Introduction

In an account of the history of higher systematics

in the Aleyrodidae, Russell (2000) stated that five
whitefly subfamily names have been used for extant
taxa. Of these, Uraleyrodinae Sampson & Drews
(1941) was found to be synonymous with Aleyro-
dinae Westwood (1840), based on a study of adult
characters by Russell (1986). Takahashi (1932) had
erected the subfamily Siphonaleyrodinae solely for
his new species Siphonaleyrodes formosanus, which
is clearly a member of the psylloid family Triozidae,
and which was placed as a junior synonym of Trioza
cinnamomi (Boselli, 1930) by Mound & Halsey
(1978), a view with which Russell (2000) concurred.
The oldest-established subfamily, Aleyrodinae, is
generally accepted and regarded as well defined by
adult and nymphal [puparial] characters (Gill 1990).
This leaves Udamoselinae Enderlein (1909) and
Aleurodicinae Quaintance & Baker (1913) whose
controversial relationship is the subject of this paper.
The genus Udamoselis, the species U. pigmentaria
and the subfamily Udamoselinae were all proposed
by Enderlein (1909), based upon his study of a
single adult male specimen. Enderleins specimen has
subsequently never been traced, and is thought to
have been lost during the upheavals of the Second
World War. As well as being described from a single
specimen, no satisfactory collecting locality is known
and Enderlein simply gave this as in all probability
South America, indicating that the specimen must
have been given to him. Enderlein also included
Aleurodicus Douglas (1892) in his new subfamily,
without any discussion.
Quaintance & Baker (1913) discussed whitefly wing
venation in detail, illustrating a range of actual and
theoretical patterns (Fig. 33). They proposed an-
other new subfamily, Aleurodicinae, accommodat-
ing Aleurodicus, Dialeurodicus Cockerell (1902),
their own new genus Leonardius and Paraleyrodes
Quaintance (1909), whilst continuing to accept
Enderleins subfamily Udamoselinae for Udamoselis
alone. Their reason for supporting a separate sub-
family for Udamoselis was the more complex wing
venation described and illustrated by Enderlein
(Fig. 7), but the insects enormous size (Table 1) may
well have also been a factor in their decision. The
Giant whiteflies (Sternorrhyncha, Aleyrodidae):
a discussion of their taxonomic and evolutionary
significance, with the description of a new species
of Udamoselis Enderlein from Ecuador
Jon H. Martin
Three adult male whitefly specimens from Ecuador are described as Udamoselis
estrellamarinae sp. n. This genus and its subfamily are reappraised on adult
characters, including wing venation, paronychium structure, and distribution of
abdominal wax glands. In the absence of associated puparia nomenclatural caution
is preferred, but the subfamilies Udamoselinae and Aleurodicinae are likely to be
synonymous. Wing venation of other very large whiteflies is illustrated, and is
discussed in comparison with fossil taxa. Speculation is made on the possible biology
of such giant whitefly species.
Jon H. Martin, Department of Entomology, Natural History Museum, Cromwell
Road, London SW7 5BD, UK. j. martin@nhm.ac.uk
Tijdschrift voor Entomologie 150: 1329, Figs. 133, Table 1. [ISSN 0040-7496]. http://www.nev.nl/tve
2007 Nederlandse Entomologische Vereniging. Published 1 June 2007.
Tijdschrift voor Entomologie, volume 150, 2007 14
relative complexity of the wing venation and great
body size, in turn, have been regarded as possible evi-
dence that Udamoselis might be a particularly primi-
tive whitefly, and thus form a link between the other
present-day taxa and species described from the fossil
record (such as by Schlee 1970, Shcherbakov 2000,
and Hamilton 1990).
U. pigmentaria was regarded as a nomen dubium by
Mound & Halsey (1978: 250), on the basis of in-
adequate description of the adult, combined with
the absence of knowledge of the puparial stage upon
which most whitefly taxonomy is now based. Ender-
leins description of the solitary male did indeed omit
mention of some characters that are now thought
likely to be important in the systematics of adults,
and this supported the proposal that it be regarded
as nomen dubium. Perhaps most importantly, End-
erleins description and illustration of the fore and
hind wings (see Fig. 7, here) showed a venation that
is considerably more complex than had been seen in
any other known extant whiteflies: this raised a ques-
tion as to whether all the firm lines in Enderleins
drawings were truly veins and, hence, whether his
illustrations were accurate.
Leaving aside the uncertainty over wing venation,
the absence of any detail of such characters as ab-
dominal wax glands or tarsal paronychium, the loss
of the antennal flagellum in his sole specimen, and
the lack of optical resolution available to Enderlein
[he stated that no empodial paronychium was vis-
ible with his magnifying glass], his description was
nonetheless remarkably detailed if sometimes rather
ambiguous. With the considerable importance of
this taxon Quaintance & Baker (1913) provided a
complete English translation of Enderleins descrip-
tion of U. pigmentaria and this translation has been
extensively consulted in the course of the present
study. In the absence of study material, Mound and
Halseys (1978) decision to regard U. pigmentaria as
nomen dubium was pragmatic, allowing the continu-
ing use of Aleurodicinae as the name for the numeri-
cally smaller of only two extant whitefly subfamilies,
accommodating about eight percent of described
whiteflies.
Schlee (1970) stated that The systematic position of
Udamoselis within the Aleyrodina cannot be elabo-
rated until a new find is made, because of the insuffi-
cient present knowledge based upon the single specimen,
which has probably been destroyed. The assumed close
kinship relation between Udamoselis and the Aleu-
rodicidae [i.e. Aleurodicinae] is unproved. In contrast,
Shcherbakov (2000) said: . despite an incomplete
knowledge of the type genus, the name Udamoselinae
should be used in the broad sense of Enderlein (1909)
and Sampson (1943), i.e. including Aleurodicinae.
Three new specimens recently collected in Ecuador
correlate with Enderleins description sufficiently well
to be regarded as belonging to Udamoselis, thus al-
lowing this intriguing controversy to be reappraised.
