pinus sylvestris,
as yet only collected (twice) as single apterae within colonies of C. nuda
(Pintera, 1966). Life cycle and other morphs unknown.
Cinara kiusa Hottes Appearance in life not recorded. Not collected since the
original description; BL of aptera 3.2-3.7 mm, other morphs unknown. On
Abies concolor in Oregon, USA.
Cinara longipennis Matsumura Apterae dark brown to black with dark legs,
body dusted with wax laterally and ventrally (Moritsu, 1983); BL 6.0-7.0 mm.
On Abies spp. (firma, holophylla, sachalinensis), in large spring colonies with
abundant alatae on trunk and older branches of young trees, apterae moving
down onto the roots in summer (Inouye, 1970). In Japan and Korea. Oviparae
Cinara 635
in Korea in late October (Paik, 1972; but the aphid illustrated by Paik as
longipennis is much paler, yellow-brown with black markings).
Cinara lyallii Bradley Apterae dark brown to dull black, dusted with grey
wax powder; BL c. 4.0 mm. In small loose colonies on upper sides of twigs
of previous year's growth of Larix lyalli at high elevations (above 2000 feet)
in Alberta, Canada. Oviparae and both apterous and alate males are recorded,
and there is possibly an abbreviated life cycle (Bradley, 1961). Closely related
to C. laricifoliae.
Cinara maculipes Hille Ris Lambers Apterae pale brown with legs, espe-
cially femora, mottled with dark brown in pantherine fashion; BL 3.7-
4.9mm. On Pinus griffithii in northern India (Himachal Pradesh, Uttar
Pradesh, Jammu, Kashmir) and Pakistan (Muree). Feeding site not recorded.
Oviparae and apterous males in December (Chakrabarti and Raha, 1988). Das
and Chakrabarti (1989a) reared a new Pauesia sp. from this aphid. 2n = 12
(Kurl and Chauhan, 1986).
Cinara melaina Boudreaux Apterae are shiny dark brown, almost black; BL
2.8-3.3 mm. Normally apterae have an uninterrupted cross-band across ABD
TERG 8, whereas in C. atlantica, with which it may form mixed colonies, this
band is usually interrupted medially. On current and previous year's growth
of Pinus spp. of subsect. Australes, feeding at needle bases; there are also
single records from P. resinosa and P. virginiana (Pepper and Tissot, 1973).
In southeastern USA, north to Delaware. Males unrecorded, and oviparae are
rare; probably anholocyclic in southern states. Patti and Fox (1981) studied
populations on P. taeda in South Carolina. Very similar when slide-mounted
to the western US species, C. ponderosae.
Cinara moketa Hottes Apterae shining dark brown with cross-bands of grey
pulverulence; BL 3.3-4.1 mm (Voegtlin, 1976). In colonies on branches, or on
trunks of young trees, of Pinus lambertiana in California and Washington,
USA. Sexuales unknown, apparently anholocyclic in California (Voegtlin,
1976). Tilles (1984) studied ant attendance, and predation by an elaterid
beetle. Cinara hirticula Hottes is probably a synonym.
Cinara nigrita Hottes and Essig Only known from a single alata, distin-
guished by its uniformly dark wings, collected in Arizona, USA. The host is
unknown but surmised to be Pinus ponderosa (Hottes and Essig, 1953b).
Cinara nimbata Hottes Apterae have black head and thorax dusted with
grey wax and abdomen grey-green dorsally with lateral areas cinnamon-brown
(Hottes, 1954c); BL 3.7-4.2 mm. The legs are characteristically patterned, with
a contrasting pale band across the dark distal part of each femur and tibiae
spotted with black. Alatae have diffuse patches of fuscous on forewings as
in C. costata, to which it is closely related. Living singly particularly on stubby
branches close to trunk of tree and attended by ants (Hottes, 1954c). On Picea
Cinara 639
Cinara obscura Bradley Apterae dark brown to black, with pale sections on
tibiae; BL c. 2.5 mm. In large dense colonies on bark of small branches of
Picea glauca in Canada, from Alberta to Newfoundland (Bradley 1953, 1961)
and also recorded from P. engelmannii in British Columbia. Fundatrices were
found on P. glauca in Ontario (BMNH colln, leg. Bradley), but sexuales are
not recorded. C, pallidipes Hottes, described from P. glauca in Maine, USA,
may be a synonym.
generation are nearly all alate and fly to new cones. Oviparae and apterous
males in September-October; eggs are laid on undersides of scales of new
cones.
Cinara parvicornis Hottes Only the ovipara and alate male of this species
are described, collected on Pinus contorta in Montana in September. BL of
ovipara 2.5-3.2 mm (Hottes, 1958d).
Cinara pectinatae (Nördlinger) Apterae are bright olive green with three
diffuse paler green longitudinal bands; BL 2.8-5.0 mm. On Abies spp.,
especially A. alba ( = pectinata). It occurs throughout Europe eastward to
Turkey, living singly on small branches where it sits on the needles, feeding
at the junction of petiole and stem. Oviparae and alate males in October. A
copious honeydew-producer, important to forest beekeepers in Central
Europe and its population dynamics has been much studied because of this
(e.g. Bloc, 1987, in France; Liebig, 1988, in Germany). 2n = 6* (Germany,
2 samples; but Rukavishnikov (1979) recorded 2n = 12 from the former USSR).
Cinara pini (Linnaeus) Apterae are dark grey or greyish-green with black
markings, and with either a slight bronze iridescence or a dusting of grey wax
(Carter and Maslen, 1982); BL 2.5-4.4 mm. On young shoots of Pinus
sylvestris in spring (Pintera, 1966), later on undersides of older foliated twigs
or branches. There are also records from many other Pinus spp., but many
of these are likely to be misidentifications or misapplications of the name pini
(Eastop, 1972). Most European records of C. pini from P. mugo should prob-
ably be referred to P. montanicola Börner. Binazzi and de Silva (1993)
distinguished this as a valid species and noted that it lived in dense colonies
preferably on two- to three-year-old often bare branches of mountain pine.
Throughout Europe and apparently also in Siberia (Pashchenko, 1988b) and
Japan (Moritsu, 1983). North American records cited as pini are probably of
pinea or a native species. Oviparae and apterous and/or alate males in
September-October. Larsson (1985) studied within-crown distribution of C.
pini, Sudd (1983) studied ant attendance, and Michelena Saval and Gonzalez
Funes (1988a) recorded parasitoids. To judge from type material recently
remounted, C. oblonga (Del Guercio) is a new synonym. 2n = 10.
Cinara piniradicis Bradley Apterae and alatae are dark brown, without wax;
BL 4.0-4.8 mm. In colonies on roots of young Pinus banksiana in central
Canada and USA (Michigan). Apterous males and oviparae in September-
October; eggs are laid singly on inner side of needles just above the follicle
sheath (Bradley and Wighton, 1959; Pepper and Tissot, 1973).
Cinara pinivora (Wilson) Apterae have shiny dark brown head, lighter
brown thorax and abdomen with dark dorsal sclerites and spots of grey wax,
black steep-sided siphuncular cones and all legs with extensive pale yellow
sections (Tissot, 1944; as osborni); BL 3.3-4.2 mm. In dense colonies at tips
of branches, or scattered along older sections of twigs, of Pinus spp. of sub-
sections Australes and Contortae in eastern USA and Canada. Oviparae and
alate males in October-November (Pepper and Tissot, 1973).
Cinara pinona Hottes Apterae have brown wax-dusted head and thorax,
dark brown or greenish-brown abdomen with a metallic sheen; BL 3.1-
3.3 mm. In large colonies on twigs of Pinus edulis among the needles, usually
on older trees with yellowish to brownish bark (Hottes, 1960b). In western
USA. Oviparae and alate males in October (Hottes, 1956f).
Cinara ponderosae (Williams) Apterae have dull brown head and thorax
and shiny golden-brown abdomen, with a dorsal pattern of wax patches; BL
2.6-4.2 mm (Voegtlin, 1976). On bark usually near tips of twigs and small
branches of Pinus ponderosa and its varieties; sometimes on other Pinus spp.
in subsection Ponderosae and also recorded from planted P. canariensis and
P. roxburghii. Records from other pines may be misidentifications. In western
North America. Oviparae and alate males in October-November in Colorado
(Palmer, 1926), but populations studied in the Californian Sierra Nevada were
anholocyclic (Voegtlin and Dahsten, 1982).
Cinara russellae Pepper and Tissot Apterae rather shiny black; BL 2.0-
2.5 mm. Found in July in small groups at bases of needles of Pinus banksiana,
feeding on the sheath area rather than on the branch, but in autumn oviparae
and apterae were found in dense colonies with C. nigra on the trunk and
branches (Pepper and Tissot, 1973). Only known from two collections in
Michigan, USA. Oviparae and alate males in late September.
Cinara rustica Hottes Apterae mostly black dusted to grey with wax
powder, except for the posterior abdomen which is dull chocolate brown
(Hottes, 1960b); BL 3.2-3.4 mm. On bark of older twigs or on trunks of Pinus
edulis, with a preference for young vigorous trees. In Arizona and Colorado,
USA. Oviparae (Hottes, 1956d) and apterous males (Hottes, 1960b) in
October.
Cinara saskensis Bradley Apterae dark grey, without wax; BL 5.1-5.6 mm.
On bark of main roots of Picea glauca, near the trunk and just below ground
level (Bradley, 1962). In Manitoba and Saskatchewan, Canada. Life cycle and
sexuales unknown.
Cinara schimitscheki Börner Apterae and alatae broadly oval and somewhat
flattened dorsoventrally, dark brown covered with wax powder; BL 3.3-
5.1 mm. In spring at twig apices among previous year's needles, later on older
branches under bark (Pintera, 1966). Host is usually Pinus nigra (incl, var.
maritima), sometimes P. mugo, P. pinea or P. leucodermis. It occurs through-
out Europe (except Scandinavia and the Baltic), east to Crimea and Turkey.
Life cycle incompletely known; Scheurer (1976) found and described oviparae
Cinara 647
Cinara schuhi Hottes Apterae probably with light brown abdomen but
darker head, thorax and legs (Hottes, 1957a); BL 2.5-2.9 mm. On Abies con-
color in Oregon, USA. Not collected since the original description; biology
and sexuales unknown.
Cinara shinjii Inouye Apterae have pale brown head and thorax and dark
greyish-brown abdomen, with an extensive pattern of white wax powder
(Moritsu, 1983); BL 2.5-3.0 mm. On twigs, branches and stem of Pinus
parviflora and P. himekomatsu in Japan, forming large colonies on lower
parts of tree in summer (Inouye, 1970). Alatae are produced in July, oviparae
and alate males in September-October. Paik's (1965) account of C. shinjii on
P. koraiensis in Korea is of a different, shorter-haired species.
Cinara sonata Hottes Apterae broadly oval, dull reddish-brown to dark grey
with numerous small irregularly distributed black spots, and at least in the
ovipara there are patches of white wax powder on pronotum and narrow cross-
bands of wax on abdominal tergites (Hottes, 1955c, 1957b); BL of aptera
5.3-7.0 mm. On undersides of small branches of Abies magnifica var.
shastensis, and also recorded from A. concolor, in western North America.
Oviparae and alate males in late September.
Cinara subterranea Bradley Only the sexuales are recorded (Bradley, 1956a).
Oviparae are greyish-brown with dark brown legs and antennae, BL c. 3.3 mm,
Cinara 649
Cinara todocola (Inouye) Apterae have brown head and dark green to
greenish-black abdomen, lightly dusted with wax (Inouye, 1936); BL 2.6-
3.0mm. On trunk and branches of young todo-fir trees (Abies spp. in Japan
and Sakhalin). Often in ant shelters. Injurious. Oviparae and alate males in
September-November (Inouye, 1970). Yamaguchi (1976) studied ecology and
morph determination, and Furuta (1984) studied the impact of natural
enemies. Cinara smaragdina is a similar species on Abies spp. in Siberia, but
apparently with a different biology as it feeds on shoots between needle bases
(Pashchenko, 1988b).
Cinara tonaluca Hottes and Wehrle Apterae dusted with white wax, with a
black stripe and two longitudinal rows of black spots showing through the wax
(Hottes, 1952b); BL 2.1-2.6 mm. On woody stems of young trees of Juniperus
monosperma in Arizona, USA. Not recorded since the original description
(Hottes and Wehrle, 1951), sexuales and life cycle unknown.
USA (incl. California and Oregon: BMNH colln) and across Canada. Ovi-
parae and apterous males are produced early, in July-August (Palmer, 1926).
Cinara wanepae Hottes, described from Picea pungens in Colorado, is a pro-
bable synonym (Palmer, 1936b).
Cinara wahhaka Hottes Apterae shining dark brown without wax; BL 1.5-
1.8mm. On foliage of small twigs of Juniperus osteosperma ( = utahensis),
only known from Colorado, USA. All the second generation were alate
(Hottes, 1952c). Oviparae and apterous males (Hottes, 1953b) in September-
October.
Cinara wahtolca Hottes Apterae brownish to dark grey with black marking
and black siphunculi, usually extensively covered with white powder (Hottes,
1953a); BL 2.6-4.6 mm. In large colonies on trunk and needle-free limbs of
Pinus spp. of subsect. Cembroides (pinyon pines) in southwestern USA.
Oviparae and alate males in October (Hottes, 1954b, 1956e). Hottes (1955b)
described a short-haired form on P. edulis in Arizona as a subspecies,
C. wahtolca curtiwahtolca; similar short-haired specimens have also been
collected in Utah on P. monophylla (BMNH colln).
Cinara watanabei Inouye Apterae have head and thorax dark brown,
abdomen shiny brown; BL 4.8-5.4 mm. In large colonies on three- to five-
year-old wood of branches of older trees, or on trunk of young trees, of Pinus
spp. (e.g. koraiensis, parviflora) in Japan, Korea and Sakhalin (Pashchenko,
1988b). Oviparae and alate males in Japan in October-November (Inouye,
1970). Cinara pinikoraiensis, described from China (Zhang and Zhong, 1989),
is possibly a synonym.
Cinara watsoni Tissot Apterae have head and thorax dark chestnut-brown,
abdomen cinnamon-brown with dark brown markings, the whole body dusted
with greyish-white wax (Tissot, 1939); BL 2.7-4.3 mm. In small loose groups
on new shoots, or near ends of previous year's growth, of Pinus spp. of
652 Clethrobius
subsect. Australes in eastern USA. Oviparae and alate males occur in October-
November (Pepper and Tissot, 1973). Population trends of C. watsoni in
South Carolina were studied by Brooks and Warren (1964), and by Patti and
Fox (1981).
Cinara westi Tissot and Pepper Apterae dark brown; BL 2.1-2.8 mm. On
bark of Pinus taeda in southwestern USA, often associated with lesions of
the rust Cronartium fusiforme and usually in ant (Crematogaster) shelters
(Tissot and Pepper, 1967). Sexuales and life cycle unknown. This species is
possibly synonymous with C. rigidae.
Three or four species of large (2.8-5.6 mm) hairy brown aphids living on twigs
and young branches of Alnus and Betula species. All adult viviparae and males
are alate, the ovipara being the only apterous morph. Accounts are available
for Europe (Stroyan, 1977; Heie, 1982; Wood-Baker, 1983) and India (Chak-
rabarti, 1988).
Clethrobius comes Walker Alate viviparae are dark brown, forming clusters
on branches and twigs of Betula spp., often where the new growth is dying
back, or on twigs of Alnus spp. overhanging streams. The populations on
Alnus are regarded by some authors as a separate species, C. giganteus
Cholodkovsky, and attempts to transfer aphids from birch to alder have not
succeeded (Wood-Baker, 1983). No consistent morphological differences have
been found, however, and both birch and alder populations share the same
peculiar structurally heterozygous karyotype (2n = 11; Blackman, 1988).
Clethrobius comes occurs throughout Europe and across Asia to Korea and
Japan. Holocyclic, with sexuales in October-November.
A genus related to Kaltenbachiella. Eastop and Hille Ris Lambers (1976) listed
eight species, but after revisions by Akimoto (1985a) and Smith (1985) only
four species now remain in Colopha, three host-alternating between Ulmus
and Cyperaceae (see below), and one (kansugei) anholocyclic on Carex. One
additional species, caucasica, is included here although it differs biologically
and its true generic position is uncertain.
the veins (Fig. 134C, p. 898). Alatae (BL 1.0-1.6 mm) emerge in June-July,
migrating to roots of Leersia spp.; an attempted transfer to Eragrostis was
unsuccessful (Patch, 1910b, cf. ulmicola). Secondary hosts must also include
Aira caespitosa and Agrostis plumosa, from which the species was originally
described (Monell, 1882). Sexuparae develop to adult on the leaves of the
secondary host, returning to elm in October (Patch, 1910b). Widely distributed
in North America. Some authors (e.g. Palmer, 1952) have synonymized
graminis with ulmicola, but Smith (1985) considered that there was sufficient
evidence to regard the two species as distinct.
