A Revision of Aleurodicus Douglas With Two New Genera Proposed For Paleotropical Natives An Identification Guide To World Genera

ZOOTAXA
1835
A revision of Aleurodicus Douglas (Sternorrhyncha, Aleyrodidae),

with two new genera proposed for palaeotropical natives and
an identification guide to world genera of Aleurodicinae
JON H. MARTIN
Magnolia Press
Auckland, New Zealand
JON H. MARTIN
A revision of Aleurodicus Douglas (Sternorrhyncha, Aleyrodidae), with two new genera proposed for
palaeotropical natives and an identification guide to world genera of Aleurodicinae
(Zootaxa 1835)
100 pp.; 30 cm.
30 July 2008
ISBN 978-1-86977-225-3 (paperback)
ISBN 978-1-86977-226-0 (Online edition)
FIRST PUBLISHED IN 2008 BY

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ISSN 1175-5326 (Print edition)

ISSN 1175-5334 (Online edition)
Cover photo: Mature colony of Aleurodicus mirabilis (Cockerell) on Annona, Guyana, J. H. Martin
2 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Zootaxa 1835: 1–100 (2008) ISSN 1175-5326 (print edition)
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Copyright © 2008 · Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)
A revision of Aleurodicus Douglas (Sternorrhyncha, Aleyrodidae), with two new

genera proposed for palaeotropical natives and an identification guide to world
genera of Aleurodicinae
JON H. MARTIN
Department of Entomology, Natural History Museum, Cromwell Road, London SW7 5BD, UK. E-mail j.martin@nhm.ac.uk
Abstract
The genus Aleurodicus is discussed in the context of the whole subfamily Aleurodicinae, and is fully revised with 35 spe-
cies now included, 10 of them newly described here. The genus Lecanoideus Quaintance & Baker is here regarded as a
junior synonym of Aleurodicus syn. nov., and Aleurodicus fucatus Bondar is proposed as a junior synonym of A. capian-
gae Bondar syn. nov.. Palaealeurodicus gen. nov. is described to accommodate six native Asian species hitherto
included in Aleurodicus, along with one new species here described. Aleuroctarthrus gen. nov. is proposed to accommo-
date Aleurodicus destructor Mackie. Two species are here transferred from Aleurodicus to other genera. Puparial keys
are provided to the worldwide genera of Aleurodicinae, and to the described species of Aleurodicus and Palaealeurodi-
cus. Lectotypes are here designated for 11 species of Aleurodicinae, and a neotype for one species. A check list is pro-
vided to all the species treated in this paper. The paper is illustrated with 144 figures including 7 plates of habitus
photographs.
Key words: Aleurodicus, Palaealeurodicus, Aleuroctarthrus, whiteflies, new genera, new species, lectotypes, neotype,
check list, keys to genera and species
Table of contents
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Abbreviations for specimen depositories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Morphological terminology, with special reference to dorsal puparial pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Designation of primary type specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Check list of taxa treated in this paper. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Key to the world genera of Aleurodicinae – puparia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
ALEURODICUS Douglas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Key to Aleurodicus species – puparia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Aleurodicus species accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
PALAEALEURODICUS gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Key to Palaealeurodicus species – puparia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Palaealeurodicus species accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
ALEUROCTARTHRUS gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Aleuroctarthrus destructor (Mackie) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Revised generic placements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Illustrations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Accepted by C. Schaefer: 7 May 2008; published: 30 Jul. 2008 3

Introduction
Aleurodicus Douglas (1892) is an important whitefly genus, being the type for the subfamily Aleurodicinae
and including several pest species. The species with greatest notoriety as pests are A. dispersus Russell
(widely known as the spiralling whitefly), and A. dugesii Cockerell (often called the giant whitefly, although
its size is entirely typical for the whole Aleurodicus group). The genus has hitherto accommodated 33
described species (Martin & Mound, 2007). The great majority of these species were described from, and are
native to, the Neotropical Region. Most other genera of Aleurodicinae (13 of the 17 other genera regarded as
valid by Martin & Mound) are also neotropical natives. Amongst these neotropical genera, the present study
has indicated that Lecanoideus Quaintance & Baker should be regarded as a junior synonym of Aleurodicus
(syn. nov.) and its two included species, floccissimus (Martin, Hérnandez-Suarez & Carnero) and mirabilis
(Cockerell) transferred to Aleurodicus (comb. nov. and revived comb., respectively). Native to the Old World
are the aleurodicine genera Nipaleyrodes Takahashi [West Malaysia, Sulawesi, New Caledonia], Pseudosyn-
aleurodicus Gillespie [Australia], Stenaleyrodes Takahashi [Malagasian Region, eastern Africa] and Synaleu-
rodicus Solomon [Australia].
However, seven species hitherto placed in Aleurodicus were described from the Oriental or Austro-orien-
tal Regions and are native to the palaeotropics. It has been clear for some time that the generic positions of
these native palaeotropical Aleurodicus species require reappraisal (Martin, 1996: 1849; Martin & Watson,
1998: 86–87; Martin & Polaszek, 1999: 1546). Such reappraisal has been one main purpose of the present
paper and six of the seven palaeotropical species are here assigned to Palaealeurodicus gen. nov., along with
one new species here described. The seventh species, Aleurodicus destructor Mackie, is here placed in its own
monobasic genus, Aleuroctarthrus gen. nov..
In addition to revising the genus Aleurodicus, a major aim of this communication is to provide the means
for all of the world’s genera of Aleurodicinae to be distinguished one from another. A puparium-based key to
the aleurodicine genera, illustrated by many figures, is presented here (p. 11). The definition of the Aleurodic-
inae has been discussed by Martin & Streito (2003) and by Martin (2007).
Abbreviations for specimen depositories
BMNH – Natural History Museum, London SW7 5BD, U.K.

CDFA – California Department of Food and Agriculture, Sacramento, U.S.A.
INBio – National Biodiversity Institute, Santo Domingo de Heredia, Costa Rica
LNPV – Laboratoire National de la Protection des Vegetaux, Station d'Entomologie, 34060 Montpellier
cedex 01, France
MNHN – Museum National d’Histoire Naturelle, 75005 Paris, France
MOV – Museum of Victoria, Abbotsford, Melbourne, Australia
MZUSP – Museu de Zoologia da Universidade de São Paulo, Brazil
UCD – University of California, Davis, U.S.A., including the R.M. Bohart Museum of Entomology.
UG – University of Guyana, Turkeyen Campus, Georgetown, Guyana
USNM – U.S. Department of Agriculture, Beltsville, Maryland, U.S.A. (custodians of the Sternorrhyncha col-
lections of the U.S. National Museum of Natural History, Washington D.C.)
Materials & methods
This investigation is based primarily upon a study of the extensive whitefly collection of the BMNH. Type
material of several species not included in the BMNH collection had already been examined in connection

with a study of the Aleurodicinae of Belize (Martin, 2004), through the generous assistance of curators of sev-
eral other collections, notably MZUSP, UCD and USNM. Material of several species has been kindly made
available for this present study through the kindness of CDFA, UCD and USNM.
In addition to type material, USNM made available almost 700 slides of unidentified and partially identi-
fied Aleurodicinae for investigation, revealing material of two Aleurodicus species here described, as well as
many slides belonging to aleurodicine species described in the past few years: determinations for these speci-
mens were made as part of this study. Some taxa were labelled as belonging to an undescribed genus close to
Aleurodicus: these have in fact been recently described as Aleurodicus species, and they are here retained in
Aleurodicus (see A. niveus, p. 37, A. rugioperculatus, p. 41 and A. talamancensis, p. 43). Around 140 slides of
Aleurodicinae were also kindly made available from the CDFA collection, providing further useful data.
For new slide preparations, the author has employed the slide-making protocols detailed by Martin (2004:
84–86). These provide methods for temporary “quickmounts”, or for permanent Canada balsam mounts for
archival collections. The use of water-based mountants (Hoyer’s, Berlese, etc) for permanent preparations is
always to be avoided if possible.
The taxonomy employed here is almost exclusively puparium-based. The use of adult characters certainly
does hold promise for future studies (Martin, 2003), but is severely limited by a continuing dearth of adults
reliably associated with their puparia. Accordingly, adult features are only discussed in a few cases, including
where they provide supplementary features for distinguishing the taxa discussed here. Whenever it is possible,
the present author continues to urge collectors of whitefly material to retain (or, better still, to rear) adults that
are clearly associated with puparia. A much greater proportion of species needs to be represented by both
puparia and reliably associated adults before significant progress may be made with adult-based taxonomy.
Quoted distributional data are from three main sources: Mound & Halsey (1978), material in the collec-
tion of BMNH, and freshly identified specimens from the large batches of USNM and CDFA slides borrowed
for this study. Distribution data do not, therefore, reflect all records that may exist in the USNM or CDFA
main collections, or in other collections.
Almost all line drawings, and the great majority of photographs, are the work of the author. Exceptions are
clearly stated in the captions to figures, and individual photographers are also credited in Acknowledgements.
Figures 1–4, 5–8, 9–12, 13–16, 117–120 and several of the individual line drawings originally appeared in an
account of the Aleurodicinae of Belize (Martin, 2004): in figure captions, therefore, the absence of a quoted
country of origin indicates drawings or photographs featuring Belize material. Many of the author’s other
original line drawings used here have previously appeared within plates in earlier publications, especially in
Zootaxa and Journal of Natural History, several of them having been modified here.
Morphological terminology, with special reference to dorsal puparial pores
Puparial descriptions and discussions presented here generally follow the terminology detailed by Russell
(1965), Martin (1987) and Gill (1990), with some characters unique to the Aleurodicinae following Iaccarino
(1990) and Quaintance & Baker (1913). Figure A (p. 6) depicts a typical New World aleurodicine puparium,
Aleurodicus pulvinatus (Maskell), specially annotated to illustrate the main characters (modified after Martin
& Watson, 1998). Puparial taxonomy of aleurodicine taxa often utilises the form and distribution of dorsal
pores and figure B (p. 8) depicts the major types of simple dorsal pore discussed in this paper, including in the
key to aleurodicine genera.
MARTIN Zootaxa 1835 © 2008 Magnolia Press · 5

FIGURE A. Aleurodicus pulvinatus (Maskell) – dorsal detail of puparium, annotated with general anatomical terminol-
ogy used in descriptions and keys.