Many of the attributes described by Enderlein for
U. pigmentaria are apparently accurate, although
other parts of his description remain ambiguous
through the absence of the original specimen, com-
bined with Enderleins failure to provide any illus-
trations beyond the wings. Nevertheless the author
now considers it quite likely that U. pigmentaria it-
self will prove to be identifiable, in the event of new
material becoming available, and its identity should
no longer be regarded as nomen dubium. However,
the subfamilial position of Udamoselis remains some-
what uncertain, as will be discussed later in this
paper.
Examination of the three males from Ecuador has
revealed their wings (Figs 5, 6) to display the identi-
cal venation illustrated by Enderlein (Fig. 7), but has
confirmed that not all veins are as distinct as implied
by Enderleins simplified line drawings. Comparison
of the Ecuadorean material with the description of
U. pigmentaria leads to the conclusion that the two
taxa are congeners but are distinct species. Despite
the small sample size, and frustrating lack of females
and (especially) of puparia, it is felt that naming the
Ecuadorean species is valid because of the wider in-
terest in higher systematics that these specimens are
likely to generate. Udamoselis estrellamarinae is there-
fore here described, and is named for its discoverer
(see below).
Materials, methods and terminology
Background
In 2005 the author visited Ecuador, in company
with Dra Estrella Hernndez-Suarez and Sr Elicio
Tapia. The purpose was to search for whitefly col-
onies that might yield natural enemies of the pest
species, Lecanoideus floccissimus Martin, Hernn-
dez-Surez & Carnero, 1997, in connection with
achieving its natural control in the Canary Islands.
Whilst sorting collected material for possible rearing
of parasitoids, Hernndez-Suarez noticed three very
large, darkly-pigmented and relatively wax-free adult
male whiteflies inside a bag containing a substantial
colony of Lecanoideus mirabilis (Cockerell, 1898)
on Annona leaves. An extensive search of other bags
of material from the same garden tree failed to re-
veal any additional specimens. The three specimens
were brought back to the laboratory at the Natural
History Museum, London, for further study.
Martin: Giant whiteflies and new Udamoselis 15
Treatment of adult whiteflies for examination
It is not easy to dissect any adult whitefly for mi-
croscopic examination on slides, in contrast with the
situation with Psylloidea (jumping plant lice) which
are routinely dissected prior to slide-display. Never-
theless, the interest in wing venation of larger white-
flies leads the author now to remove all wings from
such specimens, prior to maceration of the body, and
to mount the completely untreated wings under a
separate coverslip. This avoids distortion and dete-
rioration of any subtle wing pigmentation, and the
separate coverslip allows for a very thin (hence flat)
mount for the wings.
With very large taxa a partial dissection of the body
also assists in better displaying certain characters for
study, but with very small sample sizes this inevita-
bly results in some other characters being distorted
or obscured. The author usually dissects the head
(sometimes with the rostrum still attached) for exam-
ination frontally, the abdomen for lateral mounting,
male forceps for dorso-ventral display, and the aedea-
gus for lateral examination. The thorax, with legs still
attached, may be mounted laterally (Fig. 1) for the
practical reason that whitefly coxae are so strongly
ventrally-directed that normal dorso-ventral display
is virtually impossible. Alternatively, the specimen
may be more extensively dissected (Fig. 4), allow-
ing more constituent parts to be displayed parallel
to the slide surface. With the much thicker mounts
that are needed for the thorax / legs and abdomen, in
comparison with that required for the genitalia and
head, the constituent parts of the specimen may be
distributed between two to four coverslips of 10 mm
or 13 mm diameter (Fig. 4), but always with all the
parts of one adult on a single slide. The practice of
dividing the constituents of one individual between
two slides, sometimes used by workers on Psylloidea,
is to be avoided.
When adult whiteflies are not dissected, the favoured
alignment is generally lateral, with wings dorsally (as
in the resting position of Rhopalocera) and with
legs and rostrum displayed ventrally as seen in Fig.
1. With much smaller adults, usually members of
the Aleyrodinae, displaying them is more challeng-
ing, because manipulation requires even more care
and osmotic collapse becomes a greater risk. When
a large sample is at hand, the author usually resorts
to placing many individuals under a single coverslip,
with minimal effort being expended on each indi-
vidual: the result is that usually all characters may
be seen, but only by examining several specimens.
With very small adults, better results are sometimes
achieved by clearing and dehydrating un-macerated
specimens and placing them directly onto a slide.
Wing venation terminology
The wing venation of Udamoselis pigmentaria is
important for the discussion of its genus and other
larger whiteflies. Enderlein (1909) employed a termi-
nology (Fig. 7) that is not accepted today, but there
appears to be little consensus on which alternative
to use. Accordingly, for this communication I have
followed the system used by Quaintance & Baker
(1913; see Fig. 33, here), Solomon (1935) and Gill
(1990). For Udamoselis this is shown in figures 5 and
6, where ? indicates my uncertainty over whether
the posteriormost feature on each wing is truly a
vein. This venation terminology is also used here in
discussion of the wings of several other large whitefly
species (Figs 19-28).
Depositories
BMNH The Natural History Museum, London,
U.K.
USNM Entomological collections of the U.S. Na-
tional Museum of Natural History, housed
at U.S. Department of Agriculture, Belts-
ville, Maryland, U.S.A.
The holotype and paratype 2 of U. estrellamarinae
are deposited in BMNH; paratype 1 of U. estrellama-
rinae is deposited in USNM, and all three specimens
have been slide-mounted as described above, with
varying degrees of dissection. All the other specimens
discussed and illustrated here are also housed in the
collection of BMNH.
Udamoselis Enderlein
Udamoselis Enderlein, 1909: 230. Type species: Udamoselis
pigmentaria Enderlein, by monotypy, but sole original
specimen not traced.
Udamoselis estrellamarinae sp. n.
Figs 16, 818
Type material. Holotype: adult ?, Ecuador, Manab
Province coast, Brisea, 5 km north of San Vicente,
on Annona sp., probably A. muricata (Annonaceae),
10.ii.2005 (Hernndez-Suarez coll., Martin ref.