Colophina arma Aoki The galls are globular, loosely closed, on the margins
of leaves of Zelkova serrata in Japan (Fig. 125B). Alatae (BL 2.2-2.4 mm)
emerge in July and fly to Clematis stans. where dense colonies (including
sterile 1st instar soldiers) are formed on stems or peduncles. Sexuparae are
produced in autumn and presumably return to Zelkova, but overwintering also
occurs as midget first instar larvae under bark of the secondary host (Aoki,
1980b). 2n (female) = 10, 2n (male) = 8 (Blackman, 1986).
green immatures are dispersed along veins on both upper and lower leaf
surfaces. Alatae, the most commonly collected morph in summer, have black
head and thoracic lobes, paler prothorax and an ill-defined and variably
developed brown patch across ABD TERG 5-6; BL 2.2-3.0 mm. Oviparae
and males occur in October-November. Patti (1983) gives some biological
information.
A genus for one oriental species associated with Juglandaceae, the apterae of
which have fused head and prothorax, reduced antennae and long hair-bearing
processes - although, according to Tseng and Tao (1938), an apterous morph
without such tubercles can also occur in colonies.
1.1-1.8 mm. Even the hind legs lack claws and are often concealed beneath
body. Dorsal median area of prosoma with only 4 pairs of hairs, which are
shorter than the marginal hairs (Takahashi, 1958a). On branches of Quercus
glauca and Lithocarpus sp. (Shinji, 1941; as Thoracaphis takahashii) in Japan
and on Quercus sp. in Korea (Paik, 1972). Ant attendance is not recorded.
A genus for one species related to Nipponaphis, living without host alternation
on Distylium. The alate sexuparae have clear wings and characteristic sensoria-
tion of the antennae. A.K. Ghosh and Raychaudhuri (1973a) revised the genus.
apterous viviparae are known, collected in April (BMNH colln, leg. R. van
den Bosch) and September-October.
One species producing galls on Distylium in Java, but also having free-living
generations on the same host. First instar 'soldiers' occur in the galls
(Noordam, 1991).
Distylaphis foliorum (van der Goot) Galls on Distylium have not been
described. Alatae emerging from galls in late January have BL 2.6-3.3 mm,
long forewings with a broad black pterostigma and the wing membrane
fuscous basad of Cu lb , and secondary rhinaria distributed III 20-26, IV
8-12, V 7-10. Apterae are bright greenish, margins of body with thick
powdery or flocculent wax, and with 2 longitudinal rows of wax patches or
a thick coat of wax on dorsum; BL c. 1.5mm. On undersides of leaves of
Distylium stellare in Java. Apparently monoecious holocyclic, but life cycle
needs further clarification. Oviparae are present on Distylium leaves in
April-August; males are unrecorded (Noordam, 1991).
660 Doraphis
A genus for one or two poplar-feeding species with no obvious close relatives,
the large aleyrodiform apterae of which were originally described as a scale
insect. Alatae have an unbranched media in the forewing and a single oblique
vein in the hindwing. Tao (1969) gave an account of the genus (in Chinese).
Drepanaphis choanotricha Smith and Dillery Alatae black with pale legs.
Head and pronotum striped with white wax, meso- and metanotum and dorsal
abdomen with wax dots, denser posteriorly; BL 1.1-1.7 mm. On low branches
of Acer saccharum (always within 50 cm of ground), in Illinois and North
Carolina, USA.
Drepanaphis knowltoni Smith and Dillery Alatae white with wax, very like
idahoensis except for dark legs; BL 1.6-2.4 mm. On Acer grandidentatum in
Idaho and Utah, USA and also recorded from other Acer spp. in eastern USA
(Smith and Dillery, 1968).
Drepanaphis monelli Davis Alatae have body entirely covered with white
wax powder except for dark thoracic lobes and a brownish-yellow U-shaped
line between the tubercles on ABD TERG 3 and the dark siphunculi; BL
1.9-2.3 mm. On Aesculus glabra, especially on yellowed leaves (the only
Drepanaphis species not associated with Acer). In eastern USA and Canada
(New Brunswick).
Drepanaphis nigricans Smith Alatae have black body with 2-3 longitudinal
wax stripes on head and pronotum and small spots or flecks of wax on
thorax and abdomen. Legs pale, wings clear except for dark spots at ends
of veins. On Acer rubrum, especially on terminals of rapidly growing shoots
662 Drepanaphis
Drepanaphis parva Smith Alatae have reddish-brown head and thorax, the
head and pronotum having 5 longitudinal white wax stripes, and greenish-
grey-brown abdomen with white posterior tergites and longitudinal rows of
white wax dots; BL 1.4-2.6 mm. Legs pale, forewings somewhat cloudy with
veins diffusely bordered. On Acer rubrum in eastern USA and Canada.
Drepanaphis tissoti Smith Alatae black with pale legs, strongly resembling
nigricans (q.v.); BL 1.3-2.0 mm. On Acer rubrum in eastern USA, usually on
growing terminals of young shoots (water-shoots); one record from A. sac-
charum. Doubtfully distinct from D. nigricans.
Drepanaphis utahensis Knowlton and Smith Alatae yellow, with head and
pronotum edged with black, thoracic lobes brown, tubercles on ABD TERG
3 dark and entire body frosted with white wax; BL 1.5-2.0 mm. A very pale
form occurs in midsummer. On Acer grandidentatum in western USA (a
record from A. saccharum in Nova Scotia, Canada, seems likely to be another
species). 2n = 30.
Drepanosiphum 663
A genus for one palaearctic species related to the nearctic genus Drepanaphis,
but the primary rhinaria have no fringe of hairs and dorsal body hairs are long
and stiff as in Chaitophorinae. Both apterous and alate viviparae occur.
A genus for eight species of medium-sized to large elongate aphids with long
antennae and tubular siphunculi, all associated with Acer. All viviparae are
alate, forming 'spaced-out aggregations' on undersides of leaves in shade
(Kennedy et al., 1967). Males are alate, oviparae are large and apterous with
the end of the abdomen extended like an ovipositor (Fig. 6). Discritulus
planiceps, which pupate like Praon in a tent under the aphid, Monoctonus
pseudoplatani and Trioxys cirsii (which emerge from pale mummies), and
Aphelinus thomsoni (which emerge from black mummies) are specialized
parasitoids. Stroyan (1977) provided a generic diagnosis and keys to British
species, and Heie (1982) gave an account of the species of northwest Europe.
664 Drepanosiphum
Dysaphis devecta (Walker) Rolling and reddening the edges of the leaves of
Malus in spring; the galls contain a mixture of normal bluish-grey wax-
powdered apterae and dark green to reddish alatiform apterae with different
degrees of pigmentation and sclerotization of head and thorax. Monoecious,
with an abbreviated life cycle of only three-four generations; oviparae and
alate males are produced within the galls (Hille Ris Lambers, 1945; Forrest,
1970). Forrest and Dixon (1975) studied gall formation, Llewellyn and
Hargreaves (1986) studied the energetics of gall-feeding, and Alston and
Briggs (1977) investigated the genetic basis of resistance to this aphid in apple.
Hitherto restricted to Europe, but Zhang et al. (1990a) record this species from
Dysaphis 669
China. Despite its unusual biology, comparable morphs are very similar to
those of D. anthrisci and its relatives, and indicate a close relationship to that
group.
Primorsk). Life cycle not studied, but suspected secondary host is Libanotis
seseloides (Shaposhnikov, 1986).
A genus with five biologically diverse species. Three are conifer feeders,
one lives on Salix and one on Trochodendron. The Japanese species were
revised by Miyazaki (1971), although his concept of the genus is broader
including related aphids (Ericolophium, Neoacyrthosiphon) that live on
Ericaceae.
Elatobium 673
Elatobium abietinum (Walker), Plate 14a, b Apterae pale green, with two
darker green longitudinal stripes; BL 1.0-2.0 mm. Feeding on the needles of
Picea spp. and, much less commonly, Abies spp. In Europe, Tasmania, New
Zealand (Zondag, 1983a), Chile (Carrillo, 1977), Falkland Islands and western
North America (e.g. Koot, 1983). Monoecious holocyclic (with alate males)
in continental Europe (von Scheller, 1963), apparently anholocyclic elsewhere.
Spring colonies on Picea spp., especially P. sitchensis, cause discoloration
and loss of old needles, sometimes involving serious defoliation (Parry,
1969). Overwintering survival was studied by Powell and Parry (1976) and
flight activity by Carter and Cole (1977). Variation in susceptibility to
attack by E. abietinum has been studied between species of Picea (Nicholls,
1987) and between provenances (Day, 1984; Carter and Nichols, 1988) and
individual trees (Day, 1986) of P. sitchensis. Specialized parasitoids are
Lysaphidus schimitsheki (Stary, 1974) and Ephedrus koponeni (Halme, 1992).
2n = 18.
About 30 species, most of which have host alternation between galls on Ulmus
and secondary hosts in Pyroidea, Grossulariaceae or Compositae. Feeding by
the immature fundatrix on a growing elm shoot in spring induces a galling
reaction in one or more leaves distal to the feeding site, the fundatrix moving
into the leaf-roll after it is formed (Akimoto, 1981). Typically, alatae and
apterous exules of Eriosoma have rather conspicuous siphuncular pores with
partially chitinized rims and surrounding hairs. Alatae usually have a once-
branched media in the forewing, but specimens of E. yangi can have both
forewings with unbranched media.
The limits of the genus are difficult to define; the East Asian Colophina
and the North American Georgiaphis are closely related and often included
as subgenera (Smith, 1985; Eastop, 1987), but are treated separately in
this book. Eriosoma s. str. comprises four or five North American species
forming rosette leaf galls on Ulmus spp. of the americana group, with
676 Eriosoma
two long wax filaments extending from the end of the abdomen; BL
2.0-3.0 mm. Widely distributed in North America. Crataegus is the usual host,
but aphids apparently of this species are also sometimes collected on bark of
Fagus, in colonies of Grylloprociphilus imbricator (Smith, 1985). Smith
distinguished several forms of apterous exules on Crataegus, differing in
dimensions of wax glands. Sexuparae fly to Ulmus (americana, fulva) and
produce sexuales on elm bark in September-October. Heriot (1938) described
differences in the sexuparae, males, oviparae and eggs between E. americana,
E. lanigerum and E. crataegi. Leaf rosette galls similar to those of E. lanigerum
are presumably formed on Ulmus in spring, but strangely, despite the
biological studies of Heriot (1938) and Cox (1939), the spring generations of
crataegi on elm have not been specifically identified; separation of crataegi
in the Ulmus key is based on data provided by Cox for spring migrant alatae
collected on Crataegus in May-June. 2n = 12.
ananassa; Akimoto 1983, 1985b). In Japan, China (Zhang et al., 1990a) and
Taiwan (Tao, 1970; as E. ulmosedens). 2n = 10.
About 14 nearctic species, all living on Pinus needles except one species
(wilsoni) specific to Pseudotsuga. They are long-bodied aphids, as befitting
their needle-feeding habit, and are the nearctic equivalent of Eulachnus, but
smaller and with 5-segmented antennae. Apterae are spindle-shaped, with
tapered abdomen, but the 'spindle' is truncated at the head end, which is
relatively broad. They are often overlooked, as they feed near the bases of
the needles and move quickly when disturbed; their biology is consequently
little known. Sorensen (1988, 1992, 1994) revised Essigella, synonymizing
many of the species described by Hottes (1957c, 1958f).
Active, delicate insects on Tiliaceae, with all viviparae alate. The anal plate
is deeply cleft. Parasitoids specializing on Eucallipterus and related genera
such as Chromaphis, Myzocallis, Tinocallis and Tuberculatus are: Trioxys
pcllidus (emerging from pale mummies), Praon flavinode (forming 'tents'
under the dead aphids) and Aphelinus subflavescens (emerging from black
mummies).
Eucallipterus tiliae (Linnaeus), Plate 9f Alatae are pale yellow with black
markings, including lateral stripes on head and prothorax and two rows of
black dorsal abdominal spots; BL 1.8-3.0 mm. On undersides of leaves of
Tilia spp. in Europe, southwest and Central Asia, North Africa, and intro-
duced to North America and New Zealand. Monoecious holocyclic; apterous
oviparae and alate males occur in late August to October. Numerous aspects
of the biology of E. tiliae have been studied including population dynamics
(Dixon, 1971c; Barlow and Dixon, 1980; Heimbach, 1986); aggregation behav-
iour (Kidd, 1976); flight behaviour (Kidd, 1977); sexual morph production
Euceraphis 685
(Dixon, 1972a); effect on tree growth (Dixon, 1971b; Llewellyn, 1972); suscep-
tibility of different Tilia species (Carter and Nichols, 1986); interactions with
predators in UK (Wratten, 1973; Glen and Barlow, 1980); and parasitoids in
USA (Hajek, 1986). Biocontrol has been used in California, USA (Zuparko,
1983). 2n = 10 (female), 2n = 8 (male); the female has 2 pairs of X
chromosomes and sex determination is X 1 X 1 X 2 X 2 /X 1 X 2 0 (R.L. Blackman,
unpublished data).
[Besides the well-known E. tiliae, there may be at least two other species
in this genus: (i) Specimens with a slightly longer ultimate rostral segment from
Tilia tomentosa in Greece, Turkey and Iran may represent an undescribed
species; (ii) Shinji (1933) described Therioaphis tilicola from Tilia japonica
and T. miqueliana in Japan. T. tilicola has not since been recognized (Higuchi,
1972); it seems more likely to belong in Eucallipterus, rather than in the East
Asian Tilia-feeding genus Tiliaphis.]
A genus of six or more species living on Betula and Alnus. They are rather
large aphids with a relatively short antennal terminal process, knobbed
CAUDA and rounded anal plate. The alder-feeding aphids are somewhat
distinct from the birch-feeders. All adults apart from oviparae are winged,
active insects which secrete bluish-white wax, often in the form of tufts on
their legs. The birch-feeding species often have black dorsal abdominal mark-
ings, but these vary greatly according to season and are often absent in mid-
summer. They seem very host-specific, although adults fly a lot and may be
found on many plants besides their true hosts (but see Pashchenko, 1984). The
oviparae are large, apterous, yellowish to dark brown with dark dorsal
abdominal markings and with the posterior abdomen prolonged into a conical
'ovipositor'. Keys are available for Europe and North America (Blackman,
1977, 1988) and Siberia (Pashchenko, 1988b).
Euceraphis betulae Koch, Plate 10a This name is currently applied to a com-
plex of morphologically similar species with different host associations, distin-
guishable by karyotype differences. Adults of the nominate species feeding on
B. pendula (= B. verrucosa) have a pale green to pale yellow abdomen with
bluish-white wax and a very variable degree of black pigmentation; BL
3.0-4.2 mm. The head and thorax are normally black, both dorsally and ven-
trally, the legs and antennae variably pigmented, sometimes quite dark and
the dorsal abdomen may be unmarked or have transverse black bands (in
spring), or black patches on ABD TERG 3 and 4 only (in autumn). Oviparae
and alate males occur in September-November. Euceraphis betulae occurs
throughout Europe and on introduced B. pendula in North America, Australia
and New Zealand. A morphological comparison with E. punctipennis in
686 Euceraphis
Euceraphis gillettei Davidson Adult viviparae are active, pale green alate
aphids with pale brown thorax, dorsal abdomen never with dark markings and
only a small amount of wax; BL 2.5-3.5 mm. On Alnus spp. (rhombifolia,
rubra, rugosa, tenuifolia) in North America. Oviparae and alate males occur
in September-November. The name may be being applied to more than one
species, as indicated by the variation in karyotype. 2n = 15, 16, 18 or 19
(Blackman, 1988).
Euceraphis ontakensis Sorin Adult alate viviparae are pale green with dark
legs, often with black bars on ABD TERG 4 and 5 and secreting bluish-white
wax; BL 3.0-4.0 mm. On Betula ermanii in Japan; also recorded from Alnus
fruticosa in Japan (Sorin, 1970), and from several other Betula and Alnus spp.
Eulachnus 687
About ten biologically diverse East Asian species, of which three described
from India roll the leaves of woody Rosaceae in spring. They are probably
690 Euthoracaphis
all heteroecious and holocyclic, but with unknown secondary hosts. Eumyzus
have a spinulose, Myzus-like head, but with divergent antennal tubercles, and
rather long stiff dorsal body hairs with tuberculate bases. Chakrabarti and
Bhattacharya (1985) revised the genus.
D.S. Hill). Zhang and Zhong (1985f) distinguished a population from Yun-
nan, China (which included apterae with 5-segmented antennae) as spp.
yulongshanense. Devi and Singh (1987) studied ant attendance. 2n = 20.
Described from Litsea sebifera in Manipur, India. Other morphs and biology
unknown. In every respect except host plant this species seems to resemble
Eu. flavum Takahashi.
host) and Quercus dentata in Japan. Green and yellow alate males and alate
oviparae occur on Castanopsis in May-June in Osaka Prefecture, but not until
autumn in Tokyo (Takahashi, 1962b). Eutrichosiphum sclerophyllum Zhang
(1980a), described from Castanopsis sclerophylla in China, looks like a
synonym.