COMPOUND PORES
The most obvious of the dorsal pores are the so-termed compound pores. These are comparatively large,
even in their smallest forms, and are sometimes very large indeed (see figs 90, 110). They are usually deeply
invaginated from the plane of the dorsal surface, with a ring of loculi at the bottom of the invagination, these
loculi termed “spinneret cells” by Quaintance & Baker (1913). Such “deep” compound pores usually each
secrete a conspicuous tubular filament of waxy material which issues from the pore, with tangles of such fila-
ments often obscuring colonies of feeding nymphs (Figs 126, 128). In most aleurodicine species each com-
pound pore has a conspicuous axial process (“central process” of Russell, 1965), often extending far beyond
the pore opening at the dorsal surface (see figs A, 2, 54, 55, 81, 83 etc.). The axial process appears to guide the
pore’s waxy filament as it is extruded; axial processes vary considerably in their degree of visibility on slide-
mounted specimens, and this may reflect varying resistance to the chemicals used in slide-making, perhaps
combined with maceration time. At least some species that are usually depicted without axial processes (e.g.
Aleuroctarthrus destructor, fig. 90) do possess them, and they can be seen in some mounts (Fig. 92). How-
ever, in some members of the Aleurodicinae (e.g. Paraleyrodes species, fig. 100, Kaieteurodicus cereus Mar-
tin, fig. 110) there is no axial process, but separate chitinous “splines” issue from the pore-base loculi, and
these appear to fulfil the same purpose, of guiding the growing filament (Martin, 1996: 1854). Some palaeo-
tropical taxa possess compound pores that differ greatly from the forms discussed above (see next paragraph,
and key to genera on p. 11).
Typically, the puparia of aleurodicine species have one cephalic pair and up to six abdominal pairs of
compound pores, subdorsally located (Figs A, 1–12), although some genera and/or species have very few (e.g.
Fig 95), or none at al (Figs 109, 111, 112). The presence of compound pores in second- and third-instar
nymphs is a useful taxonomic character, and their absence in the third-instar does not imply absence or sparse
provision in the puparial stage (e.g. Kaieteurodicus spp).
The palaeotropical species Nipaleyrodes elongata Takahashi (1951) possesses more than 15 large submar-
ginal compound pores in a single row on each side of the puparium (Fig. 96), but these are of a rounded-
polygonal form that is apparently unique amongst the Aleurodicinae, as well as being much more numerous
and situated much closer to the margin than the compounds in other aleurodicine genera. Takahashi mistak-
enly described and illustrated the compound pores of N. elongata as octagonal but type specimens in BMNH,
remounted and examined in connection with this study (see p. 10), clearly possess compound pores that are
pentagonal or hexagonal, and this is also seen (but more subtly) in material from Sulawesi used to prepare fig-
ure 96. Puparia of an undescribed species of Nipaleyrodes (one slide, New Caledonia, coll. Krauss, BMNH)
have much more numerous, and smaller, pores in several ranks, and these have more of an appearance of
being large simple pores although seemingly of the same structure as the larger pores seen in N. elongata.
AGGLOMERATE PORES
As the term suggests, these are aggregations of pores, and together they form units that may be paired on
either side of the body and can, thus, be ennumerated in the same way as can compound pores. This category
of pore is seen in two forms, in two geographically and taxonomically disparate groups.
In the Australian genus Synaleurodicus Solomon (1935) each agglomerate pore comprises a small group
of pore openings clustered within a rounded plate (Figs B xiv, 102b, 103), eight such agglomerate pores being
located subdorsally on either side of the body (Fig. 102). The agglomerate pores of Synaleurodicus species
may be tracheal in function, rather than secretory (Martin, 1999: 21, and see ducts illustrated here in figure
103).
In the neotropical genus Leonardius Quaintance & Baker (1913), and in Ceraleurodicus assymmetrus
(Bondar, 1922), each agglomerate pore comprises a circular or ovoid porous area of very fine glandular gran-
ules. In Leonardius species (Fig. B xiii, 116) the agglomerate pore area also accommodates a few bright sim-
ple pores, and the compound pores are surrounded by some of the agglomerate porous areas (Fig. 116).

FIGURE B. Major types of dorsal simple pores (includes types not seen in Aleurodicus but discussed in text). i-v usually
submarginal zone – i, modified 8-shaped, just within puparial margin; ii, wide-rimmed, usually in a distinct submarginal
band but occasionally scattered on dorsal disc; iii, double-rimmed septate / 8-shaped; iv, double-rimmed concentric; v,
double-rimmed notched: vi-xi usually subdorsal and submedian zones – vi, notched, usually numerous and mesal to
compound pores; vii, angular septate (not Aleurodicus); viii & ix, septate, may be extremely small; x, loculate, always
extremely small (seen in several neotropical genera but not Aleurodicus); xi, asymmetric 8-shaped: xii-xiv subdorsal
zone of dorsal disc – xii, bright, usually in small clusters adjacent to compound pores or vasiform orifice; xiii, Leonar-
dius-type agglomerate pores; xiv, Synaleurodicus-type agglomerate pores.

SIMPLE PORES
These are important characters that can be used in grouping, and distinguishing between, aleurodicine
taxa (both genera and species). Within the Aleurodicinae simple pores are all very much smaller than com-
pound pores. However, puparia in one Asian genus of Aleyrodinae (Dialeuropora Quaintance & Baker, 1917)
usually possess five pairs of exceptionally large “simple” pores that are similar in size to the compound pores
seen in Aleurodicus and other aleurodicine genera – see comments on Aleurodicus (p.15). Russell (1965) rea-
lised the taxonomic importance of the simple pores. These had first been mentioned by Quaintance & Baker
(1913), but it was Russell who was the first to divide them into categories and provide names for them, as part
of her description of A. dispersus. Russell defined four types of simple pore in her paper of 1965. In the
present paper, figure B (p. 8) depicts the main forms of simple pore that have been recognised as part of this
study, and Russell’s names have been amended slightly.
Unfortunately, deciding on the category of a simple pore is not always easy, with a number of factors
affecting the clarity of the observed image of a particular specimen on a particular slide, viewed through a par-
ticular microscope. However, the expected locations for the different types of pore are indicated in figure B
and this will, it is hoped, further aid their recognition.
Designation of primary type specimens
In the author’s experience, syntypes of species described by Bondar and Hempel, on original slides (mostly
still residing in Brazil), are usually extremely poor preparations. Slides of several species were kindly loaned
to the author by MZUSP in connection with a study of Aleurodicinae in Belize (Martin, 2004). Most were
extremely difficult to study because of the condition of the mountant, and this was combined with an obvious
lack of maceration prior to slide-mounting of the specimens, resulting in waxy secretions obscuring even
more detail. It is understandable that the custodians of these original slides are unwilling to risk attempts to
remount. However remounting would, in the author’s opinion, reveal much valuable detail and enhance the
value of the specimens concerned – see discussion of A. jamaicensis, p. 33.
Martin (2004: 56) discussed material of 25 species that Bondar sent to BMNH in 1923, concluding that
such samples were probably syntypic. Further investigation of the background to the donation to BMNH, and
the discovery that similar donations were made to USNM by Bondar, leads the author to the conclusion that
these samples are indeed syntypic. Given the syntypic status of this duplicate material that was sent (it is
believed that it was then dry and unmounted) by Bondar to BMNH and to USNM, and given the condition of
original slides and the fact that remounting is not permitted, the designation of lectotypes from BMNH or
USNM duplicate material is desirable, and is helpful in stabilising such species through the revealing of finer
characters. Lectotypes of two of Bondar’s species are here designated, along with one Costa Lima species and
eight other species described by other authors: material bearing identical data has paralectotype status, wher-
ever it resides.
In one neotropical species treated here, Aleurodicus antillensis Dozier, no material bearing original data
has been traced. However, suitable topotypic material has been examined and compared with the description
and other data, enabling the selection of a neotype specimen. The purpose is the same as for lectotype desig-
nation – to provide the stability offered by the existence of a type specimen that is in good enough condition
for future comparative examination.
In the Old World, specimens of species of Aleurodicinae described by Takahashi were also originally
mounted by simply placing many untreated specimens onto one slide and pressing a coverslip down onto
them. The resulting mounts were unnecessarily thick, and most specimens were poorly displayed. Original
Takahashi slides are slightly better than the Bondar / Hempel examples discussed above, because the moun-
tant itself is Canada balsam that is still in good condition. The opportunity has been taken here to remount