#8103) (BMNH). Paratypes. adult ?: same data as
holotype (BMNH; USNM).
Adult male (n=3)
Measurements. For basic measurements of many
body parameters, see Table 1.
Coloration. Body colour very dark, with little ap-
parent waxy bloom covering the cuticle, resembling
small Psylloidea to the naked eye. Pigmentation
of the body cuticle as shown in the photographs
(Figs 1, 4), but is generally evenly dark brown; the
Tijdschrift voor Entomologie, volume 150, 2007 16
middle parts of the femora and the whole of the
tibiae, tarsal segments and antennal flagellar seg-
ments are paler. The wings are rather leathery and
their dark pigmentation (Figs 23) can best be de-
scribed as blotchy; each fore wing has two ovoid pale
patches, and each hind wing a single trapezoidal pale
zone; the degree of darkening of the wings varies be-
tween the three specimens, with the holotype (Figs
23) the darkest.
Legs. Femoral hairs normal on all legs, bristle-like;
many hairs on distal parts of tibiae thickened but
still apically acute; each middle tibia with a group
of thickened hairs forming a distinct comb at three-
quarters length, each hind tibia with a more subtle
comb formed by a smaller number of thickened
hairs, but each fore tibia without such a comb of
hairs; each hind tibia with an apical arc of 911 pale
dagger-like spines (Fig. 18) that may be saltatorial in
function (as in Psylloidea), but these are absent from
other legs. Tarsi two-segmented, basal segment much
longer than apical segment; hairs on basal segment
stouter and more numerous than those on apical
segment; hind basal tarsal segment with two rows of
stout ventral spines (Fig. 17), which (like the api-
cal hind tibial spines) may be saltatorial in function;
claws paired, each pair with a single stout ventrally-
directed spine-like paronychium between them
(Fig. 9). Tibial and tarsal surfaces with very fine
spinules, becoming dense enough on tarsi to be re-
garded as a fine pubescence.
Abdomen. With a pronounced furrow running along
each side of segments IIIVII, above the mid-line;
segment VIII about as long as deep, slightly ex-
panded at distal end, without a ventral spur at its
distal extremity; segment IX about 3 times as long as
deep; sternites IIIV each with a pair of wax plates
(Fig. 14), the anterior two pairs significantly larger
than the posterior pair; each anterior wax plate with
a deep invagination from its anterior side, lined with
hairs and accommodating a patch of rounded reticu-
lations; middle wax plates with a similar reticulate
patch situated in an indentation of the anterior edge
of the plate; hind wax plates evenly ovoid, without
indentations and not fringed with hairs; surfaces
of wax plates extremely finely imbricate-reticulate
(400 magnification), with a few hairs on their sur-
faces as shown in figure 14; segments VIIIIX devoid
of hairs but other segments with hairs surrounding
wax plates (Fig. 14) and near postero-ventral margins
of sternites; tergites IIIIV each with a subcircular
patch of rounded reticulations on each side (Fig. 16);
whole of abdomen punctuated by tiny pale ovoid
pores (Figs 1416). Lingula and presumed oper-
culum (Fig. 15) situated between one-third and
half way from base to apex of segment IX; lingula
with fine spinule-pubescence but no evident hairs;
Figs 14. Photomicrographs of Udamoselis estrellamarinae. 1, laterally-mounted body, holotype; 2, fore wing,
holotype; 3, hind wing, holotype; 4, dissected thorax / legs and abdomen, paratype 2.
1
2
4
3
Martin: Giant whiteflies and new Udamoselis 17
presumed operculum with similar spinule-pubes-
cence but only with a single apical hair visible in
lateral aspect [it is to be expected that there is a pair
of such hairs]; claspers evenly curved in dorso-ven-
tral aspect (Fig. 10) but flat in lateral aspect (Fig. 1),
generally smooth and with a few fine hairs as shown
in figure 10; aedeagus with a distinct elbow at half-
length (Fig. 12), about 0.325 mm from apex to el-
bow, apically with two large and possibly four much
smaller finger-like protrusions (Fig. 13).
Head. As shown in antero-ventral view in figure 8;
small for the size of the insect (1 mm wide), much
wider than long; compound eyes very large and in-
dented to accommodate the antennal insertions;
lateral ocelli well-developed, as shown in figure 8,
but median ocellus not evident in any of the three
mounted specimens; between the eyes is situated a
conical protrusion from the frons; clypeus densely
hairy; basal two antennal segments dark brown,
smooth and with fine hairs; the five-segmented
antennal flagellum (Fig. 11) without hairs but with
finely spinulose transverse striations throughout its
length (Fig. 11, expanded detail); flagellar segments
with many extremely small, pale spots that are likely
to be sensoria, with a few slightly larger sensoria
on the apical segments; ultimate rostral segment
pigmented dark brown in apical third, with many
hairs.
Thorax. Thoracic dorsal plates smooth, pigmented
brown and sparsely provided with fine hairs, except
for a fringe of longer hairs anteriorly on prothorax.
Wings. Rather narrow, dimensions given in Table 1,
fore wing margin flattened to slightly emarginate op-
posite apices of veins R
s
and M (Fig. 5). Pterostigma
well defined by surface roughening (Fig. 5) rather
than by differentiated coloration (Fig. 2). Venation
of fore wing as shown in figure 5; Sc contiguous with
C until diverging as it approaches the pterostigma;
R
1
short, becoming indistinct in the pterostigma;
R
s
almost reaching wing margin; M very long,
C
Sc
R1
Rs
M
Cu
?A
R
R
1
R
s
6
7
5
R
r
1
r
m
cu
an
ax
r
cu
an
m
?Cu
Figs 57. Wings of Udamoselis species. 5, U. estrellamarinae sp. n., fore wing with venation annotated using ter-
minology adopted here; 6, U. estrellamarinae sp.n., hind wing with venation annotated using terminology adopted
here; 7, U. pigmentaria, original figure from Enderlein (1909), showing Enderleins venation terminology. [Scale
bars = 1 mm]
Tijdschrift voor Entomologie, volume 150, 2007 18
9
b
13
a
8
10 11
14
12
c
15
Figs 815. Udamoselis estrellamarinae. 8, head, anteroventral view, paratype 2; 9, claws and empodial spine;
10, clasper, paratype 1; 11, antenna, with detail of apical 2.5 segments, paratype 1; 12, aedeagus, paratype 2;
13, aedeagal apex of (a) paratype 2, (b) paratype 1, (c) holotype; 14, wax plates on abdominal sternites IIIV,
holotype; 15, lateral aspect of operculum (left) and lingula (right), paratype 1.