A genus for one phylloxerid which has apterae with fine hairs and alatae with
a round distal rhinarium, found on oak roots in southern Europe.
An Old World genus of eight or nine species with host alternation between
Pistacia spp. (or, in one case, Toona ciliata) and roots of Gramineae. Galls
on Pistacia are formed by folding and rolling the leaf margins. The fundatrix
of most species initially forms a small temporary gall (e.g. Fig. 126F), usually
near the apex of the leaf, from which the fundatrigeniae emerge to induce the
definitive leaf-edge galls. Alatae emerging from these galls have 6-segmented
antennae, unlike those produced on the secondary hosts which always have
5-segmented antennae. The holocycle takes two years (see Fig. 3, p. 9). Anholo-
cyclic populations of several species occur outside the range of Pistacia on
roots of grasses and cereals, and several species are only known from their
secondary hosts (B & E, 1984, pp. 274-275). For accounts of the gall genera-
tions see Davatchi (1958) and Roberti (1939, 1983).
Forda marginata Koch Galls are formed by upward rolling of the leaf
margins of Pistacia spp. (terebinthus, palaestina) into an elongate spindle
shape (Fig. 126H), yellowish or pale green in colour. Emigrant alatae (BL
1.5-2.1mm) leave the galls in September-November to found colonies on
roots of Gramineae. Holocyclic populations occur in southern Europe,
southwest Asia and northwest India. Anholocyclic populations on grass roots
occur in northern Europe and North America (see B & E, 1984, p. 275).
Roberti (1983) fully described the gall generations and their phenology in Italy,
and Chakrabarti (1987) described the life cycle in northwest India. Karyotype
variation was studied by Blackman (1987); 2n = 17-20 (soma) or 25-40 (germ
line cells).
Fig. 126. Galls of Fordini on Pistacia. A, Aploneura lentisci on P. lentiscus (after Mordvilko, 1935); B,
Asiphoniella cynodonti on P. khinjuk (after Rübsaamen, 1902); C, Baizongia pistaciae on P. terebinthus;
D, Chaetogeoica foliodentata (after Remaudière and Tao, 1958); E, Fordini 'sp. B' (final gall) on P.
atlantica; F, Forda formicaria, fundatrix gall (after Roberti, 1939); G, F. formicaria, final gall; H, F.
marginata on P. terebinthus; I, F. hirsuta on P. vera (after Mordvilko, 1935); J, F. riccobonii on
Formosaphis 701
galls have not been described and on re-examination, A.K. Ghosh (1984b,
p. 381) considered the alatae from them to belong to a different, unidentified
Forda species. Forda kaussarii on P. khinjuk has also been suggested as the
possible primary host form of F. orientalis (Eastop and Hille Ris Lambers,
1976), which is only certainly known from grass roots in India, Pakistan and
Iran (see B & E, 1984, p. 275).
Forda riccobonii (Stephani) Pouch galls are formed on the leaves of Pistacia
atlantica; many of these may be clustered around the edge of a leaflet so that
it completely loses its shape (Fig. 126J). Emigrant alatae (BL 1.9-2.1mm)
emerge in September-November and found colonies on roots of Gramineae.
In Mediterranean area and southwest Asia. Populations on grass roots are
recorded from India (Chakrabarti et al., 1982). This species is keyed on the
basis of specimens from P. atlantica in Morocco (BMNH colln, leg. Bailie)
amd Malta (BMNH colln, leg. unknown); alatae from allegedly similar galls
in Israel (leg. E. Swirski) were much more like F. hirsuta. 2n = 18 in somatic
cells and 30 in germ line (Blackman, 1987).
One species associated with Magnoliaceae in East Asia. The alatae have
antennae with peculiar, reticulate rhinaria, and hind wings with the two oblique
veins united basally as in Pemphigini.
Six rather large North American species living on roots of Salicaceae, closely
allied to Plocamaphis, but more hairy and with siphunculi much reduced or
absent, probably as a result of their subterranean habit. The wing venation
of the alatae seems to vary remarkably within species. Biology has been little
studied. Richards (1966b) reviewed the species (in Plocamaphis).
Fullawaya saliciradicis Essig Apterae have dark brown head and thorax and
yellow-brown to blackish abdomen, with a dusting of fine wax powder; BL
2.1-3.5 mm. Alatae generally have a twice-branched media. Holocyclic on
roots of Salix spp. (laevigata, longifolia). Widely distributed in North America.
Oviparae occur in October-November (Palmer, 1952). Possibly F. braggii
is a synonym, as suggested by MacGillivray (1963) and Richards (1966b).
Geoica 703
About nine species of Geoica are known from roots of Gramineae, and
several of these are known to host-alternate from galls on Pistacia, the
life cycle taking two years (see Fig. 3). One species (mimeuri) is monoecious
on Pistacia. The taxonomy is difficult, particularly around G. utricularia
(Brown and Blackman, 1994). Davatchi (1958) reviewed the morphs on
Pistacia.
Geoica harpazi Brown Galls on Pistacia atlantica in Israel are not clearly
identified, but probably similar in size and shape to those of G. utricularia;
they may be the P. atlantica galls described by Wertheim (1954) as green, of
rather irregular shape and with a wrinkled and uneven surface. Alate
emigrants fly in October to found colonies on roots of Gramineae
(Ammophila, Hordeum, Triticum). See Brown and Blackman (1994).
1958); the gall on P. khinjuk in Israel illustrated by Koach and Wool (1977)
is probably that of G. setulosa. They open in late August-October. Emigrant
alatae (BL 1.8-2.3 mm) found colonies on roots of grasses. Anholocyclic
populations on grass roots occur in Europe (B & E, 1984, p. 276). 2n = 20
and 24 (from grass roots).
Three or four species, related to Old World Baizongia and Asiphoniella, form
galls on Pistacia mexicana and P. texana in Mexico and southern USA, and
probably migrate to roots of Ipomaea spp. and various Compositae. There
is as yet no published description of the galls and gall-feeding generations,
and experimental work is required to associate the known gall-forming
and root-feeding stages. [Geopemphigus floccosus (Moreira) was described
from Ipomaea stipulata in Brazil and was subsequently found on I. batatas
and/or trapped in southern USA, Central America, the Caribbean and as
far south as Säo Paulo, Brazil (B & E, 1984, p. 279). A sexupara trapped
Globulicaudaphis 705
Georgiaphis ulmi (Wilson) Gall is formed by curling one edge of the elm leaf
into a spindle shape, 2-5 cm long. Galled leaves turn yellow, then red, then
drop off (Wilson, 1911). Fundatrix reddish-brown. Monoecious holocyclic,
and apparently specific to Ulmus alata. Alate sexuparae, reddish-brown with
black head and thorax, are produced in second generation (Wilson, 1911),
leaving galls in May-June. In eastern North America.
about two-thirds of length of body. Legs pale with black knees, antennae pale,
annulated with dark markings at joints' (Hille Ris Lambers, 1966c); BL
1.4-1.8 mm. Alatae have 2-3 rather large rounded secondary rhinaria near
base of ANT III. In colonies on undersides of leaves of Quercus dilatata in
Pakistan and India (H.P.; BMNH colln, leg. S.K. David). Biology and life
cycle unknown; apterous viviparae were collected in June, November and
December (BMNH colln). ['Myzocallis floribundi', recorded as a new species
on Q. floribunda (=dilatata) by Verma (1965) but not described, seems to
be this aphid.]
Five species, three palaearctic and two nearctic, associated with Alnus and
Betula. Blackman (1989) reviewed and keyed the palaearctic species, which
feed on aerial shoots, whereas the two North American species are appar-
ently both subterranean (Richards, 1968a). Parasitized by Aphidius aquilus
(= sicarius), which also attacks other Betula-feeding genera such as Betulaphis
and Calaphis (Kallistaphis spp.) (Stary, 1973).
Glyphina betulae (Kaltenbach), Plate 4c, d Apterae are dark green to almost
black with a pale spinal stripe; BL 1.2-2.0 mm. Immature stages are green.
Monoecious holocyclic, in colonies on young shoots of Betula spp., especially
B. pendula, and occasionally on Alnus spp. Ant-attended. The life cycle is
abbreviated; oviparae appear in July and apterous males in August (Heie,
1980). Not known outside Europe. 2n = 28 or 56; the significance of the
karyotype variation is uncertain (Blackman, 1989).
cycle is reported to be like that of G. betulae (Heie, 1980), but the sexual
morphs have apparently not been described. 2n = 8.
One species in East and Southeast Asia, characterized by lack of frontal horns
and by apterae bearing spine-like dorsal hairs, and with yeast-like extracellular
symbionts indicating an affinity with Cerataphis and Tuberaphis. Noordam
(1991) provided a generic diagnosis.
Glyphinaphis bambusae van der Goot Apterae are small pale olive green
to reddish- or greenish-brown, somewhat shiny aphids scattered on young
shoots or at leaf bases of bamboos (Bambusa spp., Phyllostachys sulphurea,
Pleioblastus latiflorus, Schizostachyum brachycladum, Shibataea kumasasa).
Immatures are spotted (Moritsu, 1983, p. 213), and alatae (BL c. 1.9 mm) have
a bluish-black abdomen (Noordam, 1991). Life cycle not recorded. In India
(Assam), China, Japan, Java and Singapore.
very similar and possibly synonymous, differing only in that some of the dor-
sal hairs are blunt or bifurcate.]
About ten species in East and Southeast Asia described from members of
taxonomically diverse plant families. Host plants of several species, including
two from the Philippines (see Raychaudhuri, 1972), are unknown. Greeni-
deoida are rather elongate aphids distinguished from other Greenideinae by
the short last rostral segment with no clear division between R IV and R V.
The siphunculi have no reticulation and are often curved abruptly outwards
near their apices. Alatae where known are distinctive, having forewings with
an elongate, dark pterostigma and the media only once-branched, and hind-
wings much reduced without oblique veins (except in subgenera Neogreeni-
deoida and Pentatrichosiphum, which have twice-branched forewing media
and hindwing with one oblique vein). Raychaudhuri (1956) revised the species
then known, and more up-to-date accounts will soon be available.
A genus for one North American species with distinctive wax plates in the
fundatrix, oval secondary rhinaria in alatae and first instar exules with
thickened hind femora.
A genus of about six species with similar host plant associations to Hor-
maphis, but with a gall (or 'pseudogall') dwelling stage on Betula as well as,
or instead of, Hamamelis. Alatae have two oblique veins in the hindwing.
Species-level taxonomy is problematic and needs more biological studies for
clarification.
Fig. 127. Hamamelistes betulinus on Betula leaves, showing blisters on upper surface and aphids
feeding beneath (from Mordvilko, 1935).
716 Hamamelistes
A genus for one East Asian species related to Calaphis but with fewer hairs
on the first tarsal segment.
Two species, one known only from galls on Zelkova in Japan and the
other known only from a monocotyledenous secondary host in India. The
taxonomie position of the genus was reviewed by Akimoto (1983).
Hemipodaphis persimilis Akimoto Forming leaf-roll galls on Zelkova ser-
rata. Emigrant alatae are very small (BL 0.6-0.7 mm), brown, emerging in
July-August to migrate to an unknown secondary host (Akimoto, 1983).
Populations persist in the galls until October (Akimoto, 1992). Aggressive
behaviour by first instar larvae was described by Aoki (1978; as Colopha
sp. nr caucasica). 2n = 36 (Blackman, 1986).
A genus for one small oak-feeding species in southern Europe and southwest
Asia. Colonies include apterous viviparae with very long dorsal hairs; they
appear to be adapted to living in leaves rolled by caterpillars.
HOPLOCHAITOPHORUS Granovsky
Drepanosiphinae: Phyllaphidini
HOPLOTHORACAPHIS Pramanick,
Samanta and Raychaudhuri Hormaphidinae: Cerataphidini
* [Quednau (1971) recorded H. quercicola as feeding specifically on upper sides of leaves of 0. macrocarpa, but this is
clearly not the case. In fact Monell (1879) described his aphid from the same host and feeding site as Quednau's
heterotrichus. The latter was described from individuals intermediate between apterae and alatae, and further work may
reveal it to be a synonym of quercicola; besides varying seasonally, apterous specimens with different degrees of alatiformity
may have varying proportions of spine-like and finer dorsal hairs and also differ correspondingly in the length and form
of antennal hairs.]
Hormaphis 721
without host alternation (von Dohlen and Gill, 1989). Both life cycles occur
together at intermediate locations, so it is likely that the same name is being
used for two sibling species. 2n = 18 (for both forms)*.
A Myzus-like genus with seven North American and three Indian species,
related to Ovatus but with swollen siphunculi and with a strongly wrinkled
tergum in apterae. Three American species apparently host-alternate between
woody Rosaceae (usually Crataegus) and secondary host plants in several
families; however, as pointed out by Voegtlin (1984), there is in no case any
published evidence of host transfer tests to verify the life cycle. The other
species are known only from herbaceous plants. Smith (1982) keyed the North
American species and Voegtlin (1984) reviewed their biologies. L.K. Ghosh
(1986) reviewed the species in Himachal Pradesh, India.
Hyalomyzus tissoti Nielsson and Habeck The only morph described from
Crataegus spp. (praeformosa, vicana) is the alate ?fundatrigenia; it has brown
thorax and yellowish-brown to brown abdomen; BL 1.0-1.5 mm (Nielsson and
Habeck, 1971). 'Secondary hosts' are recorded as Drosera spp. and Isnardia
intermedia, but the species is only known from Florida, where anholocyclic
populations are likely, so the host alternation requires confirmation.
Hydronaphis 723
A genus once used in a much broader sense to include most Aphidinae with
small siphunculi, but now restricted to two or three very similar species of
elongate-bodied, mealy aphids that alternate between Prunus spp. and
Phragmites. Stroyan (1984) and Heie (1986) reviewed the genus; see also B &
E, 1984, p. 282-283. Basky and Szalay-Marszo (1987) studied morphological
differences between H. amygdali (from peach) and H. pruni (from plum). The
couplet provided in the Prunus key takes account of their work, but will
still be unreliable for single specimens. Spampinato et al. (1988) studied
electrophoretic differences which indicate that 'H. amygdali probably includes
two sibling species, one mainly colonizing P. amygdalus and the other favour-
ing P. persica as primary host. Basky (1982) studied natural enemies and
Huang et al. (1986) studied phenology of H. amygdali in China. 2n = 10
(for pruni).
724 Hysteroneura
One originally North American species in which the CAUDA is long and pale
and the hindwing has only one oblique vein.
Hysteroneura setariae (Thomas) Apterae are dark rusty brown with dark
siphunculi, contrastingly pale sections of the antennae and tibiae and a
pale CAUDA; BL 1.3-2.3 mm. In spring on twigs and leaves of tender new
shoots of Prunus domestica in North America. Host alternation occurs
to aerial parts of Gramineae, but ant-attended populations may persist on
plum throughout the year (Gillette and Taylor, 1908). Anholoyclic popula-
tions of H. setariae have become widely distributed throughout the warmer
regions of the world on Gramineae (see B & E, 1984, pp. 285-286), including
bamboos. Carver (1976) found sexuales and eggs on Prunus persica in
southern Australia, but it is not known if the holocycle was completed.
2n = 12.
Illinoia alni (Mason) Apterae spindle-shaped, pale green with a dark green
median stripe and dark spots at bases of siphunculi; BL 2.5-3.1mm.
Immatures pale green. In small colonies on undersides of leaves of Alnus
rugosa in northeastern USA and eastern Canada (MacGillivray, 1958).
Monoecious holocyclic. 2n = 10.
undescribed. Introduced from North America into Europe, New Zealand and
South Africa. 2n = 10.
INDIOCHAITOPHORUS Verma
Drepanosiphinae: Phyllaphidini
One elm-feeding species related to Sinochaitophorus, but the aptera has short
antennal hairs and a different pattern of dorsal sclerotization.
Eight species primarily associated with Ulmus and related to Colopha, but the
hindwing has two oblique veins, and the feeding position of the first instar
fundatrices is such that the gall arises from the mid-rib of the leaf near its
base, rather than from the leaf lamina. A.K. Ghosh (1981) reviewed the genus,
and Akimoto (1985a) reviewed the species in East Asia.
gall are anomalous and need additional confirmation, being based on only one
collection. In India (Uttar Pradesh).
Kaltenbachiella ulmifusa (Walsh and Riley) Galls on Ulmus rubra are large,
spindle-shaped, bag-like, about 2.5 cm long, green when young and becoming
730 Kurisakia
About seven species in East Asia, described from Juglandaceae (or from
perhaps misidentified trees with similar leaves) and Fagaceae. Closely related
to Glyphina and in fact, although Kurisakia are generally paler and less
sclerotized than Glyphina, there are no good morphological distinguishing
features between these two genera. Many of the characters used to distinguish
between species and subspecies of Kurisakia are subject to environmental
variation, so that the separate identity of most species needs experimental con-
firmation; possibly there are only two or three valid species in the genus. There
is no information on sexual morphs or life cycles for any species of Kurisakia.