type material in BMNH of two of Takahashi’s species, Aleurodicus cinnamomi (here transferred to the new
genus Palaealeurodicus, see discussion on p.49) and Nipaleyrodes elongata.
The 11 puparia on the original type slide of Nipaleyrodes elongata in BMNH were remounted as part of
this study: 10 of them are parasitised, but one parasitised specimen nevertheless displays almost all characters
very well, especially ventral details, and this has been here designated the lectotype. Paralectotype puparia of
N. elongata are now distributed between six new slide preparations of which one has been presented to
USNM.
Check list of taxa discussed in this paper
ALEURODICUS Douglas
antillensis Dozier, neotype here designated
araujoi Sampson & Drews, syntypes examined
capiangae Bondar, lectotype here designated
charlesi sp. nov.
chirripoensis sp. nov.
clusiae sp. nov.
coccolobae Quaintance & Baker, lectotype here designated
cocois (Curtis), lectotype designated by Martin, 1997
darienpalmae sp. nov.
decemensis sp. nov.
dispersus Russell, paratypes examined
dugesii Cockerell, lectotype here designated
etiennei sp. nov.
flavus Hempel, probable syntypes examined
floccissimus (Martin, Hérnandez-Suarez & Carnero) comb. nov. , holotype examined
guppyii Quaintance & Baker, syntypes examined
inversus Martin, holotype examined
jamaicensis Cockerell, holotype examined
juleikae Bondar, lectotype here designated
magnificus Costa Lima, lectotype here designated
maritimus Hempel, syntypes examined
mirabilis (Cockerell), revived combination, syntypes examined on author’s behalf
neglectus Quaintance & Baker, lectotype here designated
nicaraguensis sp. nov.
niveus Martin, holotype examined
ornatus Cockerell, lectotype here designated
pauciporus Martin, holotype examined
pseudugesii Martin sp. nov.
pulvinatus (Maskell), lectotype designated by Martin & Watson, 1998
rugioperculatus Martin, holotype examined
septatissimus sp. nov.
spectabilis sp. nov.
talamancensis Martin, holotype examined
trinidadensis Quaintance & Baker, lectotype here designated
vinculus Martin, holotype examined

PALAEALEURODICUS gen. nov.
antidesmae (Corbett) comb. nov., no material available for examination
borneensis sp. nov.
cinnamomi (Takahashi) comb. nov., lectotype here designated
holmesii (Maskell) comb. nov., lectotype designated by Martin, 1999
indicus (Regu & David) comb. nov., no type material available for examination
machili (Takahashi) comb. nov., type material not examined
wallaceus (Martin) comb. nov., holotype examined
ALEUROCTARTHRUS gen nov.
destructor (Mackie) comb. nov., neotype designated by Martin, 1999
SPECIES HERE EXCLUDED FROM ALEURODICUS
Bakerius marmoratus (Hempel) comb. nov., no puparial type material available for examination
Metaleurodicus essigi (Sampson & Drews) comb. nov., lectotype here designated
LECTOTYPE DESIGNATION HERE BUT NOT DISCUSSED IN DETAIL
Nipaleyrodes elongata Takahashi, lectotype here designated
Key to the world genera of Aleurodicinae – puparia
• This key includes all genera of Aleurodicinae, as understood by the present author. Genera that are natives
of the New World are indicated in bold italics, the others in italics only.
• Quoted counts of submarginal setae include the nominal caudal pair, but not the anterior and posterior
marginal setae.
• Note that the pair of posterior abdominal spiracles (ventral, abdominal segment VIII) can be mistaken for
small post-vasiform orifice compound pores (dorsal). Figure 3 is a good example, being without any small
compound pores, but with the spiracles clearly visible in a similar position.
1 Puparia with agglomerate (Figs B xiii & xiv, 102b, 116) or compound (Fig. A) pores present, although
sometimes much reduced in number and/or size (e.g. fig. 95) ....................................................................2
- Puparia with neither agglomerate nor compound pores (Figs 109, 111, 112), but sometimes with loose
clusters of simple pores (Fig. 107). ...........................................................................................................19
2 With 15+ small compound pores on each side of body, distributed in a single submarginal row from ceph-
alic region to beyond vasiform orifice (Fig. 96). Longitudinal moulting suture reaching anteriorly only to
a point in cephalic region, where a pair of cephalothoracic moulting sutures diverge towards edge of
puparium and then curve posteriorly into thoracic submargin .................... Nipaleyrodes Takahashi, 1951
[N. elongata Takahashi, but see discussion on p. 7]
- With up to 7 pairs of compound pores (Fig. A), or up to 8 pairs of agglomerate pores (Fig 102, 116). Lon-
gitudinal moulting suture reaching anteriorly to puparial margin (Fig. A), or entirely absent (Fig. 102) ...3
3 With 8 pairs of small agglomerate pores (Fig. B xiv) present, in an evenly-spaced arc from cephalic region
to abdominal segment VII/VIII, lateral to vasiform orifice (Fig. 102); without compound pores. Longitu-
dinal moulting suture absent (Fig. 102)........................................................Synaleurodicus Solomon, 1935
- With up to 7 pairs of compound pores present (Fig. A), sometimes with associated large agglomerate
porous areas (Fig. 116). If cephalothoracic compound pores present then only a single cephalic pair (as
fig. A), or one on one side only (Fig. 113), with no further compound pores present anterior to transverse
moulting sutures. Longitudinal moulting suture present ..............................................................................4
4 With 1–4 pairs of submedian cephalothoracic setae present. Note: these setae are usually obvious (Figs
54, 55, 63, 65, 67, 69, 100 etc.), but they are sometimes very small (Figs 71, 90, 114) ..............................9

- Submedian cephalothoracic setae entirely absent (Figs 73, 77, 80–83, 94, 97). .........................................5
5 Legs without apical claws (Figs 78, 81, 82, 86); 8th abdominal setae absent; usually with only 0–2 pairs of
submarginal setae (if present they are posterior to vasiform orifice, fig. 83) [one species has 12 pairs, fig.
81]; with few dorsal simple pores (of one type only) or none discernible; margin usually conspicuously
dentate (Figs 76–78, 80, 82, 83) ........................................................................ Palaealeurodicus gen. nov.
- Each leg with a pronounced apical claw (Figs 94, 97) or (in a few Ceraleurodicus species) an apical seta;
8th abdominal setae present anterior or anterolateral to vasiform orifice (Figs 74, 94); with 12 or more
pairs of dorsal submarginal setae (Figs 94, 97); usually with many dorsal simple pores, often of several
types; margin usually smooth to irregular ....................................................................................................6
6 With 12 pairs of dorsal submarginal setae (Figs 94, 97). ........................................................................... 7
- With 15 pairs of submarginal setae. [On closer examination, such specimens will usually prove to have
very small cephalothoracic submedian setae present (Figs 113, 114) – see couplet 4]................................9
7 Cephalic pair, and 5 abdominal pairs of compound pores present, all subequal in size (40–65 μ in outer
diameter), the abdominal pairs evenly spaced on segments III–VII (Figs 94, 120); compound pores each
with a protruding, dagger-shaped axial process. Submargin with large double-rimmed pores of two
types—concentric (Figs B iv, 94c) and 8-shaped (Figs B iii, 94a); dorsal disc with angular septate pores
(Figs B vii, 94b)........................................................ Azuraleurodicus Martin in Martin & Polaszek, 1999
- Cephalic pair and 4–6 abdominal pairs of compound pores present (Figs 97, 117, 118); usually at least
some compound pores much smaller than 40 μ, without a protruding axial process; sometimes abdominal
compound pores not all evenly spaced (Figs 117, 118). If submargin with double-rimmed pores they are
not of the concentric type ...............................................................................................................................
......................................................................................................................................................................8
8 Abdominal compound pores distributed in a smooth arc, almost concentric with curve of puparial margin,
with at least posteriormost 4 pairs evenly spaced (Fig. 97)..... Metaleurodicus Quaintance & Baker, 1913
- Abdominal compound pores not all distributed in a smooth arc, at least 1 of the posteriormost 4 pairs inset
or offset; spacing between the abdominal compound pores not even (Figs 117, 118) .................................
....................................................................................................................... Aleuronudus Hempel, 1922a
9 Pair of cephalic setae present but thoracic submedian setal pairs absent (Fig. 100). Cephalic and posterior
4 pairs of abdominal compound pores larger than anterior 1 or 2 abdominal pairs. With 14 pairs of sub-
marginal setae. [Adult forewing with R simple (Fig. 104).] ...................... Paraleyrodes Quaintance, 1909
- Cephalic setal pair present or absent but thoracic submedian setae present in 2 or 3 pairs (Figs 54, 55, 67,
69, 71, 101, 110, 114, 115 etc). Compound pores present in a different combination. Usually with 12, 13
or 15 pairs of submarginal setae. [Adult forewing with R forked (Figs 138–140).] .................................10
10 With 1 cephalic pair and 6 abdominal pairs of ovoid porous agglomerate pores (Figs B xiii, 116) present.
Compound pores present, comprising a pair on each of abdominal segments III & IV, and sometimes also
a cephalic pair, each situated within a larger agglomerate pore area (as in fig. 116) .....................................
........................................................................................................ Leonardius Quaintance & Baker, 1913
- Paired agglomerate pores absent, but occasionally one such porous area present on one side of puparial
abdomen (in Ceraleurodicus assymmetrus Bondar)...................................................................................11
11 Compound pores (Fig. 95) only present on abdominal segment VIII (each possessing a seta-like axial pro-
cess long enough to reach puparial margin) and on one or both sides of abdominal segment III (without a
long and seta-like axial process). With 13 pairs (bodkini) or 15 pairs (undescribed sp, fig. 95, BMNH) of
submarginal setae.................................................................... Eudialeurodicus Quaintance & Baker, 1915
- When compound pores present on abdominal segment VIII, then axial processes much shorter, never
more than twice pore diameter in total length. With at least one cephalic compound pore often present.
With 12, or with more than 13, pairs of submarginal setae ........................................................................12
12 Lingula with 2 stout setae clearly visible (Fig. 90), with a much smaller second pair occasionally visible