Martin: Giant whiteflies and new Udamoselis 19
16 17
18
19
20
21
22
Figs 1622. 1618, Udamoselis estrellamarinae. 16, oval patches of reticulations on abdominal tergites III and
IV, paratype 2; 17, distal half of hind basitarsus, paratype 1; 18, apex of hind tibia, showing fringe of spines and
thickened setae proximad, paratype 1. 1920. Wings of male syntype of Parudamoselis kesselyaki Visnya. 19, fore
wing; 20, hind wing. 2122. Wings of unidentified male whitefly from Tena, Ecuador. 21, fore wing; 22, hind wing.
[Scale bars for wings only = 1 mm, one per wing pair]
Tijdschrift voor Entomologie, volume 150, 2007 20
23
24
25
26
27
28
Figs 2328. Wings. 2326, Wings of unidentified whitefly (JHM 8078) from Bartola, Nicaragua. 23, fore
wing, male; 24, hind wing, male; 25, fore wing, female; 26, hind wing, female; 2728. Wings of Dialeurodicus
caballeroi, male. 27, fore wing; 28 hind wing. [Scale bars = 1 mm, one per wing pair]
Martin: Giant whiteflies and new Udamoselis 21
diverging from R near wing base, almost straight
but slightly angled posteriorly in its apical quarter;
Cu (see discussion, below) faintly-indicated basally
and apically, parallel to and closest to M at half-
length, markedly paler than the heavily pigmented
wing surface; posterior to Cu is a subtle and irregular
fold that is clearly what Enderlein regarded as the
Axillaris [anal vein, here], but it is unclear whether
this really warrants such status. Hind wing with R /
R
1
& R
s
clearly marked, with M diverging from R
at wing base; in a similar situation to that seen in the
fore wing, a subtle thickening / fold indicates what
Enderlein regarded as the Analis [vein Cu here].
Coloration and patterning of wings as shown in
Figs 23.
Whichever fore wing venation system is followed the
entity named Cu (Fig. 5) is clearly associated with
the claval furrow (sutura clavi of Enderlein), always
almost straight for much of its length and distinctly
pale on pigmented wings (a very distinct bright line
as described by Enderlein for his fore wing Analis
in figure 7). In Psylloidea, the same observations are
true for vein Cu
2
which doubles as the clavus (Hod-
kinson & White 1979). Indeed, the present author
questions whether this is really a vein at all.
Remarks. U. estrellamarinae differs from U. pig-
mentaria in some obvious respects, assuming End-
erleins description of the latter to be accurate. The
wings of U. estrellamarinae are more elongate in
shape, clearly seen by comparing figures 5-6 with
figure 7, and by comparing the parameters given in
Table 1. The margin of the fore wing of U. estrel-
lamarinae is distinctly emarginate opposite the apex
of vein Rs, and slightly less so opposite the apex of
M, in contrast to the evenly curved distal margin in
U. pigmentaria. Enderlein described the hind wing
of U. pigmentaria as densely sprinkled with dark
brown spots, in contrast to the almost uniformly
dark hind wing, punctuated by tiny pale spots
(Fig. 3), in U. estrellamarinae: otherwise, the descrip-
tion of the wing pigmentation in U. pigmentaria is
torturous and demonstrates well why photographs
are the best way to illustrate such characters.
Despite the wings of U. pigmentaria being
somewhat longer, and much broader than in
U. estrellamarinae the probability is that the body of
U. pigmentaria is smaller than in U. estrellamarinae,
with abdominal segment IX (see Table 1) being a
particularly clear indicator despite the ambigu-
ity over whether Enderleins stated body length for
U. pigmentaria was inclusive or exclusive of the
forceps.
The conical protuberance on the frons was described
as black in U. pigmentaria but is concolorous with
the remainder of the head in U. estrellamarinae.
Giant size and wing venation in the
Aleyrodidae
From the description and discussion, above, Uda-
moselis is clearly a genus with exceptionally large
males. With no females available for study, it is by no
means certain that females will prove to be as large.
There is a good reason for sounding such a caution-
ary note (see below).
Figures 19 and 20 show the fore and hind wings of
the sole syntype (an adult male) of Parudamoselis
kesselyaki Visnya (1941) in BMNH. Described as a
gigantic whitefly by Visnya (from a large introduced
population in a Hungarian glasshouse), the BMNH
male is actually even larger than the Udamoselis males
(see Table 1), but the body cuticle and wing colora-
tion are more typical for a whitefly, not being darkly
pigmented. Although P. kesselyaki is now regarded as
a junior synonym of Ceraleurodicus varus (Bondar,
1928), based on comparison of syntype puparia,
(see below) of both species in the USNM collection
(Martin et al. 2000), the discussions here are based
upon the BMNH adult male syntype of P. kesselyaki
and Visnyas observations on the Hungarian mate-
rial from the 1940s; no other males are known at
the present time. The fore wing of the BMNH syn-
type of P. kesselyaki (6.00 mm long, 2.95 mm maxi-
mum width) displays the same much-curtailed R
1
,
terminating in a pronounced pterostigma, which is
also seen in Udamoselis. There is also a suggestion
that thickening of the proximal boundary of the
pterostigma could be the equivalent of the putative
vein Sc in Udamoselis. As in the males of both spe-
cies of Udamoselis and other large species discussed
here, the result is a more robust fore wing. Visnya
also remarked upon distinct adult size-dimorphism
in P. kesselyaki. Four females reared from puparia on
Protium copal in Belize, and identified as C. varus
from their puparia, are considerably smaller than the
only (male) syntype of P. kesselyaki in BMNH, and
also smaller than Visnyas quoted measurements for
females of P. kesselyaki.