Accounts are available for Japan (Takahashi, 1960a) and China (Chang and
Zhong, 1979a).
Kurisakia indica Basu Apterae oval, colour in life not recorded, antennae
and legs pale; BL 1.6-1.9 mm. Alatae have paired dark markings on most
abdominal tergites. On undersides of leaves of Engelhardtia spicata in
West Bengal, feeding without evident injury to host (A.N. Basu, 1968).
Similar aphids were collected in Sikkim (BMNH colln, leg. M.R. Ghosh and
S. Chakrabarti), but from an unlikely host, Fluggea macrocarpa (Euphor-
biaceae). Life cycle unknown.
About 14 palaearctic and one nearctic species of medium to large 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 Europe (Heinze, 1962), Poland
(Szelegiewicz, 1978), Hungary (Szelegiewicz, 1977), India (A.K. Ghosh,
1982b), Middle East (Bodenheimer and Swirski, 1957) and Japan (Sorin,
1980). Until about 1930 the species were often placed in Pterochlorus and the
name Lachnus was applied to aphids of the genus Cinara. Several species of
Pauesia are recorded as parasitoids in Asia.
Lachnus acutihirsutus Kumar and Burkhardt Apterae have black head and
thorax and brownish-black abdomen; BL 2.8-4.4 mm. Alatae have forewings
732 Lachnus
mostly pigmented, except for two large clear triangles on basal half and a
small clear spot between pterostigma and Rs. On twigs of Quercus incana and
Qu. dealbata in northern India (Uttar Pradesh, Himachal Pradesh). Sexuales
and life cycle unknown. 2n = 16 (J. Dutta and D.C. Gautam, pers. comm.).
Lachnus salicis Chakrabarti and Raha Colour in life not recorded, probably
brownish; BL of aptera 3.5-5.4mm. Alatae have clear wings. In large dense
colonies on stems of Salix babylonica and S. tetrasperma, attended by ants.
Sexual morphs not recorded; apterous viviparae and immatures may over-
winter in bark crevices (Chakrabarti and Raha, 1988). In Uttar Pradesh, India.
Possibly a synonym of L. longirostrum, a very similar aphid described from
Salix fragilis in Himachal Pradesh (David and Ghosh, in A.K. Ghosh, 1982b).
It is likely that these willow-feeding aphids, with general appearance and
biology very like Tuberolachnus salignus but lacking a dorsal tubercle, are
wrongly placed in Lachnus.
base of media to distal part of Cula. On twigs and stems of Quercus spp. and
Castanea spp. in East and Southeast Asia (India, Japan, Korea, China,
Siberia, Java, Malaya). Dense populations of a Lachnus that appears to be
this species were also collected on twigs of Casuarina equisetifolia at high
altitude in New Guinea (BMNH colln, leg. J. Szent-Ivany). Records from
Castanopsis (= Shiia) are possibly all L. shiicola. Monoecious holocyclic in
Japan, with apterous oviparae and alate males (Shinji, 1927; Moritsu, 1983);
possibly anholocyclic in warmer regions. Lachnus siniquercus Zhang, described
from Q. liaodongensis (?liaotungensis Koidzumi) in China (Zhang and Zhong,
1982b) is closely related and could be large specimens of tropicalis. The
tropicalis/shiicola group perhaps involves a complex of species or races,
paralleling that thought to occur in L. roboris. B.C. Das and Chakrabarti
(1989a) recorded a new parasitoid from L. tropicalis, in India. 2n = 12, 13 or
16 (Blackman, 1986).
Lachnus wichmanni Hille Ris Lambers Apterae 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
Germany and Italy. Monoecious holocyclic; apterous oviparae and small
apterous males in August in southern Germany (Hille Ris Lambers, 1956c).
A genus for one species on Populus, differing from Chaitophorus in the very
short processus terminalis, 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).
Three species have been described in this genus, one in North America and
the others in the Himalayas and Taiwan. Very large, bark-feeding aphids,
characterized in the alatae by the elongate pterostigma extending around tip
of forewing. The differences between the species are not clearly defined and
it seems possible that they could even represent populations of one poly-
phagous species introduced to various parts of the world.
listed 292 host records from trees in 24 genera and 16 families. Monoecious
holocyclic, with alate males, in northern USA (Washington, DC; Wilson,
1909), anholocyclic in southern states (Bissell, 1978). Marked fluctuations in
population size occur, this species being very abundant in some years (e.g.
Tissot, 1944).
Two or three palaearctic species with host alternation between Rhamnus and
Polygonum. The antennal tubercles are smooth and divergent, the apterae
have secondary rhinaria on ANT III, the siphunculi are tapering and rather
short and the CAUDA is reduced to such an extent that it is almost
indiscernible.
Only a few of the 120-or-so species in this genus are tree-dwelling, including
the subgenus Neocorylobium with five species associated with Betulaceae.
Sitobion is here treated as a separate genus. For an introduction to the genus
as a whole and bibliographic information see B & E, 1984, p. 294.
alate vivipara, ovipara and alate male. Sexuales in October. See also Mac-
Gillivray (1968).
About ten East and Southeast Asian species, almost all on Rubus (Miyazaki,
1971), but one species is described from Ficus.
One species near Tinocallis, but with differences in the ventral structure of
the head.
One nearctic species related to the East Asian genera of Fordini of subtribe
Melaphidina (Nurudea, Kaburagia, Schlechtendalia) and with a similar alter-
nation between Rhus and mosses. The host association was discussed by
Moran (1989).
Melaphis rhois (Fitch) Forming closed, sac-like red and white galls (Fig. 129)
on leaflets of Rhus spp. (glabra, typhina) in North America. Emigrant alatae
(BL 1.4-1.7 mm) emerge in late August-October and fly to mosses. Anholo-
cyclic populations also occur all year round on mosses and have been intro-
duced into Europe (Sweden, UK). Hebert et al. (1991) studied genetic variation
in local populations by enzyme electrophoresis. 2n = 26*.
Fig. 129. Gall of Melaphis rhois on Rhus typhina (after Moran, 1989).
Mesocallis fagicola Matsumura Very small pale yellow alatae; BL c. 0.6 mm.
ANT III with about 6 secondary rhinaria spaced out along segment. Described
from Fagus sylvatica in Japan (Matsumura, 1919) and not recorded since.
Biology and sexuales unknown.
Mesocallis obtusirostris Ghosh Alatae pale yellow; BL 1.2-1.5 mm. ANT III
with 3-7 rhinaria, mostly on basal part. On Alnus nepalensis (with one record
from Betula alnoides) in India (Uttar Pradesh, West Bengal) and Nepal
(Chakrabarti, 1988). Sexual morphs were reported by Chakrabarti (1988,
p. 69).
METANIPPONAPHIS Takahashi
Hormaphidinae: Nipponaphidini
About seven East Asian species distinguished from Nipponaphis by the orna-
mentation of the tergum, consisting of rounded pustules which are rather
evenly spaced and sized over the central part of the prosoma. The fused ABD
TERG 2-7 have marginal hairs, but there are no spinal hairs on ABD TERG
7 and ABD TERG 8 has 2 (or rarely, 4) hairs. Two species in Japan have been
shown to alternate between Distylium and Castanopsis (Sorin, 1987b), the
other species that probably belong in the genus are only known from Fagaceae.
Metanipponaphis is in need of revision and no key to species on Castanopsis
is attempted; A.K. Ghosh and Raychaudhuri (1973a) provided a partial key.
One species on Quercus in Japan, in which the apterae have very short
unsegmented antennae and prosoma partially fused with abdominal plate; the
alatae as well as the apterae lack siphuncular pores.
eyes and first tarsal segments with 3 hairs; BL c. 2.6mm. Probably the
species is incorrectly placed in Micromyzus, which are typically small fern
feeders.
One species related to Thoracaphis, but the aleyrodiform apterae are without
dorsal hairs except on ABD TERG 8 and have very short unsegmented anten-
nae. It was erroneously transferred to Neothoracaphis by Eastop and Hille Ris
Lambers (1976).
Mimeuria ulmiphila (del Guercio) Forming terminal leaf nests on Acer spp.,
especially A. campestre, by inhibition of shoot growth, twisting and folding
of leaves. Fundatrices are olive green-grey, covered with white wax wool; BL
3.5-4.5 mm. They give rise to numerous dark brown alatae (BL 2.6-3.3 mm),
which fly over an extended period (June-November). Apterous exules live
mainly on roots of Ulmus; they are yellow, thickly wax-powdered, BL
1.3-2.3 mm, encased singly in brown microrrhizal cysts (Marchal, 1933;
Krzywiec, 1964). Such cysts have also been found on Rubus roots close to
Ulmus (Vernon, 1957). Sexuparae are produced in autumn and return to Acer,
but anholocycly on Ulmus roots often seems to predominate especially in
Western Europe. Krzywiec (1964) suspected that anholocyclic overwintering
of immature stages might also occur on bark of Acer. Recorded from Britain,
France, Germany, Hungary, Italy, The Netherlands, Poland, Russia and
Turkey. For a full account of the life cycle in Poland and descriptions of all
stages see Krzywiec (1962, 1964).
Mindarus 749
Mindarus victoria Essig Apterae are a soft jade-green, covered with thick
white wax wool; BL c. 1.9mm. Alatae green, wax-covered, with narrow dark
dorsal abdominal cross-bands; BL c. 2.5 mm. On shoot tips of Abies grandis
in British Columbia, Canada. Oviparae and very small dusky apterous males
in June. Essig (1939) described and illustrated all the morphs.
A genus for one large palaearctic species with many distinctive features,
including a long tapering antennal processus terminalis, feeding on Betula. See
Stroyan (1977) for a generic diagnosis.
Monellia hispida Quednau All viviparae alate, pale yellow or colourless with
banded antennae, body often with 4-6 faint yellow spots internally, and later
754 Monelliopsis
generations with a broad dusky stripe running around front and sides of body,
as in M. caryella but much paler; BL 1.2-2.0 mm (Bissell, 1978). Monoecious
holocyclic on Carya spp. in eastern North America; sexuales in October-
November.
Monellia medina Bissell All viviparae alate, bright yellow with banded
antennae, and summer/autumn generations with a heavy black band running
around front and sides and along leading edge of forewing as in M. caryella;
BL c. 2.3 mm. Recorded from Carya spp. in Maryland and Ohio, USA
(Bissell, 1978). Very similar to M. caryella. Sexuales undescribed.
Monelliopsis caryae (Monell) All viviparae alate, yellow with banded anten-
nae and, in fully pigmented specimens, a dusky median streak on the head
and small dusky spots at bases of dorsal abdominal hairs (Richards, 1965);
BL 1.2-1.8 mm. Immatures have black patches at bases of dorsal and marginal
hairs. On leaves of Juglans nigra, widely distributed in North America.
Oviparae in Utah in October (BMNH colln, leg. G.F. Knowlton). 2n = 18.
of dorsal abdominal hairs and a dark ventral spot near apex of hind femur
(Richards, 1965); BL 1.2-1.5 mm. Immatures have dark spots at bases of
dorsal hairs. On leaves of Juglans spp. and Carya spp., widely distributed in
North America. Most records of this species on Carya illinoensis should prob-
ably be referred to M. pecanis (see below). Sexuales apparently unrecorded.
2n = 10.
Monelliopsis pecanis Bissell All viviparae alate, yellow, with darker mark-
ings (when developed) similar to M. nigropunctata; BL 1.2-1.7 mm. On leaves
of Carya illinoensis in USA, Mexico and introduced to Egypt (BMNH colln,
leg. A.A. Attia) and South Africa. Sexuales, and differences from M. nigro-
punctata, with which it was confused for many years, were described by Bissell
(1983). Alverson and English (1990) studied population dynamics, Edelson
and Estes (1987) studied natural enemies in South Carolina, and de Villiers
and Viljoen (1987) investigated control with systemic insecticides in South
Africa.
One North American species in which the alatae from galls have antennae with
processus terminalis longer than base and bearing 'clear spots' of unknown
function.
Muscaphis smithi (Hille Ris Lambers) Apterae (fundatrices) are very plump-
bodied, dark brownish-black with a reddish tinge, with jet black siphunculi;
BL 3.0-4.0 mm. In tightly curled and reddened leaves of Malus angustifolia,
attended by ants. In North Carolina, USA (Hille Ris Lambers, 1962a).
Immature progeny are reddish, all developing into reddish emigrant alatae (BL
1.7-2.1mm), which fly away in July to colonize an unknown secondary
host - probably a moss.
Myzocallis boerneri Stroyan Alatae pale yellowish, with head and thorax
sometimes partly dusky, dorsal abdomen with small paired spinal and
marginal specks of brown pigment (visible with a hand lens), antennae ringed
with brown-black; BL 1.3-2.2 mm. Immatures are yellow with paired, dusky,
rather indistinct spinal and marginal spots. On undersides of leaves of Quercus
spp. (cerris, ilex, infectoria, suber, variabilis) in Europe, southwest Asia
(Israel, Iran, Lebanon) and introduced on European oaks to South Africa
(I.M. Millar, pers. comm.), New Zealand (Rohitha, 1982) and California.
Monoecious holocyclic; sexuales in October-November (northern hemisphere).
Very similar to, and often confused with, M. komareki or M. schreiberi (see
Hille Ris Lambers and Stroyan, 1959; López and Nieto Nafria, 1983).
2n = 14*.
markings. The antennae are ringed with brown, the forewing membrane
usually has a pattern of infuscation, but is sometimes almost hyaline and the
siphunculi are always dark. BL 1.5-2.0 mm. Immatures yellow, pink or
brownish with darker dorsal abdominal spots (Richards, 1968f). Living singly
on undersides of leaves of Quercus spp. (mostly on white oaks), widely dis-
tributed in North America and in Cuba (Holman, 1974). Monoecious holo-
cyclic; sexuales in September-October.
morph (BMNH colln). Çanakçioglu (1975) records this species from Q. cerris
and Q. aegilops in Turkey, but does not specify which morph(s) were col-
lected, and perhaps had boerneri or komareki. Life cycle requires clarification;
sexuales in late autumn and winter (Bodenheimer and Swirski, 1957), and
fundatrix-like alatae in April (BMNH colln, leg. B. Halperin). 2n = 14*.
Myzocallis (Pasekia) komareki (Pasek) Alatae yellow, with head and thorax
variably marked with black (typically there are fine, black lines on the pro-
thorax, roughly in the shape of a pair of rectangles) and dorsal abdomen with
a series of large, paired quadrate spinal sclerites, which have dark margins and
paler central areas, and similar-sized but paler and more uniformly-pigmented
marginal sclerites. BL 1.1-2.2 mm. On Quercus spp. (aegilops, cerris, cocci-
fera, ithaburensis) in southeastern Europe, Mediterranean area and Middle
East (Bulgaria, the Czech and Slovak Republics, Greece, Iran, Israel, Italy,
Turkey). Monoecious holocyclic; sexuales in Iran in November (BMNH colln,
leg. S.H. Hodjat). 2n = 14*.
Immatures are very pale, almost translucent yellow, with antennae ringed with
black. On Quercus spp. (mostly red or pin oaks) in eastern North America
(Boudreaux and Tissot, 1962; Richards, 1968f).
Myzocallis schreiberi Hille Ris Lambers and Stroyan Alatae pale yellow, with
head and thorax slightly darker; the prothorax often has short lateral streaks
of dark pigment, the dorsal abdomen has small transversely oval, often dusky
to dark, spinal spots and paler marginal sclerites, the legs are mainly pale and
the siphunculi usually dark. BL 1.3-2.2 mm. Immatures whitish to pale straw
yellow with variable pigmentation of dorsal sclerites (Hille Ris Lambers and
Stroyan, 1959). On undersides of leaves of Quercus ilex in Europe (England,
France, Italy, Greece, Yugoslavia); rarely on other oaks. Very similar to
M. boerneri; the distinction between these two species was discussed by
Barbagallo and Stroyan (1982). Sexual morphs unknown; apparently anholo-
cyclic, at least in England and Sicily. C.E.J. Kennedy (1986b) investigated
the method of attachment of the tarsi to the smooth leaves of Q. ilex.
have brown head, black pterothorax and pale green abdomen with marginal
sclerites (Nevsky, 1929a). In colonies on undersides of leaves of Prunus
amygdalus and P. mahaleb in spring (April-July), causing leaves to curl
spirally, but not change colour (Vereshchagin and Narzikulov, 1961; Nar-
zikulov, 1965b). In Central Asia (Turkmenia, Tadzhikistan, Uzbekistan). Life
cycle apparently unknown.