under the highest magnification (Fig. 91). With 6 pairs of very large abdominal compound pores, all simi-
lar in size (largest 80–100 μ) and distinctly larger than the cephalic pair (Fig. 90). [Adults with 8-seg-
mented antennae.] ................................................................................................. Aleuroctarthrus gen. nov.
- Lingula with 4 stout setae (Figs 21, 22, 25, 27, 33 etc). [Some species have the lingula included within the
vasiform orifice (Figs 105, 108, 109, 110, 115) with the anterior pair of setae sometimes difficult to
resolve at lower magnifications.] If with 6 pairs of abdominal compound pores that are all similar in size,
then they are much smaller and are similar in size to the cephalic pair (Fig. 119). [All known adults with
7-segmented antennae.] .............................................................................................................................13
13 With 2 pairs of cicatrices on dorsum of thorax (Figs A, 41, 54, 55, 67, 69 etc). Cephalic pair of compound
pores similar in size to those on each of abdominal segments III–VI (4 pairs, figs 67, 69, 71, 72 etc) or III–
VIII (6 pairs, fig. 119); if with only 4 abdominal pairs similar in size to cephalic pair, then usually with 1
or 2 much smaller pairs on segment VII and/or VIII (Figs A, 69, 72 etc), but sometimes without these
smaller pairs (Figs 13, 70, 71). With 12 pairs of submarginal setae. .........................................................14
- Cicatrices absent from dorsum of thorax (Figs 101, 110, 114, 115). If cephalic compound pores present,
paired, and similar in size to only 4 other pairs, on abdominal segments III–VI, then puparium usually
with 15 pairs of submarginal setae .............................................................................................................15
14 Cephalic pair and 6 abdominal pairs of compound pores, on segments III–VIII, all similar in size and
form, the abdominal pairs describing an even arc on each side of puparium (Fig. 119); anterior marginal
setae usually visible, even if very small ..................................................... Austroaleurodicus Tapia, 1970
- Cephalic pair of compound pores similar in size and form only to the 4 pairs forming an arc on each side
of abdominal segments III–VI (Figs 67, 69, 71, 72 etc); an additional 1 or 2 pairs often present, of differ-
ent form and usually much smaller than others, situated on abdominal segments VII and/or VIII (Figs A,
54, 55, 57, 67, 69, 72); anterior marginal setae absent .................................... Aleurodicus Douglas, 1892
15 Submarginal setal bases set almost on the rather thickened (and often slightly down-curved) puparial mar-
gin (Fig. 101), the setae robust, almost entirely reaching beyond edge of puparium. Puparia symmetrical
in compound pore provision and, usually, also in outline ........................... Nealeurodicus Hempel, 1922b
- Submarginal setal bases inset from the much more planar (flatter) margin, the setae finer and shorter, not
or hardly overlapping puparial margin at all (Figs 110, 114, 115). Puparial outline sometimes distinctly
asymmetrical (Fig. 113). .............................................................................................................................16
16 With 12 pairs of submarginal setae. Puparia slightly asymmetrical in outline but compound pore provision
symmetrical, with cephalic pair and 5 abdominal pairs which appear to be located on segments II and IV–
VII (Fig. 110). Puparia without radial “rays” leading from margin towards subdorsum ...............................
....................................................................................................................... Kaieteurodicus Martin, 2008
- Normally with 15 pairs of submarginal setae (Figs 113, 114, 115). Puparia may be noticeably asymmetri-
cal in outline and/or compound pore provision (Figs 113, 134). Puparia often with up to 9 pairs of radial
“rays” (Figs 113, 134), some of which may be associated with ventral tracheal folds .............................17
17 Compound pores comprising only a cephalic pair and one pair on each of abdominal segments III & IV
(Fig. 115); when undamaged each compound pore with a dagger-shaped axial process present, this pro-
cess similar in length to maximum pore diameter, not reaching puparial margin. Puparial outline symmet-
rical .......................................................................................................................... Bakerius Bondar, 1923
- Compound pores rarely present and paired on cephalic segment and abdominal segments III & IV only
but, if so, compound pores either without evident axial processes, or with their axial processes exception-
ally long and reaching to or beyond the puparial margin. Puparial outline often asymmetrical (Fig. 113) ...
....................................................................................................................................................................18
18 Asymmetry (when evident) affecting puparial outline only; abdomen with 4 pairs of submedially situated,
longitudinally aligned, compound pores on segments III–VI (Fig. 114). Cephalic and post-vasiform ori-
fice compound pores absent........................................................................ Octaleurodicus Hempel, 1922a

- Asymmetry (when evident) often affecting compound pore distribution as well as puparial outline (Figs
113, 134). Provision of compound pores very variable, but usually with at least one cephalic pore and/or
one pore present posterior to vasiform orifice; if compound pores present on each of abdominal segments
III–VI then these not longitudinally aligned in submedian area................. Ceraleurodicus Hempel, 1922a
19 Each leg with an apical claw (Fig. A). Lingula completely included within vasiform orifice (Fig 105,
108), with 4 stout setae usually distinct but sometimes lingular setae not readily discernible. With radial
“rays” (Figs 106, 107) leading from margin towards subdorsum, up to 9 pairs evident (Fig. 109)...............
..................................................................................................................... Dialeurodicus Cockerell, 1902
Legs with or without apical claws. Lingula included within, or extending beyond, vasiform orifice but
clearly with only 2 stout setae (Figs 111, 112). Without radial “rays” (but intersegmental folds may almost
reach margin as in fig. 112) ........................................................................................................................20
20 Lingula included within vasiform orifice (Fig. 111). Each leg rounded apically, without an apical claw or
seta. Antennae long and curved, reaching at least middle pair of legs, corrugate but unsegmented (Fig.
111) .............................................................................................................. Stenaleyrodes Takahashi, 1938
- Lingular apex just excluded beyond boundary of vasiform orifice (Fig. 112). Each leg with an apical claw
or fine seta. Antennae short but multisegmented, closely adjacent to front legs (Fig. 112 expanded detail)
......................................................................................................... Pseudosynaleurodicus Gillespie, 2006
ALEURODICUS Douglas
Aleurodicus Douglas, 1892: 32. Type species Aleurodicus anonae Morgan, 1892: 32, by subsequent designation by
Quaintance, 1908: 8. [Synonymised with A. cocois Curtis (1846: 284–285) by Mound & Halsey, 1978: 228.]
Aleurodicus (Lecanoideus) Quaintance & Baker, 1913: 70. Type species Aleurodicus (Lecanoideus) giganteus Quain-
tance & Baker, 1913: 70, by original designation. [Synonymised with Aleurodes mirabilis Cockerell (1898: 225) by
Martin, 2004: 18.]
Lecanoideus Quaintance & Baker; as full genus, Costa Lima, 1928: 133 (by inference). Syn. nov.
DIAGNOSIS.
PUPARIA—characters of most diagnostic importance are given in bold. As interpreted here, Aleurodicus
comprises species displaying the following combination of characters (Fig. A): 12 pairs of submarginal
setae present (including the nominal caudal pair); submedian cephalothoracic setae present (some species
with cephalic and three thoracic pairs, all similar to submarginal setae, others with cephalic pair wanting, and
the dispersus / flavus / coccolobae / charlesi assemblage usually possess only meso- and metathoracic pairs of
submedian setae which are also much smaller than submarginal setae); anterior marginal setae absent; sub-
margin and/or dorsal disc usually punctuated by pores of several types, but loculate pores (Fig. B x)
absent; 4 pairs of large subdorsal abdominal compound pores, evenly spaced on segments III – VI, usually
each with an axial process extending beyond the pore mouth; most species also additionally with one or two
much smaller abdominal pairs of compound pores, on segments VII and/or VIII; two pairs of cicatrices
present on thoracic area (scars of third-instar compound pores); lingula always protrudes beyond vasiform
orifice, its four subapical setae on the protruding part; ventrally, each leg bears a large and distinct claw,
antennae long, curved and reaching at least to middle legs, often to hind legs (Figs 69, 71). [2nd-instar
with 3 pairs of compound pores, on cephalus, prothorax and last abdominal segment; 3rd-instar with 3
pairs of cicatrices in same positions.]
ADULTS—with 7-segmented antennae; forewings with R forked (Figs 138–140); antennae of males
only with tiny and discreet sensoria; abdominal wax plates numbering 4 pairs in females (Fig. 141) and
3 pairs in males.