Visnya further noted that the male puparia of
P. kesselyaki were significantly larger than those of fe-
males, as follows: The length of most L
4
is between 3.8
and 4.3 mm. These are all females and the well devel-
oped eggs can be seen in matured ones before emerging.
There are L
4
of 5 mm and above, these seem to be males,
so far however I could only find one preparation with a
larva of 5.3 mm length and 2.5 mm wide, in which the
penis and claspers are visible.
[Authors note: the observations below concern some
undescribed whitefly taxa. The characteristics de-
tailed here are not intended to be for nomenclatu-
ral purposes, but only to broaden this discussion
topic.]
Tijdschrift voor Entomologie, volume 150, 2007 22
In 2004 a congregation of extraordinary adult white-
flies was discovered under a single leaf in Nicaragua
[Uncaria tomentosa (Rubiaceae), Rio San Juan / Rio
Bartola confluence, 22.vi.2004, Martin #8078]. The
generic position of this undescribed species is un-
certain, because almost nothing is known about the
relative importance of different adult characters in
whitefly generic definitions. This situation has arisen
because for 150 years whitefly taxonomy has been
based almost solely upon puparial [i.e. final nymphal
stage] characters (Martin 2003). This species is here
referred to simply by its collection number, JMH
8078, and it is notable for two reasons. Firstly, one of
the five males (Fig. 29) may be the largest individual
whitefly ever collected, with an overall body length
of over 1 cm (see Table 1). Despite this exceptional
length, the body is possibly less massive than the
males of Udamoselis, with the extreme length being
due to abdominal segments VIII and IX (Fig. 29).
At 3.78 mm long in this largest specimen, its ab-
dominal segment IX alone is longer than almost all
other whitefly species, including those in the Aleuro-
dicinae. However, the fore wings of these males (3.80
mm long in the largest specimen, down to 3.20 mm
in the smallest) are considerably smaller than those
of Udamoselis males, presumably reflecting a lower
body mass.
JMH 8078 is also notable for extreme adult dimor-
phism (Fig. 29). The adult females (body length
2.102.60 mm, fore wing length 1.752.20 mm,
n=7) are all very much smaller than the males, and
are similar in size to many other Aleurodicinae. The
difference in wing size, between males and females
of JMH 8078, is at once obvious from comparison
of figures 2324 with 2526 (all shown to the same
scale). Size apart, while the female fore wing displays
an entirely typical aleurodicine venation, that of the
male has vein R
1
converging with the costal margin,
there forming a small pterostigma, and then contin-
uing towards the wing apex and almost paralleling
the curve of R
s
. The increased complexity of the fore
wing venation in the male results in a more robust
structure.
Figures 21 and 22 show the fore and hind wings
of a single large (dissected) male specimen (fore
wing 3.92 mm long, 2.19 mm maximum width)
from Ecuador [Tena, 23.ii.1923, F.X. Williams coll.],
which also remains unplaced generically. Here, R
1
is
Fig. 29. Unidentified whitefly (JMH 8078) from Bartola, Nicaragua largest male specimen (left) and a female
specimen (right) from same collection, in alcohol prior to slide-preparation, showing extreme dimorphism.
Martin: Giant whiteflies and new Udamoselis 23
Figs 3032. Likely models for Udamoselis puparia habitus photographs of puparia of species of Aleurodicinae with
sparse wax, radial rays and additional ventral tracheal folds, that do not occur in aggregations. 30, Ceraleurodicus
keris Martin, on Lunania parviflora, Nicaragua; 31, Octaleurodicus sp., on ?Melastomataceae, Ecuador; 32, Ceraleu-
rodicus sp., on Clusia grandiflora, Guyana.
30
31
32
Tijdschrift voor Entomologie, volume 150, 2007 24
Fig. 33. Reproduction of Plate 1 from Quaintance & Baker (1913), showing wing venations of present-day white-
flies, theoretical origin of whitefly wing venation, wing venation of present-day Trioza sp. (Psylloidea) and theoreti-
cal origin of triozid wing venation.
Martin: Giant whiteflies and new Udamoselis 25
reminiscent of JMH 8078 but, instead of a pte-
rostigma, there is a curious fusion with the putative
Sc. Also, M is broken at its proximal end, with the
basal stub displaced towards Cu (this curious fea-
ture is identical in both fore wings). Once again, the
result is a more robust structure, possibly to support
a larger-than-usual body mass.
Although very large males are sometimes associated
with much smaller females (JMH 8078 and P. kes-
selyaki [= C. varus], see above), there are also very
large female whiteflies in existence. One such female
[Chiococca alba (Rubiaceae), Belize, Chiquibul forest
Reserve, 01.vi.2004, Martin #7967], also of uncer-
tain generic position, has fore wings 3.65 mm long,
2.00 mm maximum width and hind wings 3.10 mm
long, 1.40 mm maximum width. The abdomen has
four pairs of wax glands, the posteriormost two pairs
with coarsely reticulate facets (which are reminiscent
of those in Aleurodicus dispersus). The body, is not
dark brown and the wings have only isolated spots of
pigment, so it is not thought likely that this individ-
ual will prove to be a species of Udamoselis. Despite
the large size of the wings of this female, the venation
is typically aleurodicine, with R widely-forked and
no pterostigma.
Finally, figures 27 and 28 show the wings of the male
of Dialeurodicus caballeroi Martin, 2004, a much
smaller species than those discussed above (fore wing
1.83 mm long, 0.86 mm maximum width). This is a
wing type commonly seen in aleurodicine species of
medium size, with R
1
simply but well developed in
the fore wing, similar to that seen in the hind wing,
but contiguous with a large pterostigma in the fore
wing. There is no indication of size dimorphism in
D. caballeroi, with female fore wings even slightly
longer and broader than in most males. [The pter-
ostigma in both sexes is much less distinct in speci-
mens whose wings have been through the macera-
tion process prior to slide-mounting.]