Myzus cerasi (Fabricius) Apterae shiny, very dark brown to black, with
antennae and legs bicoloured yellow and black and siphunculi and CAUDA
wholly black; BL 1.5-2.6 mm. Alatae have a yellow-brown abdomen with an
extensive dorsal patch. In dense, often ant-attended colonies at shoot tips of
Prunus spp., especially P. cerasus and P. avium, in spring, curling leaves and
often doing severe damage. Heteroecious holocyclic, migrating to secondary
hosts in Rubiaceae (Galium), Scrophulariaceae (Veronica and, especially in
North America, Cruciferae. However, part of the population may persist
through summer on Prunus (Gilmore, 1960). In Europe, across Asia to
Pakistan, India, Siberia and Korea. Introduced into Australia, New Zealand
and North America. Rakauskas (1984) studied phenology and natural enemies
of M. cerasi in Lithuania. Apparently a complex of species or subspecies with
different life cycles and/or secondary host associations is involved (Dahl,
1968; Gruppe, 1988a); electrophoretic differences have been found between
populations on P. avium and P. cerasus (Gruppe, 1988b). North American
emigrant alatae from P. pennsylvanica and P. virginiana have a relatively
short antennal PT. In Japan, M. cerasi umefoliae occurs on P. mume and is
768 Myzus
morphologically like European M. cerasi, but does not colonize cherries and
migrates to Artemisia capillaris (Takahashi, 1965c); Myzus prunisuctus,
described from Prunus sp. in China (Zhang and Zhong, 1980a), could perhaps
be this aphid. See also B & E, 1984, p. 311. 2n = 10 (European and North
American populations) or 2n = 12 (India, possibly umefoliae; Khuda-Bukhsh
and Pal, 1986a; Kurl and Chauhan, 1988).
Myzus cornutus Medda and Chakrabarti Apterae pale, colour in life not
recorded; BL c. 2.0mm. Alatae have a dark central dorsal abdominal patch
on ABD TERG 4-6 and separate cross-bands on other tergites. Spring popula-
tions occur on Prunus cornuta in India (Uttar Pradesh), feeding on undersides
of leaves, which are rolled longitudinally and turn red or yellow (Medda and
Chakrabarti, 1986b). Apparently heteroecious holocyclic; migration occurs
during June to an unknown secondary host (Medda et al., 1986). B.C. Das
and Chakrabarti (1990) described a new parasitoid species (Aphidius staryi)
from this aphid.
Myzus ligustri (Mosley) Apterae shiny yellow, with distal parts of siphunculi
brownish-black; BL 1.0-1.5 mm. Sporadically common on privet hedges
(Ligustrum ovalifolium, L. vulgare); leaves are rolled longitudinally into nar-
row tubes and spotted with yellow. It was described from England (1841)
before the introduction of the East Asian L. ovalifolium to Europe, so pre-
sumably the native European L. vulgäre is the type host. Monoecious holo-
cyclic, with oviparae and alate males in November. In Europe and introduced
into North America. 2n = 12.
Myzus padellus Hille Ris Lambers and Rogerson Apterae bright yellow
with blackish-brown head; antennae, femora and tibiae bicoloured yellow and
blackish-brown, siphunculi black. BL 2.0-2.3 mm. Alatae have a yellow
abdomen with a black dorsal patch. Forming red and yellow blistered leaf-
galls on Prunus padus in spring (Hille Ris Lambers and Rogerson, 1946). In
Europe (England, Norway, Sweden, Finland, Austria). Heteroecious holo-
cyclic, apparently migrating to Labiatae (Galeopsis spp. - records from
Norway and Sweden; Tambs-Lyche, 1957) and Scrophulariaceae (Pedicularis
verticillata, Rhinanthus sp. - specimens from Austria; BMNH colln, leg.
H.L.G. Stroyan and R.N.B. Prior). The host plant transfers to confirm the
life cycle have not been carried out, however, and sexuales are unknown.
growing season (Alma and Arzone, 1983). Presumably native to East Asia,
where it is recorded from China, Taiwan, Korea, Thailand and Japan.
Introduced into Europe and southwest Asia, where it has become a serious
pest on peaches (Arzone and Alma, 1984; Tkachuk, 1988). It is a vector of
plum pox virus (Sharka disease). Myzus varians was described from North
America, but presumably only occurs there as anholocyclic populations on
secondary hosts. (See also B & E, 1984, p. 315.) 2n = 12.
Myzus yamatonis Miyazaki Apterae black in life, with antennae and legs
pale except at apices and siphunculi and CAUDA black; BL 1.4-1.6 mm
(Miyazaki, 1971). Alatae black in life, but with variably developed abdominal
sclerotization, the central patch being irregular and with large perforations.
On Prunus spp. (cherry trees), especially P. yedoensis, spring populations
causing heavy leaf curl. Heteroecious holocyclic, migrating to Labiatae (Salvia
nipponica, Plectranthus spp.). although populations on Prunus may persist
throughout the growing season. Yano et al. (1984) studied predation by
Elateridae.
Nearctaphis californica Hille Ris Lambers Described only from the secon-
dary host, Melilotus albus, in California, USA (Hille Ris Lambers, 1970c),
but apterae collected in British Columbia on Sorbus aucuparia appear to be
Nearctaphis 771
An oriental genus on Betulaceae, nominally with four species, but the dif-
ferences between N. alba in Japan and N. pusilla in India are those to be
expected between small and large specimens respectively of the same species,
and the other two species are also doubtfully distinct from each other.
Chakrabarti (1988) and A.K. Ghosh and Quednau (1990) reviewed the three
Indian species.
Neobetulaphis alba Higuchi Apterae are pale yellow in life, BL c. 1.5 mm.
Alatae are undescribed. On Betula platyphylla var. japonica in Japan (Higuchi,
1972). Life cycle unknown. Perhaps a synonym of N. pusilla.
Both the known species live on Magnolia in East Asia. They are reviewed by
Takahashi (1965b), Higuchi (1972) and Moritsu (1983; including colour photo-
graphs of both species). Sexuales and life cycle are not recorded for either
species.
Neocalaphis magnoliae (Essig and Kuwana) All adult viviparae alate, pale
shiny straw yellow to yellow-green with antennae and bases of tibiae banded
with black, and forewing veins conspicuously black-bordered; BL 1.4-2.2 mm.
On leaves of Magnolia kobus and M. liliflora in Japan and Korea. 2n = 20
(Blackman, 1986).
Neocalaphis magnolicolens (Takahashi) All adult viviparae alate, almost
translucent yellowish-white to very pale green; BL 2.3-3.3 mm. On undersides
of leaves of Magnolia obovata in Japan. Immatures feed alongside main veins.
2n = 20 (Blackman, 1986).
Two East Asian species feeding on twigs and branches of Betulaceae. They
are stout-bodied aphids with pigmented wings, long dorsal abdominal hairs
and a very short processus terminalis. Higuchi (1972) reviewed the Japanese
species. (A third, western palaearctic species on Ostrya, ostryae Börner, was
placed in this genus by Eastop and Hille Ris Lambers (1976), but is here
regarded as a Pterocallis.)
Neochromaphis carpinicola (Takahashi) All viviparae alate, stout-bodied,
dark brown with forewings almost wholly pigmented (see Moritsu, 1983);
BL 1.7-2.0 mm. On undersides of twigs and branches of Carpinus spp. in
Japan and Korea. Alate males and apterous oviparae in October-November
(Takahashi, 1923).
774 Neohormaphis
A genus for one species with prosoma distinctly separate from ABD TERG
2-7 and bearing numerous fine hairs.
Neophyllaphis viridis Ilharco Apterae dark green, covered with thick wax
powder; BL 1.5-1.8 mm. On Podocarpus falcata in southern Africa, found
in a mixed colony with N. grobleri (Ilharco, 1973; Dürr and Martin, 1976).
The alate vivipara was described by Russell (1982). Sexual morphs unknown.
One nearctic species with host alternation between Acer and Smilax.
A little-known genus of three species, two from basal and underground parts
of Salix in Europe, the third from Populus in China. No alate morph is
known, and in the one case where sexuales have been collected the males
appear to be paedogenetic.
About ten species resembling Symydobius but associated with nearctic oaks.
They usually feed on bark of terminal twigs, extending onto leaf petioles and
along basal parts of main veins. Remaudière (1982a) provided a key.
nigra and Q. cinerea in Florida (Tissot, 1932a) and Louisiana. Biology and
sexuales unknown.
NEOTHORACAPHIS Takahashi
Hormaphidinae: Nipponaphidini
About ten species mainly known from Quercus in East Asia. Apterae are
black, sclerotic, lack siphuncular pores and have much reduced tarsi, with the
fore and mid-legs concealed but the hind legs usually projecting a little behind
the body. Host alternation and gall formation on Distylium is only established
for one species (yanonis); several species seem to be anholocyclic on evergreen
Quercus, with the apterous vivipara the only known morph. Most species were
originally described in Thoracaphis. Takahashi (1958d) and A.K. Ghosh and
Raychaudhuri (1973a) gave accounts of the genus and keyed species under
Microthoracaphis, which is here included within Neothoracaphis. Tao (1966)
keyed the species in Taiwan.
dorsal wax and black hind legs projecting behind body; BL 0.5-0.6 mm. Alate
sexuparae return to Distylium in October-November and produce abundant
sexuales on undersides of leaves (Takahashi, 1958d). Shen (1986) studied the
life cycle and chemical control measures in China and Ito (1984) recorded
invasion of galls by the kleptoparasitic moth Nola innocua in Japan. Neo-
thoracaphis hangzhouensis Zhang, described from China (Zhang and Zhong,
1982b), is not clearly distinct from N. yanonis and may be a synonym.
Two species in India, one on Eugenia and the other described from an
unnamed parasitic plant on a tree (Pal and Raychaudhuri in Raychaudhuri
et al., 1980c). Raha and Raychaudhuri (1981) provided a key.
species is similar to bulbicauda but the apterae have only 4 processes on ABD
TERG 7 and few ventral cephalic hairs (Quednau, pers. comm.).]
About 15 East Asian species, some with host alternation between galls
on Distylium and Fagaceae, others apparently anholocyclic on Fagaceae,
Lauraceae and Moraceae. Apterae on secondary hosts are aleyrodiform, with
the strongly sclerotized prosoma bearing pustulate ornamentation. Alatae
Nipponaphis 785
Fig. 130. Galls of Nipponaphidini on Distylium. A, Dinipponaphis autumna (after Moritsu, 1983);
B, Neothoracaphis yanonis (after Paik, 1972); C, Monzenia globuli (after Moritsu, 1983);
D, Metanipponaphis cuspidatae (after Sorin, 1987b); E-G, (3 figs) various galls of Nipponaphis monzeni
(all from Sorin, 1958); H, N. litseae (after Sorin, 1987b); I, N. ?machili (after Sorin, 1987b);
J, Schizoneuraphis gallarum (after Docters van Leeuwen-Reijnvaan and Docters van Leeuwen, 1926).
A-l, all on Distylium racemosum; J, on D. stellare.
786 Nipponaphis
Nipponaphis ficicola Hille Ris Lambers and Takahashi Apterae very broadly
oval, wrinkled, dark purple to blackish in life; BL 1.0-1.3 mm. On the bark
of about 1 cm diameter branches and green twigs of Ficus benjamina and Ficus
sp. in Java (Hille Ris Lambers and Takahashi, 1959). Apterae collected in
June-November. Life cycle unknown.
About five species forming galls on Rhus in East Asia and migrating in
autumn to unknown secondary hosts, likely to be mosses. Tsai and Tang
(1946) provided a review of the Chinese species and Tao (1970) keyed the
species in Taiwan (although his synonymy is not followed here). See also B
& E, 1984, pp. 188-189.
Nurudea ibofushi Matsumura (= sinica Tsai and Tang) Galls pale yellowish-
brown with a pinkish tinge, irregular in shape, max. diameter 4.5-5.5 cm; on
leaves of Rhus semialata in China and Japan. Alatae (BL c. 1 mm) with brown
head and thorax and pale yellow-green or yellow-brown abdomen emerge in
autumn (Tsai and Tang, 1946; as N. sinica).
Nurudea meitanensis (Tsai and Tang) Galls are rosy-red, somewhat flat-
tened, irregularly branched from the base, maximal dimension up to 10cm,
arising from leaflets of Rhus punjabensis var. sinica in China (Kweichow).
Alatae emerging from the gall in autumn have black head, pale green pro-
thorax, dusky pterothorax and pale yellowish abdomen (Tsai and Tang, 1946;
as Floraphis meitanensis). A closely-related species, N. choui Xiang, produc-
ing branched galls on Rh. potaninii in Shansi Province, differs in colour and
shape of gall, and the emergent alatae have longer ANT III and dense long
hairs on hind tibiae (Xiang, 1980/81).
One species with 1-segmented tarsi, forming galls on leaves of Japanese elm.
About ten species mostly host-alternating between Populus and roots of Picea.
The fundatrices lack wax glands and the alate spring migrants have fore-
wings with once-branched media. Accounts are available from Heie (1980;
Fennoscandia and Denmark), Danielsson (1976, gall generations in Sweden;
and 1990a, root-feeding generations), Carter and Danielsson (1991, root-
feeding generations in Britain) and Smith (1974a; North America). Asiphum
is a synonym.
One species closely related to Pachypappa, but the fundatrix has wax glands
on the thorax and abdomen.
Large distinctive aphids with short antennae living on Juglans regia (see B &
E, 1984, p. 248, as Callaphis). The three described taxa are closely related
and could perhaps be geographic variants of a single species. Previously the
name Callaphis Walker was widely used for this genus, but was suppressed
following a plea by Quednau (1983), on the grounds of potential confusion
with Calaphis Walsh.
in Nepal and Tibet. Sexuales are unrecorded. Zhang in Zhang and Zhong
(1985c) described a subspecies, P. nepalensis yunnanensis, from Yunnan
Province, China, also collected on Juglans regia.
Paoliella chiangae van Harten and 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 and Ilharco, 1972). Living singly or in
small colonies on undersides of young leaves of Combretum sp. (?psidioides)
in Angola.
Paoliella nachensis (Eastop) All viviparae alate, head and thorax dark,
abdomen yellow with dark dorsal processes; BL 1.1-1.4 mm. Living singly
on undersides of leaves of Combretum spp. (incl. apiculatum, ?psidioides).
Recorded from Angola, Burundi and Tanzania. Life cycle unknown.
Paoliella papillata (Hall) Plate 7f All viviparae alate, very dark brown; BL
1.4-1.5 mm. Immatures pale reddish-brown with white dorsal processes (Hall,
1932). On Combretum spp. (incl. apiculatum, molle), widely distributed in
Africa south of the Sahara. Doncaster (1954) described the alate male and
apterous oviparae from amongst the type material, collected in Zimbabwe in
December.
796 Paracletus
Paoliella terminaliae (Hall) All viviparae alate, brick red to very dark but
with tip of abdomen always pale; BL c. 1.5 mm (Hall, 1932). On Terminalia
sericea in southern Africa (Angola, Malawi, South Africa, Zimbabwe). Life
cycle unknown. Eastop (1956) described kenyensis as a subspecies, from a
single alata trapped in Kenya.
Paoliella wettsteini Quednau Apterae pale yellow with white dorsal pro-
cesses; BL 1.0-1.2 mm. Alatae very pale with head and thorax only slightly
dusky. On leaves of Combretum gueinzii (= C. molle) in South Africa
(Quednau, 1964). Life cycle unknown.
About 4 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), but the gall-living generations of this species
are unknown.
Two species in East Asia closely related to Colopha, but the media of the
forewing of alatae from galls is unbranched and the apterae on the secondary
hosts (bamboos) always have siphunculi. The life cycle of only one species is
known. Akimoto (1985a) gave a full account.
One species in New Zealand, resembling Aphis but lacking lateral tubercles
on ABD TERG 7, and often also lacking them on ABD TERG 1.
PARANIPPONAPHIS Takahashi
Hormaphidinae: Nipponaphidini
Three species, two nearctic and one palaearctic. The nearctic species have
host alternation between Acer and Alnus; the other lives on Alnus and pro-
duces sexuparae in autumn, but its primary host generations are unknown.
Remaudière and Muñoz Viveros (1985a) summarized differences between the
species.
PARATHORACAPHIS Takahashi
Hormaphidinae: Nipponaphidini
Five species in East and Southeast Asia differing from Thoracaphis in the
complete fusion of prosoma and abdominal plate, different cuticular orna-
mentation, spine-like marginal hairs and absence of siphunculi. Four species
are on Fagaceae; the host of the fifth is only tentatively recorded as Myrica.
Only apterous viviparae are known. Some of the species may have unrecog-
nized gall-forming generations on Distylium.
Patchia winforii Miller Apterae dark brown to black; BL c. 1.7 mm. Alatae
have black dorsal abdominal cross-bands and thickly-bordered wing veins.
Forming dense clusters on twigs and leaf petioles of Quercus palustris in
Massachusetts, USA (Miller, 1933b). Life cycle unknown. Not recorded since
the original description.