COMMENTS. Confusion over the identity of the type species of Aleurodicus was investigated by Martin
(1997), confirming it to be Aleyrodes cocois Curtis—see comments on Aleurodicus cocois, p. 26.
Species with the characters detailed in the diagnosis above, but whose puparial margins are usually vari-
ably deflexed, the submargin punctuated by a broad band of crowded simple pores of only the wide-rimmed
type (Figs 69, 70), and whose compound pore axial processes are usually directed mesally when slide-
mounted, have hitherto been placed in the genus Lecanoideus Quaintance & Baker (1913) – see Martin, 2004:
16, key couplet 10. Lecanoideus was raised to a full genus by Costa Lima (1928), but with no reasons stated.
Lecanoideus was discussed by Martin et al. (1997: 1269–1270), who concluded that there was little justifica-
tion for its status as a full genus, but who decided to preserve the status quo pending further investigations.
Subgenera are not generally accepted by whitefly workers (Martin & Mound, 2007: 5), precluding that option
for Lecanoideus, in the author’s opinion. Aleurodicus inversus (described from Belize) is somewhat interme-
diate in form, reinforcing the decision, here, to regard Lecanoideus Quaintance & Baker (1913) as a junior
synonym of Aleurodicus (1892) syn. nov..
Amongst extensive material that had remained unidentified within the USNM collection, prior to being
loaned to the author for this study, were three species labelled as belonging to a new genus closely related to
Aleurodicus. These three species are included in Aleurodicus (as defined by Martin, 2004, and followed here),
and all three had been recently described – as A. niveus Martin (2004), A. rugioperculatus Martin (2004) and
A. talamancensis Martin (2005). These three species belong to two discreet groups within Aleurodicus (see
key couplet 30, p. 19) in the same way as do members of the A. dispersus-group (see diagnosis, above), but
there is no evidence to support the proposal of a new genus, in the author’s opinion – echoing the situation
with Lecanoideus discussed above. Notably, A. niveus had been intercepted on imported orchids for many
decades, while remaining undescribed (see discussion of A. niveus, p. 38).
The five pairs of exceptionally large simple pores observed in most species of a palaeotropical aleyrodine
genus, Dialeuropora Quaintance & Baker, led Chen (1997) to describe a new species from China as Aleurod-
icus photiniae. However, the even spacing of the five pairs of large pores in puparia of most species of Dialeu-
ropora is completely different from the cephalic pair + separate abdominal grouping seen in Aleurodicus
species, and photiniae was transferred from Aleurodicus to Dialeuropora by Martin & Mound (2007): speci-
mens bearing type data (discovered in BMNH accessions) have confirmed this decision.
Key to Aleurodicus species – puparia
• Note that the pair of posterior abdominal spiracles (ventral) can be mistaken for small post-vasiform ori
fice compound pores (dorsal). Figure 3 is a good example, being without any small compound pores, but
with the spiracles clearly visible in a similar position.
• An attempt has been made, here, to deal with the species in natural species groupings where possible.
These groupings may need to be reappraised if molecular or numerical studies so indicate, but this should
not affect the working of the key.
1 Cephalic and anterior 4 pairs of abdominal compound pores with long, acute axial processes that extend
far beyond puparial margin (Fig. 72). Dorsal disc punctuated by un-grouped simple pores of 3 types—
wide-rimmed, symmetric 8-shaped septate, and asymmetric 8-shaped septate (Fig. B ii, iii, xi)
..................................................................................................................................... decemensis sp. nov.
- Compound pore axial processes always much shorter, or sometimes inevident. Dorsal disc without asym-
metric 8-shaped septate pores.......................................................................................................................2
2 Submargin with double-rimmed pores that may be septate (Fig. B iii) or notched (Fig. B v): these may be
very small and situated adjacent to, or amongst, simple pores of other types (Figs 21, 22, 25, 43, 44, 46),

and may be present only at posterior end of puparium (Fig. 52)..................................................................3
- Submargin entirely without double-rimmed pores .....................................................................................22
3 Subdorsum with numerous large, crowded, prominent simple pores (Figs 13, 45, 46, 50, 58, 59, 71) that
are usually of the notched type (Fig. B vi) ...................................................................................................4
- Subdorsum with simple pores of the septate type (Fig. B ix, 48, 49, 54, 55 etc), not prominenent, usually
sparsely distributed and often extremely small ........................... capiangae / dugesii species-groups ... 10
4 With 4 pairs of submedian cephalothoracic setae, long and fine; with 6 pairs of abdominal compound
pores.......................................................................................................... magnificus species-group. ….. 5
- With 2–3 pairs of submedian cephalothoracic setae, usually very small and fine (Fig. 71); usually with
only 4 or 5 pairs of abdominal compound pores ........................................... dispersus species-group .... 6
5 Each cephalic compound pore surrounded by a cluster of notched pores (Fig. 59); density of abdominal
notched pores on dorsal disc increasing posteriorly .................................... neglectus Quaintance & Baker
- With a cluster of notched pores posterior to each cephalic compound pore, but not surrounding it (Fig.
58); density of abdominal notched pores on dorsal disc more even........................ magnificus Costa Lima
6 With 6 pairs of abdominal compound pores, posterior 2 pairs extremely small (Fig. 51); submarginal pore
band comprises only very large double-rimmed pores that are 8-shaped septate (Fig. B iii), present in 3
ranks (Figs 50, 51); submarginal setal bases lateral to pore band (Fig. 50); dorsal disc with numerous sim-
ple pores that are most clearly of the notched type submedially ................................... spectabilis sp. nov.
- With 4 or 5 pairs of abdominal compound pores; submarginal pore band comprises double-rimmed pores
that are much smaller, notched (Fig. B v, 46, 71), surrounded by wide-rimmed pores; submarginal setal
bases situated within the pore band ..............................................................................................................7
7 With only 4 pairs of abdominal compound pores (Fig. 71)..........................................................................8
- With 5 pairs of abdominal compound pores, the small 5th pair on abdominal segment VII (Figs 45, 46) ...9
8 Band of notched pores continuous posterior to vasiform orifice, but notched pores absent from median
part of abdominal segment VII between pockets and segment VI/VII division (Fig. 71)..............................
.......................................................................................................................................... dispersus Russell
- Band of notched pores interrupted posterior to vasiform orifice (Fig. 13), but a few notched pores present
on median part of abdominal segment VII between pockets and segment VI/VII division...........................
.................................................................................................................. coccolobae Quaintance & Baker
9 Pair of compound pores on abdominal segment VII extremely small, only about twice size of a notched
simple pore (Fig. 45); notched pores not extending from dorsal disc into outer subdorsum beyond arc of
compound pores; large abdominal compound pores all similar in size................................. flavus Hempel
- Pair of compound pores on abdominal segment VII more than half size of pair on segment VI (Fig. 46);
notched simple pores extending from dorsal disc, beyond arc of compound pores and into outer subdor-
sum (Fig. 46); compound pores on abdominal segments V and VI significantly smaller than those on seg-
ments III and IV.................................................................................................................. charlesi sp. nov.
10 Only a few submarginal double-rimmed pores present, posterior to the vasiform orifice (Fig. 52); only 5
pairs of abdominal compound pores present, with none on segment VIII; submargin with a broad band of
crowded wide-rimmed pores, with the submarginal setal bases lining the mesal boundary of this band;
with 4 pairs of submedian cephalothoracic setae. ........................................................... antillensis Dozier
- Submarginal double-rimmed pores present around the circumference of the puparium. ..........................11
11 With only 5 pairs of abdominal compound pores (Fig. 53); submarginal double-rimmed septate pores
evenly-spaced in a single row.......................................................................................... ornatus Cockerell
- With 6 pairs of abdominal compound pores; submarginal double-rimmed septate pores may be evenly-
spaced (Figs 47, 55, 56) or irregular (Figs 22, 26, 31, 48, 49.....................................................................12
12 With 2 ranks of large double-rimmed septate pores (Figs 48, 49), outer rim of each pore contiguous with
pores in adjacent rank and with neighbouring pores in same rank; remainder of dorsal disc with numerous