In all the taxa discussed above, the hind wings have
the putative Cu either short and poorly indicated, or
apparently absent, and are thus typical of aleurodi-
cines. The only oddity amongst these hind wings is
seen in the male syntype of P. kesselyaki (Fig. 20),
where R
1
has a curious (?vein) stub arising at its half-
length, a feature seen in both of the hind wings.
Observations on wings and other
characters, with respect to subfamily
placement in extant taxa
Added to the detailed comments on the very large
taxa, above, the following broader observations may
be made:
Almost all members of the Aleurodicinae have
the fore wing with R forked to form R
1
and R
s
,
and are larger insects than most members of the
Aleyrodinae. Udamoselis also displays these at-
tributes.
Most members of the Aleyrodinae do not have a
forked R, but only a single main vein.
The only Neotropical members of the Aleuro-
dicinae that are physically very small, and thus
similar in size to typical members of the Aleyro-
dinae, are the members of the genus Paraleyrodes,
whose fore wings have an unbranched main vein,
as is typical in the Aleyrodinae.
A few unusually large members of the Aleyrodi-
nae have their fore wings with R forked, or have
other complications to their venation. Examples
include European species of Aleurochiton Tull-
gren, 1907 and the northern Australian native,
Gagudjuia allosyncarpiae Martin, 1999 and at
least one species of Aleuroparadoxus Quaintance
& Baker, 1914 [an undescribed species from
Guyana].
All the males discussed here, including Udamose-
lis and JMH 8078, have three pairs of abdominal
wax plates normally regarded as a diagnostic
aleurodicine feature (Gill, 1990).
All the males discussed here, including Uda-
moselis and JMH 8078, have a spine- or seta-like
paronychium, again regarded as a diagnostic
aleurodicine feature.
Size dimorphism is also quite common in the
Aleyrodinae but it is then usually the adult
males, and male puparia, that are smaller. In gen-
era such as Aleurocanthus, the male and female
puparia have not infrequently been described as
separate species.
The fossil dimension
Schlee (1970) presented a detailed discussion of Cre-
taceous and Tertiary amber-fossilised whiteflies. As
part of his comparisons of these with extant taxa, he
said the following:
The recent Udamoselis is by no means the most primi-
tive recent Aleyrodid; it does not represent the Aley-
rodinas ground plan. Moreover it is one of the highly
derivative forms, exhibiting numerous autapomorphies
(one of which is the enormous size). The relatively com-
plete and distinct venation is connected with the large
body size. He further stated that The wing venation
gives no evidence for the kinship relations within the
Aleyrodina; families must not be defined by this fea-
ture. These views are supported by the quoted small-
to-medium size of the following fossil taxa: Heidea
cretacica Schlee, 1970, body length 1.02 mm, fore
wing length 0.83 mm; Bernaea neocomica Schlee,
Tijdschrift voor Entomologie, volume 150, 2007 26
1970, body length 1.30 mm, fore wing length 1.08
mm; Juleyrodes gilli Shcherbakov, 2000, fore wing
length 2.1 mm (no associated body); Burmoselis
evelynae Shcherbakov, 2000, body length 0.95 mm,
fore wing length 1.10 mm. The wing venation of
H. cretacica was not discernible, but the venations of
the other three species vary from similar to present-
day Aleurodicinae to slightly more complex than
in Udamoselis. Another fossil species, Megaleurodes
megocellata Hamilton, 1990 [Cretaceous], measured
11.5 mm in length but is clear to the present au-
thor that Shcherbakov (2000) has already correctly
regarded this particular insect as being a member of
the Fulgoroidea (Auchenorrhyncha), having three-
segmented tarsi and legs that would be characteristi-
cally angular in cross-section.
These limited data do not, of course, preclude the
possibility that very large whiteflies have occurred
in the fossil record. Nevertheless these observations
combine to suggest that there has been an evolu-
tionary trend towards the extremely simplified wing
venation seen in over 90 percent of extant whitefly
species, and that this trend may have been reversed
for larger species, with a more complex venation re-
tained or re-evolving in the cases of Udamoselis, most
members of the Aleurodicinae and some larger mem-
bers of the Aleyrodinae.
Observations on biology
All specimens of the very large taxa, and specimens
of several other larger species, are represented in
the BMNH collection only by small numbers of
adults, these sometimes having been discovered in
apparent mating congregations with no associated
puparia. A personal observation, that the author be-
lieves to be correlated, is that puparia of several large
aleurodicine species currently placed in Ceraleu-
rodicus Hempel, 1922 are found widely scattered
over their hosts, and not in the kind of aggregations
that are typical for most other species in the Aleu-
rodicinae. It is considered possible that the adults
of such species congregate for mating, but that the
females then distribute single eggs very widely. If this
is the strategy also adopted by Udamoselis then it is
likely that the three individuals of U. estrellamarinae
were a nucleus for such a mating group, and it is also
likely that puparia will be difficult to find.
Visnyas comments on male puparia of P. kes-
selyaki concur with the present authors field ob-
servations of this whitefly group (see description of
Ceraleurodicus keris and account of C. varus by
Martin 2004) that males are even more difficult
to find than are females, and no males or probable
male puparia of either C. varus or C. keris have been
discovered despite detailed searches.
A characteristic of the puparia of larger spe-
cies currently accommodated in Ceraleurodicus,
Dialeurodicus and Octaleurodicus Hempel, 1922
(Figs 3032) is that they feature nine pairs of radial
rays [peripheral intersegmental ridges of Shcherba-
kov 2000]. At least some of these rays terminate in
very fine combs of modified marginal teeth, and the
secreted glassy peripheral filaments are thus narrower
at these points, appearing more opaque and render-
ing the rays clearly visible within the glassy skirt that
surrounds the puparium (Figs 3032). Some rays in
some species also have apparent tracheal folds un-
derlying them ventrally (Martin 2004). Shcherbakov
considered that this tracheal feature indicates a more
complete complement of spiracles than is usual in
whitefly puparia. Such puparia also tend to have their
compound pores reduced in size and number (those
entirely without compound pores being assigned
to Dialeurodicus), and do not produce the copious
woolly tangles of secretions that are so commonly
associated with other members of the Aleurodicinae.