Over 70 species have been described, of which 46 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 17 of these gall-
formers are known; four are monoecious on Populus, the alate sexuparae
being produced in the galls, and in the other 13 species the alate progeny of
the fundatrix migrate to various herbaceous secondary hosts (in Cruciferae,
Compositae, Chenopodiaceae, Euphorbiaceae, Ranunculaceae and Umbelli-
ferae), 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. The species are distributed rather evenly
throughout the northern hemisphere (see B & E, 1984, p. 324). Accounts are
available for Britain (Furk and Prior, 1975); European gall-makers on
Populus nigra (Lampel, 1960); Fennoscandia and Denmark (Heie, 1980);
India (Ghosh, 1984b); eastern Russia (Pashchenko, 1988b); Japan (Aoki,
1975); China (Zhang and Zhong, 1979, 1983b); and North America (Palmer,
1952; Harper, 1959a, b; Lange, 1965; Smith, 1985).
Fig. 131. Galls on Populus. A, Epipemphigus sanpopuli on P. cathayanae (after Zhang and Zhong,
1983b); B, E. imaicus on P. ciliata (after Habib and Ghani, 1970); C, Gootiella tremulae on P. tremula
(after Danielsson, 1976); D, Mordwilkoja vagabunda on P. deltoides (after Britton, 1923); E, Pachypappa
marsupialis on P. nigra (after Börner and Heinze, 1957); F, Pachypappa populi on P. tremula (after
Danielsson, 1976); G, Pachypappa tremulae on P. tremula (after Tullgren, 1909); H, Pachypappella
lactea on P. tremula, with colony overflowing onto outside of gall (after Danielsson, 1976); I, Thecabius
affinis (fundatrix gall) on P. nigra (after Börner and Heinze, 1957); J, Th. lysimachiae on P. nigra (after
Börner and Heinze, 1957); K, Pemphigus borealis on P. laurifolia (after Tullgren, 1909); L, Pemphigus
bursarius on P. nigra (after Börner and Heinze, 1957); M, Pemphigus immunis on Populus sp. (after
Theobald, 1915); N, Pemphigus monophagus on P. angustifolia (after Aoki, 1975); 0, Pemphigus passeki
on P. nigra (after Prinsen, 1990); P, Pemphigus populinigrae on P. nigra (after Börner and Heinze,
1957); Q, Pemphigus populitransversus on P. deltoides (after Palmer, 1952); R, Pemphigus protospirae
on P. nigra (after Danielsson, 1976); S, Pemphigus spyrothecae on P. nigra (after Danielsson, 1976);
T, Pemphigus vesicarius on P. nigra (after Lichtenstein, 1855).
804 Pemphigus
extended period from late May to September, with peak emergence in late
June-July (northern hemisphere). They found colonies mainly on roots of
Compositae (except for populations in Australia and New Zealand which are
mainly recorded from Chenopodiaceae and Polygonaceae, and are perhaps a
separate species). Return migration of sexuparae to poplar is in September-
October, but anholocyclic overwintering on roots of secondary host is com-
mon. In Europe, northwest, southwest and Central Asia, Australia and
New Zealand, North and southern Africa, North and South America. Dunn
(1960a) studied gall formation and (1960b) reviewed natural enemies. Alleyne
and Morrison (1978a, b) gave accounts of the bology and natural enemies in
Quebec. See also B & E, 1984, p. 325. 2n = 20.
Pemphigus circellatus Zhang and Zhong Large galls are formed on the
branches of Populus tremula var. davidiana in China (Yunnan). Emigrant
alatae (BL c. 1.8mm) were collected in May (Zhang and Zhong, 1985a).
Life cycle unknown.
Pemphigus cylindricus Zhang Only fundatrices are described, from a gall
on an unidentified Populus sp. in Tibet (Zhang and Zhong, 1981b).
Pemphigus dorocola Matsumura The galls on twigs of Populus maximo-
wiczii in Japan are pouch-shaped, reddish when mature, with a transverse
apical slit (Aoki, 1975). Also recorded from P. koreana and P. suaveolens.
Fundatrix green; alatae (BL 1.9-2.3 mm) emerge over an extended period
(late July-November). Secondary host unknown. In Japan, Korea and Siberia
(Pashchenko, 1988b). Aoki (1978, 1980a) described aggressive and gall-
cleaning behaviour by first instar larvae, and Aoki and 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 recognize the presence of P. borealis in Japan.
Alatae in the BMNH colln from galls on Populus maximowiczii 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 (Blackman, 1986).
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). Fundatrix dull pale
green. Alatae (BL 1.5-2.2 mm) emerge in June-July and migrate to colonize
roots of Aethusa cynapium (Stroyan, 1964b). Sexuparae return in October.
Only recognized from England, but probably more widely distributed in
Europe.
Pemphigus 805
Pemphigus populi Courchet Galls on Populus 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. Alatae (BL 1.4-2.5 mm) emerge in
June-July through a secondary opening at the top of the gall, migrating to
found colonies on roots of Melilotus altissima and possibly other Leguminosae
(Furk and Prior, 1975). Sexuparae return to P. nigra in September-October.
In Europe, southwest and Central Asia, and east to China (Chang and Zhong,
1979c). Furk and Prior (1975) reported parasitization of fundatrices by
Monoctonia pistaciaecola.
Pemphigus populiglobuli Fitch 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 and Knowlton, 1929).
Primary hosts are Populus angustifolia and P. balsamifera, more rarely
P. trichocarpa. Fundatrix is variously reported to be dark green (Palmer,
1952) or pale yellowish-white (Maxson, 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 and Knowlton
(1929).
green. Alatae (BL 1.1-1.8 mm) emerge in June-July and migrate to found
colonies on roots of Chenopodiaceae. The evidence for this host alterna-
tion is work in California by Grigarick and Lange (1962), who transferred
sexuparae from sugarbeet 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 colonized roots
of B. vulgaris and Chenopodium album. However, the suggested synonymy
of both P. betae and P. balsamiferae with populivenae indicates some confu-
sion of identity and further work seems advisable to confirm that the correct
name for populations damaging sugarbeet in California (e.g. Summers and
Newton, 1989) is in fact populivenae rather than betae. There may be a
complex of species with different Populus spp. as primary hosts. Whitham
(1978) reported that P. populivenae galling Populus fremontii in Utah caused
chlorosis of the leaf distal to the gall.
Periphyllus aceris (Linnaeus) Apterae yellow, often with green flecks dor-
sally; 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
814 Periphyllus
Periphyllus aesculi Hille Ris Lambers Apterae rather elongate oval, brownish
in life with black dorsal spots (A.K. Ghosh, 1980); BL 2.5-3.5 mm. Siphunculi
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 (BMNH colln, leg. CIBC).
Chakrabarti et al. (1972a) described the apterous vivipara and redescribed the
alate vivipara; L.K. Ghosh (1986) described the ovipara; and Agarwala and
D.N. Raychaudhuri (1982) described the alate male. Aestivating morph not
recorded. Sexuales in October-December.
Periphyllus brevispinosus Gillette and Palmer Apterae brown with dark and
pale mottling, and pale siphunculi; BL 2.4-2.9 mm. Alatae brown with black
dorsal abdominal cross-bands and marginal sclerites; on undersides of leaves
of Acer glabrum in western North America. Aestivating nymphs have foliate
marginal hairs with serrate margins (Essig and Abernathy, 1952). Monoecious
holocyclic; oviparae in late September (Palmer, 1952).
Periphyllus bulgaricus Tashev Apterae shining black, with pale legs and
antennae; BL 2.0-2.8 mm. Apterae and alatae both have broad black dorsal
abdominal cross-bars. On shoot tips and undersides of leaves, usually at
base of mid-rib. Described from Acer hyrcanum in Bulgaria (Tashev, 1964);
also collected from A. hyrcanum in Turkey, from A. opulifolium in Iran,
from A. hermaneum in Lebanon (BMNH colln, leg. DHRL) and from
A. granatense and A. monspessulanum in Spain. Monoecious holocyclic;
Nieto Nafria and Gonzales Funes (1990) described fundatrices, oviparae and
alate males and discussed the very difficult separation of this species from
P. rhenanus. Aestivating nymphs not recorded.
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 green, covered with
dirty white wax wool; BL 1.2-2.2 mm. On bark and in crevices on trunks of
Populus spp. (alba, ciliata, x euroamericana, maximowiczii, nigra, suaveolens,
tadzhikistanica, tremuloides), heaviest infestations occurring on six to eight-
year-old trees. Monoecious holocyclic; oviparae and males are produced in
September-October in the northern hemisphere, the oviparae each laying two
eggs (Theobald, 1929). Anholocyclic overwintering by apterous viviparae is
also common and may predominate; no morph recognizable as a fundatrix
has been described. In Europe, North Africa (Egypt, Morocco), southwest,
Central and East Asia (incl. Japan; BMNH colln, leg. S. Aoki); USA (only
record is from Maine) and South America (Argentina, Chile, Colombia).
Arzone and 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 Doom and Hille Ris Lambers, 1962), as also were
large specimens found on P. tremuloides in Maine (as Ph. dearborni; Smith,
Phyllaphis 821
1974b). Zhang (in Zhang and Zhong, 1982b) described a subspecies in China,
Ph. passerinii zhangwuensis. It seems likely, however, that a single, somewhat
variable species is involved. 2n = 10.
Phyllaphis fagi (Linnaeus), Plate 8e, f Apterae elongate oval, pale yellowish-
green, covered with wax wool; 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, and introduced to Australia, New Zealand and North America.
Records from Japan apply to Ph. fagifoliae. Monoecious holocyclic, with
dwarf apterae living through summer months to produce apterous oviparae
and alate males in October-November. Oviparae have dark dorsal cross-
bars and dark siphunculi (cf. Ph. grandifoliae). Chandler (1968) studied its
association with a syrphid predator. 2n = 16.
with fleshy spines on Carya sp. in Illinois (the immature inhabitants of these
galls were described by Shimer as having long spiny tubercles); caryaemagna,
producing velvety galls on petioles or at base of mid-rib on C. amara
(= cordiformis) in Illinois and Missouri; and subelliptica, producing ellipsoid
mottled green galls on side of twig of C. alba ( = ovata) in Illinois (see also
Pergande, 1904). Caldwell and Schuder (1979) studied the life cycle of the
form on C. ovata in Indiana. Shimer (1869) reported on natural enemies of
galls of caryaemagna in Illinois. 2n = 8.
Phylloxera caryaefallax Riley Galls on leaves of Carya alba (= ovata) are
thin-walled, 1-5 mm diameter, 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 Tennessee. Life cycle 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.] 2n = 12 (Steffan, 1968).
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 northeastern
USA (New York, Illinois, Iowa, Virginia). Life cycle unknown. 2n = 8
(Steffan, 1968).
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 hemispherica Shimer might be a
synonym. Life cycle unknown. 2n = 22 (Steffan, 1968).
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 undescribed and life cycle 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 trans-
verse 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,
Phylloxera 825
his name spinifera is available for this form should it prove to be specifically
distinct.
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 diam. Alatae (BL 0.8-1.0 mm, with
orange abdomen) differ from Ph. devastatrix in characters of the wings and
antennae (see Pergande, 1904).
Phylloxera rileyi Riley Apterae dark brown with dorsal tubercles almost
black (but paler forms may 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 monoecious holocyclic; alatae were pro-
Phylloxera 831
duced 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).
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, DC, USA. Life cycle 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.
About seven species on Salicaceae, mostly living in dense white wax. The
species on Salix in Germany were studied by Iglisch (1965).
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. Monoe-
cious holocyclic on N. sylvatica. In Maryland and District of Columbia, USA
(Pergande, 1904).
Phylloxerina sp. Holman (1974) described (but did not name) a species of
Phylloxerina on trunks of Salix longipes in Cuba. Life cycle unknown. Similar
to European Ph. capreae (but not compared with the North American species).
834 Pineus
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 and Heinze, 1957; Heinze, 1962), India (A.K. Ghosh, 1984a),
Japan (Inouye, 1953) and North America (Annand, 1928).
Pineus abietinus Underwood and Balch Only the first instar and adult
progrediens (aestivoprogrediens) are described; adults are very small (about
0.6mm), 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 northwest 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 and Balch, 1964).
Pineus boycei Annand 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 or 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).
Fig. 132. Galls of Pineus on Picea. A, Pineus cembrae on Picea abies (after Francke-Grosmann, 1938);
B, P. orientalis on Picea orientalis (after Carter, 1976); C, P. pinifoliae on Picea mariana (after Patch,
1910a; open gall on left, closed gall on right).
Deboo and Weidhaas (1967) believed that gallicolae were recolonizing spruce
directly and not migrating to pine.
Pineus hosoyai Inouye Only the apterous progrediens and immature sex-
upara 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. Life cycle is unknown.
One species on Fagus in Japan. Only the very specialized apterous vivipara
is known, and its phylogenetic position is uncertain. Takahashi (1957) com-
pared it with Dasyaphis, but Shaposhnikov and Gabrid (1987) suggested that
it belongs in the Hormaphidinae.
Platyaphis fagi Takahashi Apterae 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
unknown.
(1966b) revised the genus (which in his concept included Fullawaya), and Heie
(1986) reviewed the European species.
Plocamaphis martini Richards Apterae light brown to pink (no wax men-
tioned); BL 2.8-3.5 mm. On roots of Salix sp(p). in arctic and subarctic
Canada (Baffin Is., Quebec). Monoecious holocyclic; apterous males and
oviparae occur in mid-August (Richards, 1963). Possibly conspecific with
P. amerinae.
About 50 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 trans-
ferred their sexual generation in relatively recent times from Populus to
842 Prociphilus
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 cornifoliae Singh, Das and 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 unknown.
apterae and alatae) live in curled and clumped leaves at tips of twigs through-
out the growing season. 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), Mexico and introduced to Chile (Carrillo,
1977) and South Africa (on planted F. pennsylvanica and F. velutina; Müller
and Scholl, 1958, and BMNH colln). Carver (1980a) described an aphelinid
parasitoid on P. fraxinifolii in Iowa; Brundrett and Kendrick (1987) studied
symbiosis between root-feeding populations and a bolete fungus. 2n = 20
(Robinson and Chen, 1969) or 2n = 22*.
according to Mordvilko (1935). Hori (1938) also noted that the secondary host
'may probably be Rumex'. Sexuparae occur in October-November, but have
not been described. In Japan, Korea and eastern Siberia.
Prociphilus oleae Leach ex Risso Fundatrices and their progeny form com-
pact spring colonies covered in white wax wool on shoots near base of
trunk of Olea europea. Also recorded from Phillyrea media. Unusually for
a Prociphilus, no leaf deformation is caused. Progeny of fundatrix all alate,
BL c. 4 mm with smoky yellowish abdomen, migrating in May. The life
cycle is unknown, but apterae described from roots of Vitis vinifera in Sicily
(Barbagallo and Stroyan, 1982) could be the secondary host form of this
species. In Mediterranean area (France, Italy, Greece, Turkey). Roberti and
Monaco (1987) give a recent account.
About ten species mostly on roots of Compositae (see B & E, 1984, p. 332).
Two poplar-feeding species only known from Canada and northwest USA,
related to Chaitophorus but with a distinctive, long, thick processus terminalis
clothed with hairs like the rest of the antenna, and pore-like siphunculi.
Richards (1972a) provided a key.
Fourteen species of small pale aphids with a short antennal terminal process
living on Betulaceae, especially Alnus. They are usually dispersed on under-
sides of leaves and are mostly not attended by ants. Apterous viviparae occur
more commonly than in most Drepanosiphinae, particularly in Pterocallis
s. str. which has 4 species in North America, 3 in Europe and 3 in East Asia.
The subgenera Paratinocallis (1 sp.) and Recticallis (3 spp.) are wholly East
Asian. Mesocallis is here considered as a separate genus. Accounts are available
for Britain (Stroyan, 1977), Fennoscandia and Denmark (Heie, 1982), Central
Europe (Börner, 1949), Japan (Higuchi, 1972), India (Chakrabarti, 1988;
A.K. Ghosh and Quednau, 1990), USA (Granovsky, 1928a; as Myzocallis)
and Canada (Richards, 1965).
Pterocallis affinis Chakrabarti Alatae pale yellow, with antennae and legs
mainly pale; BL 1.5-2.0 mm. On Corylus colurna in Uttar Pradesh, India. No
Pterocallis 855
Pterocallis alni (De Geer) Apterae and alatae are yellowish-white to yellowish-
green, with antennae ringed with black, a black spot near apex of hind femur
and black tarsi; BL 1.3-2.0 mm. Almost exclusively on Alnus glutinosa in
Europe, living dispersed on undersides of leaves and only rarely attended by
ants. Common and widespread in Europe, east to Turkey, Iran and Lebanon;
introduced to North America (Essig, 1912b), New Zealand (Lowe, 1968),
Australia (Carver, 1980a) and Chile (Carrillo, 1980). In North America it
seems able to feed on various native Alnus spp. (BMNH colln). South-
west Asian populations colonize A. orientalis and A. subcordata as well as
A. glutinosa and tend to be intermediate in morphology between P. alni and
P. maculata, so their specific identity is often uncertain. Monoecious holo-
cyclic, with oviparae and alate males in September. Gange and Pryse (1990)
investigated the effects of temperature and nutrition on development, and
Gange and Llewellyn (1988) studied egg distribution and mortality. Pterocallis
alni on alder windbreaks may be a useful alternative food for predators of
orchard pests (Gange and Llewellyn, 1989). 2n = 20.