septate pores (Fig. B ix) which in abdomen are interspersed with smaller wide-rimmed pores (Fig. 49).....
................................................................................................................................. septatissimus sp. nov.
- If submarginal double-rimmed pores nearly contiguous, then in a single rank (Figs 21, 31, 56); if double-
rimmed pores in 2 ranks then the pores not contiguous (Fig. 54); dorsal disc septate pores usually more
sparsely distributed and often smaller (Figs 54, 55).................................................................................. 13
13 With a cluster of bright simple pores anteromesal to small abdominal compound pores on abdominal seg-
ment VII or segments VII and VIII (Figs. 21, 44)......................................................................................14
- Without clusters of bright pores adjacent to small abdominal compound pores ........................................15
14 Bright pores only on abdominal segment VII (Fig. 21); submarginal double-rimmed pores large and sep-
tate, in an uneven row, surrounded by tiny wide-rimmed pores (Fig. 21) ........ araujoi Sampson & Drews
- Bright pores on abdominal segments VII and VIII (Fig. 44); submarginal double-rimmed pores very
small, little larger than size of bases of submarginal setae (Fig. 43) and sometimes difficult to distinguish
from submarginal wide-rimmed pores when margin down-curled (Fig. 44) ............... pseudugesii sp. nov.
15 Double-rimmed pores in more than a single rank (Figs 23, 54), sometimes present mesal to the abdominal
compound pores (Fig. 23)...........................................................................................................................16
- Double-rimmed pores in a single rank, although sometimes very irregular (Figs 21, 22, 26, 31, 47, 55, 56)
....................................................................................................................................................................17
16 Double-rimmed pores not in definable rows, distributed across submargin and subdorsum, several mesal
to arc of compound pores in abdomen (Fig. 23, 24); all double-rimmed pores situated within a band of
wide-rimmed pores (Fig. B ii) that is very broad and extends mesal to abdominal compound pores in dis-
tinct lobes (Fig. 23); submedian area of abdomen with only a single pair of septate pores present on each
of segments III–VI inversus Martin
- Double-rimmed pores numerous and in 2-3 ranks (Fig. 54), occupying a wide band but entirely lateral to
arc of abdominal compound pores; a narrow band of small wide-rimmed pores present between double-
rimmed pore band and puparial margin; submedian area of abdomen with septate pores more numerous...
........................................................................................................................................... etiennei sp. nov.
17 Double-rimmed pores circular, notched (as Fig. B v but more subtly indented), up to 20 μ in outer diame-
ter (when outer rim sufficiently defined), inner part of pore only slightly larger than the large, pale, wide-
rimmed pores that form a broad band laterad (Fig. 22); with only 3 pairs of submedian cephalothoracic
setae ................................................................................................................................. dugesii Cockerell
- Double-rimmed pores septate (Figs 21, 26, 31, 47, 55, 56), appearing subcircular (as fig. B iii) or ovoid,
depending on proximity of puparial margin and degree of marginal down-curling; inner part of pore much
larger than adjacent wide-rimmed pores (when these are present); with 3 or 4 pairs of submedian cephal-
othoracic setae ............................................................................................................................................18
18 Double-rimmed pores unevenly spaced, some distance from puparial margin (Figs. 26, 47). .................19
- Double-rimmed pores in a single row very close to puparial margin, more evenly spaced (Figs. 11, 21, 55,
56) ...............................................................................................................................................................20
19 Double-rimmed pores septate exceptionally large (Figs 7, 26), maximally 30–35 μ in diameter, unevenly
spaced and aligned, particularly irregular on cephalothorax (Fig. 26, to left); a narrow band of wide-
rimmed pores present in outer submargin, these pores not interdigitating with double-rimmed pores; lin-
gula usually overlapping puparial margin in slide-mounted specimens (Fig. 7); dorsal disc septate pores
sparse and also unusually large, each up to 15 μ long (Fig. 26, to right); with 3 pairs of submedian cepha-
lothoracic setae .............................................................................................................. maritimus Hempel
- Double-rimmed pores about 20 μ in outer diameter, in an unevenly-spaced row that interdigitates with
mesal boundary of a broad submarginal band of wide-rimmed pores (Fig. 47); lingula not overlapping
puparial margin in examined specimens; dorsal disc septate pores minute; with 4 pairs of submedian
cephalothoracic setae ................................................................................................ nicaraguensis sp. nov.

20 With 4 pairs of cephalothoracic submedian setae (Fig. 55); often with 2–3 pore-diameters between adja-
cent submarginal double-rimmed pores........................................................................... capiangae Bondar
- With 3 pairs of thoracic submedian setae (cephalic pair wanting); no more than one pore-diameter
between adjacent double-rimmed pores (Figs 11, 31, 56)......................................................................... 21
21 Small septate pores in submedian area arranged in linear “chains” (Fig. 30), a single chain across each
abdominal segment, but a pair of chains across meso- and metathorax; axial processes of large compound
pores [in very small type sample] truncate, not protruding beyond pore mouths; a few submarginal dou-
ble-rimmed pores displaced from even row in cephalothorax (Fig. 31).............................. vinculus Martin
- With fewer small septate pores on dorsal disc, at most in groups of 3-4 arranged linearly; axial processes
of large compound pores protruding beyond pore mouths, apically acute (Fig 56).......................................
.............................................................................................................. trinidadensis Quaintance & Baker
22 With either 4 pairs of large abdominal compound pores only (none on abdominal segment VII or VIII, fig.
70), or with additionally a pair of very small compound pores (sometimes just one on one side) on abdom-
inal segment VIII only (Fig. 69); axial processes of large abdominal compound pores often directed pos-
teromesad (Figs 69, 70) in slide-mounted specimens; puparial margin much down-curved, often
significantly deflexed.................................................................................... mirabilis species-group ......23
- Usually with 2 pairs of small abdominal compound pores, on abdominal segments VII and VIII (Figs 27,
33, 34, 37, 57, 63, 65, 67), in addition to the 4 large pairs anteriorly; if with only 1 pair of small compound
pores these are on abdominal segment VII; axial processes of large abdominal compound pores usually
not directed posteromesad; puparial margin much less, or not, down-curved / deflexed...........................24
23 Posterior half of median area of abdominal segment VI with a band of very short furrows perpendicular to
the segment VI / VII boundary (Fig. 70); between vasiform orifice and band of wide-rimmed simple pores
is a short but distinct median furrow, underlying lingula; lingular apex not reaching wide-rimmed pore
band; always only with 4 pairs of abdominal compound pores (none on segments VII and VIII, fig. 70)
..................................................................................................................................... mirabilis (Cockerell)
- Without short furrows on median area of abdominal segment VI and without a furrow underlying lingula
(Fig. 69); lingular apex slightly overlapping wide-rimmed pore band; a single pore, or a pair of tiny com-
pound pores, inconsistently present on abdominal segment VIII, underyling lingula (Fig. 69), in some
populations but entirely absent in other populations ............................................ floccissimus Martin et al.
24 Submarginal wide-rimmed pores (Fig. B ii) in a crowded broad band whose entire mesal boundary is at
least half way between puparial margin and arc of the compound pores, AND lingula with apex rounded,
with spinules covering its surface evenly (Figs 63, 65–67); operculum transversely rounded-trapezoidal,
its surface rarely corrugate....................................................................... pulvinatus species-group ……25
- If submarginal wide-rimmed pores in a crowded broad band whose entire mesal boundary extends at least
half way between puparial margin and arc of the compound pores, then lingula with its apex acute and
sometimes slightly bifid (Figs 8, 12, 16, 33) and/or dorsal surface of lingula with spinules loosely clus-
tered (Figs 20, 38, 57), rather than covering surface evenly; operculum often corrugate-rugose (Figs 16,
33, 34, 37) ...................................................................................................................................................30
25 With only 5 pairs of abdominal compound pores (none on abdominal segment VIII); submarginal setae
fine and rather short, their bases lining mesal boundary of wide-rimmed pore band (Fig. 61) .....................
................................................................................................................................. jamaicensis Cockerell
- With 6 pairs of abdominal compound pores; bases of submarginal setae usually within band of wide-
rimmed pores (Figs 41, 62, 64, 67).............................................................................................................26
26 A single row of contiguous or near-contiguous modified 8-shaped pores present immediately within
puparial margin (Figs Bi, 64, 66, 67a,), appearing different from those in the wide-rimmed pore band
mesad (Fig. B ii) usually black and appearing shallowly “w”-shaped .......................................................27
- Without a single row of such modified pores between puparial margin and wide-rimmed pore band (Figs