Three puparia of the type discussed above are shown
here: figure 30 depicts Ceraleurodicus keris, with an
asymmetric puparium and with a very long and com-
plex filament arising from each compound pore, but
otherwise highly cryptic; figure 31 shows a species of
Octaleurodicus, with short and glassy fingers of wax
secreted by eight small, submedian compound pores;
figure 32 features a species of Ceraleurodicus with ex-
tremely reduced (submedian) compound pores, and
no discernible dorsal secretions at all.
One can only speculate on the size and appearance
of puparia of Udamoselis, or those of other giant
males such as those of JMH 8078, but Visnyas ob-
servation that the male puparia of P. kesselyaki are
significantly larger than those of females is likely to
be relevant. The puparia of both C. varus (recent-
ly-collected material from Central America) and
C. keris have been recorded as reaching 4 mm in
length, but these may all be female. The widely scat-
tered puparia of at least some species currently placed
in Ceraleurodicus are highly cryptic in life, hinder-
ing field searching. This was especially marked with
C. varus in Belize (Martin 2004).
A few old puparia of C. keris were discovered on the
same host plant as the aggregation of adults of JMH
8078, and it remains a possibility that these two en-
tities may be conspecific [no adults of C. keris have
been reared from puparia].
Provisional conclusions
Based upon all the adult male characters for
Udamoselis, now available for study on actual speci-
Martin: Giant whiteflies and new Udamoselis 27
mens, there is nothing that suggests to the author
that Udamoselis should be regarded as belonging to
a different subfamily from Aleurodicus and other
members of the Aleurodicinae. This concurs with the
conclusions of Solomon (1935), Sampson (1943),
Schlee (1970) and Shcherbakov (2000), as discussed
by Martin & Streito (2003). The same may be said
of the other large taxa with relatively complex wing
venation, discussed here. In large part, this conclu-
sion is based upon the proposition that more com-
plex wing venation is a response to larger body mass
(as promulgated by Schlee 1970), and is thus not
a feature of importance for subfamilial placement.
This proposition is made because other major at-
tributes are entirely typical for Aleurodicinae.
It therefore follows that there is no compelling rea-
son for regarding Udamoselis as providing evidence
of a missing link between fossil whitefly taxa and
those alive today. It is of particular note that at least
some fossil taxa were not giants (see above), despite
their more complex wing venations, leading Schlee to
conclude that Udamoselis is actually highly derived,
rather than plesiomorphic, in some of its features.
However, the continuing (and frustrating) lack of
adult females and (especially) immature stages, reli-
ably associated with males of Udamoselis, prevents a
final conclusion from being drawn here on its subfa-
milial position. Should future collecting yield females
and puparia, which further support the view that
Udamoselis, Aleurodicus and the other taxa discussed
above do indeed all belong to the same subfamily,
then Udamoselinae is the older name that would
then take precedence over Aleurodicinae. Solomon
(1935) did express the opinion that Aleurodicinae
and Udamoselinae were one and the same, but then
incorrectly used the more recent name, Aleurodici-
nae, as valid.
Shcherbakov (2000) has expressed the opinion that,
should puparia of Udamoselis display the feature of
rays and additional tracheal openings at the puparial
margin (see above, and Figs 30-32) then all such taxa
might reasonably be placed in a separate tribe, Uda-
moselini, and those without them in a second tribe,
Aleurodicini.
A final prediction
Although still a speculative opinion, the authors
interpretation of the biological evidence discussed
above does lead him to expect the following at-
tributes to be displayed by Udamoselis puparia:
They are likely to be rather cryptic (i.e. without
copious woolly secretions);
They are likely to be scattered widely over their
host(s);
They are likely to resemble those of aleurodicine
species with relatively cryptic puparia that have
radial rays (Figs 30-32), and possibly also under-
lying tracheal folds;
Males of the size of U. pigmentaria and U. estrel-
lamarinae will need puparia of at least 5-6 mm
long, as observed by Visnya (1941) for one prov-
en male puparium of similar-sized Parudamoselis
kesselyaki;
Discovery of Udamoselis puparia will finally con-
firm that Udamoselis and Aleurodicus are mem-
bers of one subfamily.
Should the above prove to be true, Udamoselinae
will then become the valid name for the numerically
smaller of only two extant whitefly subfamilies (un-
less a case for exception is made to the International
Commission for Zoological Nomenclature).
Acknowledgements
Field work in Ecuador was made possible by the
enthusiastic logistical support provided by Profes-
sor Giovanni Onore of the Pontificio Universidad
Catlica del Ecuador, Quito (PUCE), who was in-
strumental in obtaining necessary permissons, and
who advised on collecting locations. The field work
was carried out under Investigation Authorisation
#008 IC FAU.DNBAP/MA, and export certifi-
cate #0687999. The energetic field assistance and
incredibly sharp eyes of Elicio Tapia (PUCE) were
also extremely valuable and directly resulted in the
collection of the sample that yielded the specimens
of Udamoselis that are the main subject of this com-
munication. The author would also like to thank the
Instituto Canario de Investigaciones Agrarias, La
Laguna, Spain for supporting his participation in the
Ecuador field project.
Field work in Nicaragua, which also yielded material
of importance to these discussions, was facilitated by
the 2004 Universidad de Leon entomological expe-
dition, organised by Jean-Michel Maes.
All line drawings and photography are the work of
the author, with the exception of figure 33, which
is a reproduction of plate 1 of Quaintance & Baker
(1913). Scanning of line drawings was carried out by
Pat Hart (BMNH).
My thanks are extended to Penny Gullan for her
valuable comments on an earlier draft of this paper.