Pterocallis alnifoliae (Fitch) Apterae pale green, pale yellow or whitish with
darker green irregular patches (Granovsky, 1928a); BL 1.1-1.5 mm. Alatae
have pale greenish-yellow abdomen and darker yellow to orange head and
thorax; BL 1.3-1.7 mm. On undersides of leaves of Alnus rugosa and A.
serrulata in eastern North America. Monoecious holocyclic; oviparae and
alate males in September. [Granovsky (1928a) described a very similar but
smaller species, A. rhombifoliae, from Alnus rhombifolia in California and
also from A. incana (= rugosa) in Wisconsin. Collections from Maine on
Alnus sp. (BMNH colln, leg. H.L.G. Stroyan) and Quebec on A. rugosa
(BMNH colln, leg. DHRL) seem to be this species. However, it is not clear
how apterae of rhombifoliae can be distinguished from those of alnifoliae.
According to F.W. Quednau (pers. comm.), P. alnifoliae has only alate
viviparae and all apterae are rhombifoliae. Further work is needed to clarify
the situation; in our key to Alnus-feeders, both species are therefore taken out
in two places, but discriminated only on the basis of characters of alatae.]
brown-black, forewing veins narrowly bordered with fuscous, dark hind legs,
dark finger-like spinal tubercle on ABD TERG 1, dark marginal spots on ABD
TERG 3-4 and black distal halves to siphunculi; BL 1.6-2.1 mm. On under-
sides of leaves of Alnus formosana in Taiwan, where it reproduces partheno-
genetically throughout the year (Takahashi, 1923; as Myzocallis pseudoalni).
Takahashi (1924a) provided detailed descriptions of all immature stages.
Granovsky (1928a) provided an illustrated redescription.
Pterocomma italica Barbagallo and Stroyan Apterae very dark brown, with
a fine spinal line of whitish wax crossed by two transverse wax bands on about
ABD TERG 1 and 5 and orange-yellow siphunculi; BL 2.3-3.4 mm. Alatae
with variable dorsal abdominal markings, not forming cross-bands. On twigs
or at base of stems of Salix purpurea, in mountainous areas of Italy and
Switzerland (Barbagallo and Stroyan, 1978). Sexual morphs and life cycle
unknown. Very similar to P. jacksoni. Heavy parasitization by an Aphidius
species is reported in the original description.
Pterocomma 859
Mongolia. Introduced to North America (see Smith and Parron, 1978; Forbes
and Chan, 1986a) and South America (Argentina; Chiesa Molinari, 1942, and
Chile; BMNH colln, leg. E. Zuniga). Monoecious holocyclic; oviparae and
alate males occur in October-November. Martin-Bernal and Padro Simarro
(1986) reviewed natural enemies. Börner (1950) described a form, dubium,
from P. alba in Germany, resembling populeum but differing in colour (silver
grey) and absence of marginal tubercles; Holman and Szelegiewicz (1974a)
reported similar aphids (as P. populeum dubium) from Mongolia, collected
on P. laurifolia. [There are numerous records of this species from Salix, but
these are almost certainly due to confusion with other species (pilosum and
pilosum ssp. konoi in palaearctic; ?sanguiceps in North America.]
Two or three species of very large East Asian aphids living on woody
Rosaceae. The alatae have an elongate pterostigma as in Longistigma, but it
does not curve around the tip of the wing. A.K. Ghosh (1982b) reviewed the
Indian species.
One oak-feeding species with small, flattened aptera having an oddly irregular
shape. Alatae have an unbranched media in the forewing and a hind wing
without any oblique veins.
Apterae and alatae described from Tibet (Zhang and Zhong, 198Ib) under the
name Radisectaphis gyirongensis appear to belong to a species of Pterocom-
matini. The recorded host plant is Malus pumila; presumably either this was
an error, or the aphids were vagrants.
864 Rectinasus
Probably only one species, with distinctive antennal sensoriation in alatae and
a long R IV+V. Affinities with other genera of Fordini were discussed by
Remaudière and Davatchi (1956).
Three East Asian species having aleyrodiform apterae with heavily sclerotized
and reticulate dorsal cuticle. On Ficus or Fagaceae. At least one species forms
galls on Distylium, but the taxonomy and life cycle needs further investigation.
A.K. Ghosh and Raychaudhuri (1973b) reviewed the genus.
Reticulaphis distylii (van der Goot) Galls on Distylium stellare in Java are
conical or fingertip-like, brownish-green when mature (Hille Ris Lambers and
Takahashi, 1959). The gall illustrated (as Schizoneuraphis distylii van der
Goot) by Docters van Leeuwen-Reijnvaan and Docters van Leeuwen (1926)
seems to be another species (see Noordam, 1991, p. 259). Alatae (BL 1.5-
1.9mm), secondary rhinaria distributed III 19-31, IV 8-12, V 6-10, abdomen
dark green) are present in the galls in October. Host alternation apparently
occurs to Ficus spp. (Hille Ris Lambers and Takahashi, 1959), but this needs
to be confirmed experimentally; very similar aphids, identified as subspecies
rotifera, have been collected from Quercus sp. in India (Raychaudhuri et al.,
1980c). Apterae on Ficus are dark brown to dull purplish-black, acquiring with
age a fringe of wax; BL 0.5-0.9 mm. Immature apterae are yellow-brown and
active. Immature alatae on Ficus have a woolly wax fringe plus long erect wax
Rhopalosiphoninus 865
filaments; when adult they are dark violet-black, without wax. Zimmerman
(1948) provided a well-illustrated account of this species in Hawaii. Hille Ris
Lambers and Takahashi (1959) recognized several 'subspecies' on Ficus in
Taiwan and Java and provided a key, noting that many of the differences
could prove to be within the range of variation of one species. The most
commonly encountered form on Ficus, R. distylii fici (Takahashi), has
marginal hairs about 75 µm long with acute apices, and is recorded from
China, Taiwan, Japan, Java, Nepal (BMNH colln, leg. K.C. Sharma) and the
Philippines. Another fairly distinct form, R. distylii foveolatae, is smaller and
narrower (BL c. 0.5 mm) and has no evident wax secretion; it appears specific
to Ficus foveolata in mountainous regions of Taiwan and Japan (Takahashi
1935c, 1958a).
About 20 species with large and strongly swollen siphunculi and a short
triangular CAUDA, living on a great variety of plants, often in cryptic
habitats. Three species are recorded from trees; at least two of these have host
alternation to herbaceous plants. See also B & E, 1984, p. 337.
through summer on Staphylea, are much more pigmented than the spring
generations; green, with variably-developed olive green to black dorsal
abdominal cross-bands. The holocycle is only recorded from Europe, but
anholocyclic populations of staphyleae group are almost cosmopolitan (see
B & E, 1984, p. 339). 2n = 10.
Malus, Pyrus, Sorbus and sometimes on Cydonia and Mespilus). It may also
occasionally form colonies on suckers of Prunus cerasus. Heteroecious
holocyclic; alatae fly in late May-June to found colonies on subterranean
parts of various Gramineae, but colonies may persist into summer on primary
hosts. In Europe, eastward to the Urals and Turkey; the Azores; possibly
originating from North America (where for many years the name fitchii was
used for insert m on its primary hosts). Evenhuis (1968) studied its natural
control and the role of a parasitoid in apple orchards. See also B & E, 1984,
p. 340. 2n = 10.
One (or two) palaearctic species, resembling Nearctaphis but without lateral
abdominal tubercles, and the primary host is Prunus rather than Pomoidea.
Roepkea marchali (Börner) Apterae a dirty yellowish-green to almost black,
according to degree of dorsal sclerotization; BL 2.0-2.3 mm. Alatae have
Sanbornia 869
One species in Japan, resembling Aphis in its general appearance and arrange-
ment of marginal tubercles, but with numerous marginal hairs on abdominal
segments; with somewhat developed antennal tubercles, giving the front of the
head a concave outline in dorsal view; and with secondary rhinaria on ANT
III (and sometimes IV) in apterae.
A small oriental genus with Pyrus as primary hosts, related to Dysaphis but
with long hairs on body and antennae. Accounts are available for Japan
(Miyazaki, 1971) and China (Zhang and Zhong, 1983b). The name Sappaphis
was used in a broader sense from 1952 to 1965 to include species now in
Dysaphis.
non-native pines, in western North America. Oviparae and alate males occur
in October (Hottes et al., 1954). The alata was described by Hottes (1956d).
Laviolette, 1968b), predators (Gagné and Martin, 1968) and fungal diseases
(Gröbler et al., 1962; Tyrrell and Macleod, 1975).
Six species are known, at least one of them having host alternation between
Distylium galls and Litsea (in Java; Noordam, 1991). Closely related to
Nipponaphis, but the aleyrodiform exules have long, basally thick dorsal
(prosomal) hairs.
Schizoneurata tissoti Hille Ris Lambers Only immatures and alate sexuparae
are described, and appearance in life is unknown. Apterae and immatures
are presumably wax-covered. Sexuparae have BL 1.6-2.2 mm and secondary
rhinaria distributed III 15-20, IV 2-5, V 1-3, VI 0-1. On Crataegus uniflora
and Crataegus sp. in Florida, USA (Hille Ris Lambers, 1973c).
One species in the Himalayan region, with empodial hairs reduced (in
alatae) or absent (in apterous exules); related to Eriosoma but without
siphunculi in the morphs on the secondary host plant, and distinct from
the nearctic genus Schizoneurata because it has embryos with spinulose
hind tarsi, as in palaearctic Eriosoma (Schizoneura) (Hille Ris Lambers,
1973b).
Schlechtendalia 875
Two or three East Asian species related to Melaphis, but alatae emerging from
galls on Rhus have an elongate, curved pterostigma.
Fig. 133. Gall of Schlechtendalia chinensis on Rhus semialata (after Baker, 1917b).
876 Schoutedenia
Six Asiatic species mainly associated with Celtis, with one described from
Tilia. Quednau and Remaudière (1985) keyed the species then known, and
A.K. Ghosh and Quednau (1990) reviewed the Indian species.
Shivaphis celti Das, Plate l0c, d Apterae pale to dusky greenish, sometimes
pinkish, covered in white flocculent wax; BL 1.9-2.6 mm. Alatae also wax-
covered, with forewing veins thickly bordered with fuscous distally and
large black patches on pterostigma and distal part of Cu lb . On undersides
of leaves or on shoot tips of Celtis spp. in Turkey, India, Pakistan, Nepal,
Sri Lanka, Korea, Japan, Loochoo, China, Taiwan and Philippines. Mono-
ecious holocyclic; Das (1918) gave a full account of its biology and natural
enemies in Pakistan; oviparae and alate males occurred in December, eggs
were laid in January and hatched in March. In Japan, Shibata (1955; as
Sh. celticolens) obtained sexuales in September-November. In Taiwan,
Sh. celti is mainly anholocyclic, but produces some sexuales in December-
January. 2n = 10.
One species in southern Italy on Quercus, the apterae of which have remark-
able variation in form of dorsal hairs.
Sinolachnus 879
SINOCHAITOPHORUS Takahashi
Drepanosiphinae: Phyllaphidini
Six species with head ventrally spinulose and with weakly clavate, calf-shaped
siphunculi and a long CAUDA, living on trees and shrubs in East and South-
east Asia. Accounts are available for Japan (Miyazaki, 1971), China (Tao,
1963), northeast India (Raychaudhuri, 1980) and Java (Noordam, 1986).
Two species in East Asia in which the dorsum of the aleyrodiform aptera is
densely covered with hair-like processes.
with about 20 species recorded partly or entirely from trees. Sitobion are
distributed throughout the northern hemisphere and have also undergone an
adaptive radiation in Africa south of the Sahara. See also B & E, 1984,
pp.349-350.
Sitobion congolense Doncaster and Hille Ris Lambers Apterae mid- to dark
green; BL 2.2-2.5 mm. In West Africa, usually on Vernonia spp. (Com-
positae), forming colonies at growing points, but Remaudière and Autrique
(1985) recorded it from Harungana madagascariensis, both in Burundi and
Ivory Coast (cf. S. nigeriense). Parasitism by Aphidius sp. was observed.
Life cycle unknown.
Sitobion 883
Sitobion krahi (Eastop) Apterae green, with siphunculi pale only on basal
third and a pale CAUDA; BL 2.5-3.0 mm. Alatae have siphunculi wholly
black and dark marginal abdominal sclerites, a small antesiphuncular sclerite
and a large postsiphuncular sclerite. Collected on growing points of saplings of
a forest tree, probably Rauwolfia vomitoria but possibly Conopharyngea
chippii, in Ghana (Eastop, 1959). Specimens trapped or on unidentified hosts
have been collected in Nigeria, Sierra Leone and Tanzania (BMNH colln), and
Remaudière and Autrique (1985) recorded krahi from Mangifera indica in
Burundi. Life cycle unknown.
Sitobion nigeriense (Eastop) Apterae green with dark siphunculi and pale
CAUDA; BL 1.5-2.2 mm. Mostly recorded from Vernonia (Nigeria, Kenya,
Ethiopia; BMNH colln), but Remaudière and Autrique (1985) recorded it
twice colonizing Harungana madagascariensis in Burundi. Parasitism by a
?Trioxys sp. was observed. Life cycle unknown.
Sitobion phyllanthi (Takahashi) Apterae dirty olive green with dark brown
head and dorsal markings (Müller, 1959; as S. adgnatum); BL 1.5-2.1 mm.
Prepared specimens of apterae have variably developed narrow dorsal abdom-
inal cross-bands between dark marginal sclerites. Described from Phyllanthus
sp. in Mauritius (Takahashi, 1937c). Remaudière (1988) suggested that
5. adgnatum Müller, widely distributed on Euphorbiaceae in Africa, is this
species, and the BMNH collection contains specimens collected by A. van
Harten on Phyllanthus maderaspatensis in Angola. Life cycle unknown.
One species characterized by the elongate second antennal segment and the
thick sclerotic rims of the primary sensoria on the last two segments.
Smynthurodes betae Westwood Galls on Pistacia spp. (atlantica, mutica
and, rarely, vera) are yellow-green or red, spindle-shaped, about 20 mm long,
formed by rolling of the edge of the leaflet near its base (Fig. 126P, p. 700).
These are secondary galls, produced by the progeny of the fundatrix, which
lives in a small red mid-rib gall (Burstein and Wool, 1991). Heteroecious
holocyclic, with a two-year cycle; alatae (BL 1.3-1.6 mm) emerge in September-
November and migrate to roots of numerous, mostly dicotyledonous, plants.
The holocycle is recorded throughout the range of the primary hosts; Algeria,
Morocco, Israel, Syria, Iran, southern Crimea and Transcaucasus (Mordvilko,
886 Sorbaphis
One species related to Sappaphis, but with 4 instead of 3 hairs on 1st segments
of fore and mid-tarsi.
About six species on nearctic Quercus. Alate females apparently do not occur
and males may be apterous or alate within a species. Hille Ris Lambers (1966a)
keyed the species then known and discussed the confused nomenclature.
Stegophylla essigi Hille Ris Lambers Apterae are broadly oval, pale grey-
green to olive, covered in white wax wool; BL 1.2-1.8 mm. Feeding by
fundatrices in spring causes edges of young leaves to curl upward and
become thickened and red. Monoecious holocyclic on native Quercus spp.
in California, USA, with oviparae and both apterous and alate males in
November, but populations may continue parthenogenetically on evergreen
oaks such as Q. agrifolia, especially in leaves spun together by caterpillars
(Hille Ris Lambers, 1966d). 2n = 12.
Sumatraphis celti Takahashi Apterae dull green (in Sulawesi; J.H. Martin,
pers. comm.) or yellow to deep brown (in India; A.K. Ghosh et al., 1971e);
BL 1.5-1.7 mm. On new growth of Celtis tetrandra or ?Trema orientalis;
the apterae are flattened into angles of new shoots (J.H. Martin, pers.
comm.). In Indonesia (Sulawesi, Sumatra) and India (West Bengal). Alatae
of Sumatraphis have also been trapped in Nepal (BMNH colln, leg. K.C.
Sharma). The alate morph has not been properly described, and sexual morphs
and life cycle are unknown. The colour difference noted between Indonesian
and Indian specimens suggests that two species might be involved; the host
in India was unidentified.
Symydobius 891
Six species of medium to large, rather shiny dark brown aphids associated with
Betula and Alnus, forming colonies on 'the branches and twigs, invariably
attended by ants. A seventh species will shortly be described from Betula nana
in the Czech Republic (J. Holman, pers. comm.). All the apterae (even the
oviparae) are 'alatiform', with a similar pattern of sclerotization and pigmenta-
tion to that of the fully-winged adults. Monoecious holocyclic; males are
apterous, and oviparae have the posterior abdominal segments extended into
an ovipositor-like structure. Trioxys betulae is a specialized parasitoid.