41, 42, 62) ...................................................................................................................................................29
27 Mesal boundary of submarginal wide-rimmed pore band almost linear in meso- and metathorax, parallel
to longitudinal moulting suture (Fig. 65), emphasised by a slight fold; in abdomen mesal boundary of pore
band interdigitates between large compound pores (Fig. 65)................................................ cocois (Curtis)
- Mesal boundary of wide-rimmed pore band not linear and not parallel to longitudinal moulting suture in
meso- and metathorax, and does not interdigitate with abdominal compound pores (Figs 63, 67, 68) .....28
28 Mesal boundary of wide-rimmed pore band in abdomen arcuate, not close enough to large compound
pores to be modified by their proximity (Figs 10, 67, 68)........................................... pulvinatus (Maskell)
- Mesal boundary of wide-rimmed pore band in abdomen closer to large compound pores, subtly curving
around them but not interdigitating with them (Figs 63, 64)............................................... juleikae Bondar
29 Wide-rimmed pores in submarginal band (Figs 41, 42) exceptionally thick-rimmed, appearing blackish on
slides, mesal boundary of band smoothly concentric with puparial margin (Fig. 41); scattered simple
pores on dorsal disc (Fig. 41), including small clusters adjacent to abdominal compound pores (Fig. 42)
also appearing to be of this thickened wide-rimmed type, not septate; cuticle slightly pigmented, brownish
............................................................................................................................................. clusiae sp. nov.
- Submarginal wide-rimmed pores much finer (Fig. 62), appearing pale on slides, mesal boundary of band
rather sinuous; dorsal disc simple pores appearing pale on slides (but their nature not clear on syntypic
material available for this study); cuticle pale. [Adult forewing with a large pigmented patch, fig. 138]
........................................................................................................................ guppyii Quaintance & Baker
30 Lingular apex acute, sometimes slightly bifid (Figs 8, 12, 16, 33); operculum with its surface corrugate-
rugose, its posterior margin convex, often almost filling vasiform orifice (Figs 16, 33) ...............................
......................................................................................................................... niveus species-group ...... 31
- Lingula with its apex rounded, even if lingula itself somewhat angular (Figs 18, 20, 27, 34, 37, 38, 57);
operculum rugose or not ...................................................................... talamancensis species-group ..... 32
31 Mesal boundary of submarginal wide-rimmed pore band forming mesally-directed lobes (Figs 12, 32), the
pore band abruptly narrowed immediately posterior to lingular apex; lingular apex almost reaching pupar-
ial margin (Fig. 12) ................................................................................................. rugioperculatus Martin
- Very broad submarginal band of wide-rimmed pores with its mesal boundary concentric with puparial
margin, closely approaching compound pores, the band only gradually narrowing posterior to vasiform
orifice (Figs 8, 33); lingular apex extending a little beyond vasiform orifice but not reaching band of
wide-rimmed pores (Fig. 33) .................................................................................................. niveus Martin
32 Submarginal wide-rimmed pores not in a sharply-defined band, closer together nearest margin but becom-
ing gradually more sparse mesad (Fig. 28), often with a few present mesal to line of submarginal setal
bases; dorsal surface of lingula evenly spinulose (Fig. 27) ............................................ pauciporus Martin
- Submarginal band of crowded wide-rimmed pores sometimes very narrow, but sharply defined (Figs 35,
36, 40, 57), the submarginal setal bases lining the band’s mesal boundary, sometimes the boundary inter-
digitating with the setal bases (Fig. 57); dorsal surface of lingula with spinules loosely clustered, rather
than covering surface evenly (Figs 20, 38, 57)...........................................................................................33
33 Mesal boundary of submarginal wide-rimmed pore band sinuous, interdigitating with bases of submar-
ginal setae (Fig. 57) ................................................................................................... darienpalmae sp. nov.
- Mesal boundary of submarginal wide-rimmed pore band more evenly concentric with puparial margin
(Figs 35, 36, 40)..........................................................................................................................................34
34 Operculum about twice as wide as long, its posterior margin roughened but conspicuously emarginate
(Figs 20, 34); Puparium usually less than 1.30 mm long .......................................... talamancensis Martin
- Operculum much less than twice as wide as long, its posterior margin very irregular, not conspicuously
emarginate (Figs 18, 37); Puparium 1.37–1.57 mm long .......................................... chirripoensis sp. nov.

Aleurodicus antillensis Dozier
(Fig. 52)
Aleurodicus antillensis Dozier, 1936: 144. Neotype puparium here designated, Puerto Rico [USNM].
Redescription, from neotype puparium

PUPARIUM. Outline oval, 1.08 mm long, 0.76 mm wide. Margin. Irregularly crenulate, not regularly
dentate. Dorsum. Longitudinal moulting suture reaching puparial margin; transverse moulting sutures becom-
ing indistinct in outer subdorsum; a distinct fuscous patch at confluence of longitudinal and transverse moult-
ing sutures in most specimens. Abdominal segment VII not significantly reduced medially. Vasiform orifice
(Fig. 52) broadly cordate, wider (0.17 mm) than long (0.12 mm); operculum trapezoidal, its posterior margin
almost completely straight, bearing a pair of stout setae; lingula head tongue-shaped, densely covered by seta-
like spinules, bearing the normal 4 stout setae; lingula extending to half-way between posterior edge of vasi-
form orifice and puparial margin. Chaetotaxy. A pair of posterior marginal setae present, long and stout, about
80 μ long, similar to lingular setae. Single pairs of cephalic, pro-, meso- and metathoracic and eighth abdomi-
nal setae present, finer than posterior marginal setae, about 30 μ long; submargin with 12 pairs of setae
(including the nominal caudal pair), posteriormost 2 pairs up to 70 μ long, but remainder only up to about 20
μ long. Pores. Cephalic compound pores 24 μ in outer diameter, large abdominal compound pores 28–32 μ in
outer diameter, situated on segments III–VI; a single pair of smaller compound pores located on abdominal
segment VII (Fig. 52), 20 μ in outer diameter, but none present on segment VIII. Axial processes of com-
pound pores not extending beyond pore rims, hardly evident. Submargin defined by crowded band of wide-
rimmed pores, inner boundary of band lined by the submarginal setal bases and not interdigitating with com-
pound pores; on abdominal segments VII–VIII only (Fig. 52) inner boundary submarginal pore band is also
lined by a few distinct double-rimmed septate pores (7 in neotype); remainder of dorsal disc punctuated by
small septate pores, fairly evenly distributed; a small array of bright pores anterior to each of the small com-
pound pores on abdominal segment VII. Venter. Ventral characters entirely typical for Aleurodicus.
DISTRIBUTION. Neotropical Region – Dominican Republic, Puerto Rico.
MATERIAL EXAMINED. Neotype puparium, Puerto Rico, San Juan, Parque Muñoz Rivera, near Naval
“Resv”, on Calophyllum antillanum (Clusiaceae), 20.iv.1952 (Russell #357) (USNM); 5 puparia, 1 third-
instar / puparial mid-moult specimen, same data as neotype—on two slides in total (USNM). Several other
samples from USNM (some duplicates in BMNH), collected in Puerto Rico, from Pithecellobium unguis-cati,
Cissus sycioides, Ficus laevigata, Inga laurina, and Cocos nucifera.
COMMENTS. No type depository for A. antillensis was traced by Mound & Halsey (1978), and Debra
Creel (personal communication) has confirmed that the USNM collection does not house any material with
data quoted by Dozier (1936). Given an inadequate original description, it is desirable to stabilise this species
through the designation of a type specimen. Two slides from the Dozier collection are present in USNM, and
one of these contains four empty pupal cases that are in good condition for examination, along with three
uncleared puparia: unfortunately, the data are imprecise, being merely “Puerto Rico, H.L. Dozier, ?1935”.
However, another slide from Puerto Rico contains puparia, collected more recently, that are in extremely good
condition and are clearly conspecific with those on the Dozier slide, collected from Calophyllum antillanum
which is one of the hosts listed by Dozier: one of these puparia has therefore been chosen as a neotype, here
designated, and is clearly indicated on its slide. A. antillensis is a member of the capiangae / dugesii species-
group.

Aleurodicus araujoi Sampson & Drews
(Figs 1, 21)
Aleurodicus araujoi Sampson & Drews, 1941: 145. Syntypes, Mexico [UCD, examined].
DISTRIBUTION. Neotropical Region – Belize, Costa Rica, Mexico.

MATERIAL EXAMINED. Syntype puparia, Mexico, Manzanillo, on Aristolochia sp., xii.1925 (UCD);
numerous samples, Belize, host records tabulated by Martin (2004) (BMNH, USNM); quarantine-interception
material, Costa Rica (USNM).
COMMENTS. Numerous samples of Belizean material of A. araujoi (BMNH) have been identified in
comparison with syntypic material from Mexico, loaned by UCD in connection with a study of Aleurodicinae
in Belize (Martin, 2004), in which publication host plant records in Belize were tabulated. This species clearly
belongs to the capiangae / dugesii group, and may be recognised by use of the key to species, p. 15.
A sample of puparia from mistletoe in Costa Rica (BMNH) has its puparia rather darkly pigmented, and
other slight differences from the extensive Belizean material, but is considered likely to be conspecific.
Aleurodicus capiangae Bondar

(Fig. 55)
Aleurodicus capiangae Bondar, 1923: 71. Lectotype puparium here designated, Brazil [USNM].
Aleurodicus fucatus Bondar, 1923: 74. Syntypes, Brazil [BMNH, examined]. Syn. nov.
DISTRIBUTION. Neotropical Region – Anguilla, Brazil, Costa Rica, Ecuador, Guyana, Nicaragua, Panamá,
Surinam, Trinidad.
MATERIAL EXAMINED. Type material of A. capiangae and A. fucatus as detailed and discussed below;
figured specimen, Panamá, Canal Zone, Cerro Galera, on ?Combretum, 26.iii.1983 (Martin #4147) (BMNH);
numerous samples from most countries listed above (BMNH, USNM).
COMMENTS. The USNM collection contains a single slide of A. capiangae collected by Bondar, the
data being “Aleurodicus capiangae Bondar, Bahia, Brazil, G. Bondar, #504, Q[uaintance] 23299”. Bondar
(1923: 72) clearly stated that cotype material [=syntype in currently accepted terminology] was deposited in
USNM. Accordingly, one of three puparia on the single Bondar slide of A. capiangae in USNM is here desig-
nated the lectotype (see discussions of lectotype designation, p. 9), and is clearly indicated on the slide. All
three specimens on the lectotype slide are parasitised (each with a fuscous patch in a different position and one
with an emergence hole), and the lectotype puparium also has some adult whitefly abdominal cuticle present
inside it, of which the observer should be aware.
Five specimens in BMNH are on slides prepared from duplicate dry material of A. fucatus in USNM, pre-
sumed to be syntypic – the data are “Inga & cacao, Brazil, Dec. 1922, fr. Dr G. Bondar #637, Q[uaintance]
23041”. Comparison of these specimens with the lectotype and paralectotypes of A. capiangae indicate that
the two species are one and the same. Bondar (1923) described the two species consecutively, on pp 71–73
(capiangae) and 74–75 (fucatus). He illustrated A. fucatus with considerably larger compound pores than he
showed for A. capiangae, but there is little size difference observed in the examined specimens. Bondar’s
comments compared both species with A. trinidadensis Quaintance & Baker, but not with each other. Even
though, amongst the many studied samples, there is considerable variation in the numbers and density of dou-
ble-rimmed septate pores in the outer-submarginal row, this feature is in fact almost identical in the type spec-
imens of both A. capiangae and A. fucatus. Accordingly A. fucatus is here regarded as a junior synonym of A.
capiangae, syn. nov., using the principle of page priority to decide upon the senior name.