Tijdschrift voor Entomologie, volume 150, 2007 28
Table 1. Measurements (in mm) of main body parameters of males of Udamoselis estrellamarinae sp. n.,
U. pigmentaria Enderlein, and some other very large whiteflies.
Key: L = length; W = width; abd. = abdominal; abd. L = abdominal length, including forceps in ?; ant. = antennal
segment(s); body L = body length, including forceps in ?; dts = distal tarsal segment; fch = frontal cone height in
relief; fwpd = maximum dimension of fore wax plate; f+t = femur + trochanter; HT = holotype; hwpd = maximum
dimension of hind wax plate; mwpd = maximum dimension of middle wax plate; PT = paratype; pts = proximal
tarsal segment; vas = ventral abdominal spur on abdominal segment VIII.
sp. n. HT ? sp. n. PT ? 1 sp. n. PT ? 2 U. pig- Parud. kessel JHM 8078 indet. ? indet. /
(BMNH) (USNM) (BMNH) menta- yaki ? largest Tena, JHM 7967
ria ? **** ? Ecuador Belize
body L 8.50 - - 7.00* 6.30*** 10.5 - -
fore wing L 5.00 5.20 5.00 5.50 5.00 (6.00) 3.80 3.92 3.65
fore wing W 2.10 2.25 2.20 2.91** 2.70 (2.95) 1.70 2.19 2.00
hind wing L 3.50 3.6 3.55 3.75 4.00 (4.55) 2.70 3.15 3.10
hind wing W 1.25 1.26 1.28 1.69** 1.80 (2.00) 1.03 1.62 1.40
abd. L 5.90 6.75 5.80 4.75* 4.50 (6.30) 8.95 5.85 2.40 [/]
abd. I-VII 2.10 2.40 2.10 (3.00) 2.40 1.70 -
abd. VIII 0.90 1.00 0.80 (0.90) 1.65 0.70 -
abd. IX 2.15 2.26 1.95 1.75 1.20 (1.55) 3.78 2.25 -
fwpd 0.57 0.63 0.57 (0.55) 0.49 0.45 0.76
mwpd 0.60 0.62 0.58 0.48 0.50 -
hwpd 0.36 0.44 0.37 (0.41) 0.26 ? 0.71
lingula 0.085 0.11 0.10 0.10 ? 0.10
forceps 1.35 1.45 1.32 1.25 0.80 (1.30) 1.60 1.20 n/a
fore f+t 1.35, 1.35 1.47, 1.45 1.35, - 0.93 (1.12) 0.85, 0.87 0.72 0.90
fore tibia 1.35, 1.37 1.49, 1.48 1.36, 1.36 1.09 (1.15) 0.82, 0.84 0.73 0.90
fore pts 0.38, 0.39 0.42, 0.42 0.37, 0.38 0.37 (0.42) 0.22, 0.23 0.28 0.32
fore dts 0.27, 0.29 0.30, 0.30 0.24, 0.28 0.26 (0.27) 0.25, 0.24 0.23 0.23
fore claw 0.09, 0.09 0.095, 0.09 - , 0.10 (0.08) 0.09, 0.09 0.08 0.08
mid. f+t 1.43, 1.43 1.55, 1.55 1.38, 1.38 1.12 (1.30) 0.95, 0.96 0.83 1.05
mid. tibia 1.45, 1.43 1.52, 1.52 1.42, 1.37 1.33 (1.40) 0.91, 0.91 0.95 1.00
mid. pts 0.38, 0.36 0.41, 0.40 0.39, 0.36 0.45 (0.50) 0.18, 0.20 0.29 0.33
mid. dts 0.26, 0.28 0.29, 0.27 0.26, 0.26 0.27 (0.28) 0.22, 0.23 0.23 0.22
mid. claw 0.09, - 0.09, 0.09 0.09, - 0.09, - 0.08 0.07
hind f+t 1.55, 1.50 1.60, 1.67 1.55, 1.54 1.15 (1.30) 0.97, 0.98 1.00 1.15
hind tibia 2.17, 2.17 2.18, 2.20 2.11, 2.09 1.97 (2.10) 1.29, 1.30 1.29 1.36
hind pts 0.45, 0.44 0.50, 0.51 0.46, 0.46 0.82 (0.88) 0.46, 0.48 0.64 0.51
hind dts 0.27, 0.27 0.28, 0.28 0.26, - 0.32 (0.32) 0.29, 0.27 0.27 0.25
hind claw 0.09, 0.09 0.085, 0.09 0.09, - 0.09, - 0.08 0.07
head W - - 1.00 - - 0.76
fch 0.20 - - n/a ?n/a [small]
ant. I 0.07 - 0.09, - 0.08 (0.08) 0.09, - 0.05 0.05
ant. II 0.15 0.15, 0.16 0.15, - 0.16 (0.18) 0.14, 0.14 0.15 0.15
ant. III 0.63, 0.65 0.71, 0.76 0.62, - 0.88 0.54, 0.50 0.58 0.49
ant. IV 0.31, 0.26 0.28, 0.30 0.26, - 0.35 0.28, 0.28 0.27 0.31
ant. V 0.12, 0.16 0.15, 0.165 0.14, - 0.28 0.18, 0.19 0.28 0.12
ant. VI 0.055 0.06, 0.055 0.06, - 0.17 0.07, 0.08 0.10 0.095
ant. VII 0.04 0.03, 0.035 0.03, - 0.13 0.07, 0.065 0.075 0.06
urs 0.43 0.41 0.40 (0.34) 0.33 0.34 0.34
vas no no no ? yes no no n/a

* Enderlein did not state whether this included the forceps
** Calculated from Enderleins wing-length measurements, assuming outlines are accurate
*** Body length measured in alcohol or in balsam, not stated if forceps included; sole male in BMNH in dissected
condition
**** Data from Visnya, 1941, with authors measurements of BMNH paratype in (bracketed italics)
Martin: Giant whiteflies and new Udamoselis 29
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habits and corresponding organization of differ-
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London, 587 pp.
Received: 11 October 2006
Accepted: 1 December 2006