Taiwanaphis atrovirens Noordam and Hille Ris Lambers Alatae have head,
pronotum and sides of abdominal segments 3-8 yellowish or brownish,
mesonotum and central part of dorsal abdomen dark green; BL 1.2-1.5 mm.
Immature alatae are yellowish-white with a dorsal greenish-black area.
Apterous viviparae, other morphs and life cycle unknown. On upper and
lower sides of new leaves and developing shoots of ?Syzygium sp. in Java
(Noordam and Hille Ris Lambers, 1985).
Hales (1974) described all morphs and gave life cycle details; monoecious
holocyclic, with oviparae and alate males occurring concurrently with alate
viviparae through spring, summer and autumn. Eggs are flat, colourless and
wax-covered, and are laid on leaves.
England, New Zealand and North America. Life cycle unknown in East Asia;
anholocyclic where introduced. Coffelt and Schultz (1990) studied its popula-
tion dynamics in USA. 2n = 18.
About 30 species in which the apterae have 1-segmented tarsi and the alatae
have a simple, unbranched media in the forewing and usually ANT IV much
shorter than ANT V. At least seven species have host alternation between leaf
galls on Ulmus and roots of Gramineae, and the lesser-known species are
described either from Ulmus or grass roots. The elm galls are invariably
stalked, pouch-like outgrowths on upper sides of leaves. Tetraneura seems to
be predominantly oriental and eastern palaearctic but T. ulmi appears to be
native to northwest Europe, T. caerulescens and T. africana are around the
Mediterranean and in southwest Asia. T. nigriabdominalis is probably east
Asian although holocyclic populations of this species are widespread. Hille Ris
Lambers (1970b) revised the world fauna and there are more recent accounts
from India (A.K. Ghosh, 1984b), China (Zhang et al., 1991) and Japan
(Akimoto, in prep.), but the genus is still in need of further revision. The
karyotype variation may be useful in this respect. (See also B & E, 1984,
pp. 358-361.)
Tetraneura africana van der Goot Galls on Ulmus have not been identified,
but Mordvilko (1935) recorded sexuparae collected on trunks of U. campestris
( = procera) in southern Italy.
with numerous granular protuberances and short white silky hairs, usually
crimson when mature. Alatae (BL 1.7-2.3 mm) emerge in June-July through
a lateral opening. Hille Ris Lambers (1970b) redescribed the gall generations
(as nigriabdominalis). The life cycle has not been fully investigated. 2n = 18
(S. Akimoto, pers. comm.).
Tetraneura paiki Hille Ris Lambers Described from a single alate sexupara
trapped in Korea (Hille Ris Lambers, 1970b). Moritsu's (1983) account and
illustrations of an aphid under this name in Japan should be referred to
T. nigriabdominalis (Akimoto, in prep.).
Tetraneura sorini Hille Ris Lambers Galls are stalked, hairy, pouch-
shaped, often somewhat globular, yellow and red (Moritsu, 1983). On Ulmus
spp. (laciniata, japonica, parvifolia) in Japan, Korea and China. Alatae
(BL 1.6-2.0 mm) emerge in June. Secondary host generations unknown.
2n = 16.
898 Tetraneura
Fig. 134. Galls on Ulmus. A, Olegia ulmifoliae on Ulmus japonica (after Pashchenko, 1988b); B, Colopha
compressa on Ulmus sp. (after Mordvilko, 1935); C, C. ulmicola on U. americana (after Hottes and
Frison, 1931); D, Tetraneura ulmi on Ulmus sp. (after Marchal, 1933); E, T. nigriabdominalis on Ulmus
sp. (after Moritsu, 1983, as T. paiki); F, T. chinensis on Ulmus sp. (after Mordvilko, 1935, as
Thecabius 899
upper side of the leaf becomes blistered and yellowish or reddish. These are
secondary galls containing only the alatiform progeny of the fundatrix, which
inhabits a small gall of its own formed by folding the edge of a leaf (Fig. 131I,
p. 803). Alatae (BL 2.2-3.1 mm, with green abdomen) leave the gall in late
June-July to found waxy colonies at stem bases and on runners of Ranunculus
spp. (Fig. 3). Mordvilko (1935) gave an account of the life cycle and described
all morphs. It occurs on a wide range of Populus spp. throughout Europe and
across Asia to eastern Russia, Japan and Korea. Populations on East Asian
Populus spp. are often treated as a separate species, Th. orientalis Mordvilko,
here regarded as a synonym. Karyotype is 2n = 38 in both England and Japan.
[In North America, Thecabius populiconduplifolius (Cowen) is virtually
indistinguishable from Th. affinis and has an identical biology, but with native
poplars as primary hosts (Palmer, 1952). Harper (1966) and Smith (1974a)
suggested synonymy, but Th. populiconduplifolius is not introduced Th.
affinis, as it has 2n = 28 (Harper and Macdonald, 1966); it is apparently a
good species.]
Four oak-feeding species, three in Europe and the Mediterranean region, and
one in North America. Remaudière (1982c) provided a key including charac-
ters of alatae as well as apterae, and Ilharco (1966) keyed first instar nymphs.
Thelaxes suberi (del Guercio), Plate 4a, b Apterae vary in colour from pale
green to dark brown; BL 0.9-1.5 mm. Living abundantly on young shoots,
leaves and developing acorns of numerous Quercus spp. (but not robur) in
England, southern Europe, Mediterranean region and southwest Asia. Also
recorded from Castanea sativa (in Italy). Presumably monoecious holocyclic,
but life cycle unstudied; Polaszek (1986) collected presumed sexuparae on
Q. cerris in England in November-December. 2n = 8.
Four oriental species very similar to Eucalliplerus, but with larger, pale
siphunculi and more extensive forewing pigmentation. Accounts are available
for Japan (Higuchi, 1972), China (Zhang and Zhong, 1982c) and Siberia
(Pashchenko, 1988b; Quednau and Shaposhnikov, 1988).
Tiliphagus 903
Tiliaphis shinjii Higuchi All viviparae alate, colour in life not recorded,
dorsal dark markings as in coreana but with paired dark spinopleural patches
on abdominal tergites; BL 2.0-2.7 mm. On Tilia spp. in Japan, Korea and
eastern Siberia. Sexual morphs in September (Higuchi, 1972; Quednau and
Shaposhnikov, 1988).
About 25 mostly small species usually associated with Ulmaceae, although the
three members of subgenus Sarucallis feed on Lythraceae (Lagerstroemia,
Duabanga), and species have also been described from Corylus, Dalbergia
and Sapindus. (Woody Leguminosae seem to be recorded quite regularly as
alternative or 'casual' hosts.) All viviparae are alate and usually have paired
spinal and marginal tubercular processes. Some species have conspicuous
black markings on dorsal body and/or forewings. Problems with the taxo-
nomy of Tinocallis may arise from seasonal variation; alatae in summer and
autumn are often more pigmented than in spring and may have fewer secon-
dary rhinaria. For example, specimens of Tinocallis platani collected in May
have 17-25 (usually 20-23) rhinaria on ANT III, whereas specimens collected
from the same trees in July-October have 7-21 (mostly 11-17) rhinaria on III
(VFE, unpublished data). Richards (1967) keyed the world fauna as then
known and more recent accounts are available for Britain (Stroyan, 1977),
Fennoscandia and Denmark (Heie, 1982), India (Chakrabarti, 1988; A.K.
Ghosh and Quednau, 1990), eastern Russia (Pashchenko, 1988b), China
(Zhang and Zhong, 1980c) and Japan (Higuchi, 1972).
Tinocallis nikkoensis Higuchi Alatae pale yellow with pale or dusky anten-
nae and legs; BL c. 1.5mm. On Corylus spp. (heterophylla, sieboldiana) in
Japan (Higuchi, 1972). Biology and sexual morphs unknown. [Paik's (1965)
record of T. zelkowae on Corylus sieboldiana in Korea should perhaps be
referred to this species.]
Tinocallis suzhouensis Zhang Alatae dusted all over with white powder; BL
c. 2.2cm. On Zelkova schneideriana in China (Zhang and Zhong, 1980c).
Sexual morphs and life cycle unknown.
Tinocallis 907
Tinocallis viridis (Takahashi) Alatae green, with green eyes, banded anten-
nae, forewings with dark spots on pterostigma and at distal ends of veins,
dark abdominal tubercles and dusky brown siphunculi; BL 2.2-2.4 mm. On
Zelkova formosana in Taiwan. Life cycle and sexual morphs unknown. [This
species has been confused in the literature with ulmiparvifoliae.]
One species on Prunus in northern India and Nepal. A.K. Ghosh and Quednau
(1990) gave a full account.
Tinocalloides montanus Basu All viviparae alate, with brown head and
thorax, pale yellow abdomen with two widely-separated rows of black spino-
pleural patches; BL 2.1-2.6 mm. On undersides of leaves of Prunus spp.,
especially P. cerasus, in northern India (West Bengal, Himachal Pradesh) and
Nepal. Holocyclic, with apterous oviparae and alate males in December-
January. Sexuales occur on P. persica as well as on P. cerasus (Agarwala et al.,
1984). 2n = 18 (Kurl, 1981).
About five species resembling Aphis but with a stridulatory apparatus con-
sisting of ventrolateral ridges on the abdomen and peg-like hairs on the hind
tibia (Eastop, 1952). Of East Asian origin, with three species now widely
distributed on shrubs and trees. Martin (1991) provided a key.
collection from Ficus microcarpa (leg. D.S. Hill) agreeing with Tao's descrip-
tion; also alatae with host unrecorded from China (Foochow; leg. M.S. Yang)
and one alata trapped in Nepal (leg. K.C. Sharma). Biology and life cycle
unknown.
Five East Asian species, at least four of them associated with Acer. Related
to Chaitophorus and Periphyllus, but with fused head and pronotum, and
with long thick, hollow hairs around margin of body, but not across dorsum.
Apterae have 5- or 6-segmented antennae. Chakrabarti and Mandal (1986)
revised the genus and keyed the species then known.
Trichaitophorus aceris Takahashi Apterae dark green with last two seg-
ments of antennae dusky, tarsi dark and siphunculi black (Takahashi, 1937a);
BL 1.4-1.8 mm. On young leaves of Acer spp.; originally described from
A. rubescens in Taiwan. Aphids identified as aceris have since been recorded
from various Acer spp. in northwest India and Nepal (Chakrabarti and
Mandal, 1986). However, there must be some doubt about the identity of the
Indian populations, as they are pale in life and have pale tarsi and siphunculi
(A.K. Ghosh, 1980). Alatae of Indian populations differ greatly from apterae,
having black-brown head and thorax, antennae and legs mainly dark, abdomen
with a large black patch on ABD TERG 4-6 and separate sclerites on
other tergites, and Chaitophorus-like siphunculi with polygonal reticula-
tion (Quednau and Chakrabarti, 1976; as Periphyllus pusillus). The popula-
tions identified as T. aceris in northwest India are monoecious holocyclic;
apterous males and oviparae were described by Chakrabati and Mandal
(1986), collected on Acer sp. in October-November.
2.4-2.5 mm. Stated to live on undersides of rolled leaves of a plant that was
unidentified at the time the description was published (Hille Ris Lambers and
Basu, 1966), but subsequently identified as Elaeocarpus sikkimensis (BMNH
colln, leg. DHRL). In West Bengal, India. Other morphs and life cycle
unknown. [The host plant and leaf-rolling habit is unlikely for a chaito-
phorine and needs additional confirmation. Probably the true hosts are Acer
spp., as listed by Chakrabarti and Mandal (1986) under T. aceris. However,
the size, colour in life, length of antennae and shape of marginal hairs make
it unlikely that the synonymy with T. aceris proposed by these authors is
correct.]
About eight species are known in East and Southeast Asia, mostly described
from secondary hosts which are all Loranthaceae. Recently the link has been
made with primary host forms that produce elaborate branching galls on
Styrax, described in the genus Astegopteryx (Aoki and Kurosu, 1993).
spinal processes on thorax and ABD TERG 1-3 that are all concolorous with
body (Shinji, 1924); BL not recorded. On Quercus acutissima and Q. serrata
in Japan (Higuchi and Miyazaki, 1969); no specimens are known, and it could
not be included in the key.
About ten species of Myzus-like aphids in East Asia, often with hair-bearing
siphunculi, galling the leaves of Prunus in spring. Where the life cycle is
known there is host alternation to Compositae (Artemisia, Gnaphalium). The
generations on secondary hosts look very different from the spring forms on
Prunus, and this has contributed to the taxonomie confusion in the group.
Accounts are available from Japan (Miyazaki, 1971), China (Chang and
Zhong, 1976) and eastern Russia (Pashchenko, 1988b). Moritsu and Tokumoto
(1972) gave an account of Japanese Tuberocephalus (under Myzus) on flower-
ing cherries in which they included several unnamed species. Sorin (1993) and
Remaudi re and Sorin (1993) described galls and life cycles of several species
that could not be included in our key to Prunus aphids.
only partially fused between segments. Life cycle is unknown. [N.B. Tao's
(1966) sakurae was probably Myzus yamatonis, and Miyazaki's (1971) descrip-
tion is partly of misakurae.]
Two species with a single large conical spinal process or tubercle on ABD
TERG 4, but otherwise not very closely related and placed in separate sub-
genera. A.K. Ghosh (1982b) provided an account including both species.
A large genus of medium-sized to rather large aphids (BL 2-4 mm) associated
almost entirely with Compositae and Campanulaceae. The very few records
from trees involve the two most polyphagous species, which occasionally occur
on plants in other families; U. ambrosiae (Thomas), a reddish-brown aphid
with black siphunculi and a pale CAUDA in North, South and Central
America, and U. compositae (Theobald), a shiny very dark red aphid with
siphunculi and CAUDA both black, distributed widely through the Old World
subtropical regions, and introduced to Brazil (see B & E, 1984, pp. 368-369).
About ten species of Myzus-like aphids with a wide range of host plant rela-
tionships (B & E, 1984, p. 321). The six North American species are mostly
associated with Rosaceae and/or Gramineae; one lives all year around on
Crataegus.
About six species with elongate swollen siphunculi, associated mostly with
Rosa and/or Ericaceae. One species utilizes Arbutus as secondary hosts.
XENOTHORACAPHIS Takahashi
Hormaphidinae: Nipponaphidini
One species on Quercus in Japan, apparently closely related to the type species"
of Thoracaphis (arboris), and perhaps not warranting a separate genus.
Seven oriental species associated with Acer and related to North American
Drepanaphis, but with a subapical zone of reticulation on the siphunculi. All
viviparae are alate. Accounts are available from Japan (Higuchi, 1972, 1974)
and India (Chakrabarti, 1988; A.K. Ghosh and Quednau, 1990).
One East Asian species on Acer, related to Trichaitophorus but with long,
thick spinal as well as marginal hairs, the latter being duplicated.
One species described from Sasa in Japan and not subsequently identified. A
summary of the original description is provided here.
COLLECTING METHODS
Aphids may be collected by sweeping or beating, or simply by careful search-
ing of appropriate parts of the plant. Beating onto a tray or card held under-
neath the branch may be useful for free-living aphids on leaves and shoots
of trees, especially Lachninae on conifers, but it is not very efficient for active,
winged aphids such as many Drepanosiphinae, adults of which will fly as soon
as disturbed leaving only the immatures and damaged adults to fall onto the
beating tray. Beating cannot be used to collect aphids that live in wax wool
such as Adelgidae, and is obviously useless for those making galls or feeding
on the trunk or large branches.
It is best, if at all possible, to examine the tree carefully and try to find
the aphids in situ, so that a representative sample of all available morphs and
developmental stages can be collected and information gained about the
feeding site and the size and appearance in life of the colony, if one is formed.
Aphids inhabiting the undersides of leaves can often be observed in silhouette
by looking up through a leaf against the sun. Ant-attended aphids, such as
Thelaxinae, Chaitophorinae and many Lachninae, can often be detected
by looking for ants moving up the trunk of the tree. Large numbers of
active flies and wasps may also indicate presence of honeydew-producing
Homoptera.
It is usually considered best to bring the aphids back to the laboratory
alive on a piece of the host plant, rather than to collect them directly into
926
Techniques 927
labelling; the system most frequently used is shown in Fig. 135. Slides may be
stored horizontally in trays, or vertically in slotted drawers or boxes. For
larger collections a compact and versatile system involves vertical storage of
slides in individual envelopes made of paper or cellulose acetate, with inter-
spersed tab cards providing the collection with an integral index (Eastop,
1985).
v
BIBLIOGRAPHY
N.B. B & E (1984) = Blackman, R.L. and Eastop, V.F. (1984) Aphids on the World's Crops. Wiley,
Chichester.
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966 Bibliography
967
968 Index to Species Names of Aphids
987