As stated above, the many specimens examined in connection with this study display great variation in the
number of double-rimmed septate submarginal pores (and hence the proximity of each pore with its neigh-
bours) and also variation in compound pore size: specimens with their submarginal septate pores almost con-
tiguous impinge on the characters used to define A. trinidadensis (q. v.) and there is uncertainty over the
species limits in A. capiangae, which may be part of a complex of similar species. With such variability, and
in view of the imperfect condition of the lectotype, the specimen chosen for the illustration, figure 55, is an
individual from Panamá that closely matches the lectotype – the data are detailed in Material Examined,
above.
Mound (1961) provided observational data on the biology of A. capiangae, based on a colony on Canna
indica in Trinidad. No voucher material of this studied colony is known to exist.
Aleurodicus charlesi sp. nov.

(Fig. 46)
PUPARIUM. Habitus. Appearance in life not noted. Margin. Outline oval, 0.98–1.10 mm long, 0.72–0.86 mm
wide, generally widest opposite transverse moulting suture (n=4). Margin apparently unevenly crenulate, due
to submarginal pores being in relief through down-curling, but true margin can be seen to be smooth where
fully deflexed. Dorsum. Longitudinal moulting suture reaching puparial margin; transverse moulting sutures
reaching outer subdorsum. Abdominal segment VII similar in length to segment VIII, medially; pro- / meso-,
meso- / metathoracic and abdominal segmentation well marked. Vasiform orifice (Fig. 46) cordate, rather
close to posterior margin of puparium, a little wider than long, smooth; operculum trapezoidal, occupying
over three-quarters of vasiform orifice, surface very finely punctate, its posterior margin shallowly concave,
bearing a pair of unusually long stout setae; lingula head tapering but rounded apically, densely covered by
seta-like spinules, bearing the normal 4 stout setae, its apex close to puparial margin. Chaetotaxy. A pair of
posterior marginal setae present, similar to eighth abdominal setae but shorter than the 12 pairs of long and
fine submarginal setae (including the nominal caudal pair); single pairs of submedian meso- and metathoracic
setae present, similar to eighth abdominals; submedian cephalic and prothoracic setae absent. Pores. With
cephalic pair and only 5 abdominal pairs of compound pores (none on abdominal segment VIII, fig. 46); Larg-
est compound pores on abdominal segments III and IV, up to 60 μ in outer diameter; cephalic pair similar to
pair on abdominal segment VI, about 30 μ; pair on abdominal segment V intermediate in size; small pair of
compound pores on abdominal segment VII subcircular, about 20 μ in diameter; axial processes of large com-
pound pores variable in appearance, moderately acute to rather blunt apically. Immediately within puparial
margin lies a band of small wide-rimmed simple pores which (in thoracic and abdominal areas) are relatively
sparse in submargin but becoming denser mesally, “fingers” of this band of pores extending mesad to sur-
round compound pores (Fig 46) except for the pair on abdominal segment VII; within this pore band, mesal to
submarginal setae, is a loose row of small double-rimmed notched pores with the notch bulging slightly (Fig.
46 expanded detail); remainder of abdominal dorsal disc (except abdominal segment I and most of thoracic
area) with many notched pores similar in size to submarginal double-rimmed variety but without the double
rims. Venter. Ventral characters typical for Aleurodicus, but 2 pairs of unusually large “sacs” of delicate ven-
tral cuticle present anterolateral to vasiform orifice; ventral abdominal setae very long and fine, around 90μ
long.
MATERIAL EXAMINED. Holotype puparium, GUYANA, Mazaruní-Potaro, plateau surrounding Kai-
eteur Falls, on unidentified host, 11.vi.2006 (E. Charles coll., Martin #8318) (BMNH). Paratypes: 3 puparia,
same data as holotype (BMNH, UG).
ETYMOLOGY. This species is named for its collector, Elroy Charles (UG), in recognition of his exten-
sive field work and investigations into the morphology and molecular systematics of the Aleurodicinae.

COMMENTS. This species appears to have close affinities with A. magnificus Costa Lima and A. neglec-
tus Quaintance & Baker, to judge from the shape and position of the vasiform orifice and lingula, along with
many rather large notched simple pores on the dorsal disc, and the presence of double-rimmed pores within
the submarginal wide-rimmed pore band. However, the incursion of smaller wide-rimmed pores into the com-
pound pore zone, and the presence of only meso- and metathoracic pairs of submedian cephalothoracic setae
more strongly suggest inclusion in the dispersus species-group, where the presence of small compound pores
on abdominal segment VII is shared by A. flavus Hempel.
Aleurodicus chirripoensis sp. nov.

(Figs 17, 18, 37–40, 133)
PUPARIUM. Habitus. Developing in distinct groups under leaves, with rather sparse waxy secretions (Fig.
133) limited to a peripheral annulus and a dusting of dorsal meal, with brownish subdorsal bars visible where
puparial surface is visible through the meal; broken filaments from the large compound pores visible on the
leaf surface. Morphology. Most characters are shared with A. talamancensis (q.v.) and only differences
are described below. Outline. 1.37–1.57 mm long, 0.97–1.17 mm wide, generally widest at abdominal seg-
ment I/II (n=26). Dorsum. Raised areas on abdominal segments VII and VIII (Fig. 37) broader and less well
defined than in A. talamancensis, with caudal furrow therefore less evident; submedian abdominal depres-
sions more distinct, with 2–3 contiguous pits on each side of submedian area (Fig. 39); abdominal segment
VI/VII division marked medially by a distinct suture-like fold; cephalic pair and anterior 4 pairs of abdominal
compound pores up to 65 μ maximum diameter; rim of vasiform orifice (Fig. 37) a little less sharply defined;
operculum almost completely occupying vasiform orifice, surface punctate and with longitudinal and trans-
verse rugae, its posterior edge not conspicuously emarginate, often slightly convex; clusters of surface
spinules on lingula head more clearly defined (Fig. 38); largest of the dorsal disc bright simple pores appar-
ently each with a fine transverse septum; usually more simple pores present in vicinity of each compound
pore, the most numerous clusters typically on abdominal segment VII and cephalic segment, with up to 10
pores (Figs 37).
MATERIAL EXAMINED. Holotype puparium, COSTA RICA, San José Province, Cerro Chirripó, 2600–
2800 metres, on ?Maxillaria sp. (Orchidaceae), 17.ii.1983 (J H Martin #3873) (BMNH); Paratypes: 49
puparia (of which 13 on leaf tissue in alcohol), 10 third-instar nymphs, 1 second-instar nymph, same data as
holotype (BMNH, INBio, USNM).
ETYMOLOGY. This species is named for its collecting locality, the mid-montane slopes of Cerro
Chirripó which, at 3820-metres, is the highest mountain peak in Central America.
COMMENTS. Many of the observed differences between puparia of A. chirripoensis and A. talamancen-
sis were earlier (Martin, 2005) considered likely to be a result of size variation in A. talamancensis alone, and
the material listed above was provisionally determined as A. talamancensis, but was not given paratype status
because doubt did remain. However, the size range of the orchid-feeding specimens from Cerro Chirripó does
not overlap the size range of all the examined specimens of A. talamancensis, including coconut-feeding
material from Ecuador not available at the time of describing A. talamancensis. Along with larger overall
puparial size, specimens of A. chirripoensis have their large compound pores almost twice the diameter (up to
65μ) of those in puparia of A. talamancensis (about 30–40 μ), and there are greater numbers of dorsal disc
simple pores in the Cerro Chirripó sample; also, the posterior edge of the operculum is markedly emarginate
in A. talamancensis, but not so in A. chirripoensis which also has a less clearly defined caudal furrow. These
differences are not, now, considered to be merely artifacts of differing physical size and the montane orchid-
feeding specimens from Cerro Chirripó are regarded as a distinct species.

The author recently described A. niveus and A. rugioperculatus from Belize (Martin, 2004), and these
share with chirripoensis the characteristically rugose operculum that occupies most of the vasiform orifice,
the robust and often-pigmented puparial cuticle, and the possession of submarginal simple pores of only one
type. However, both A. niveus and A. rugioperculatus have a much wider submarginal pore band than do A.
talamancensis and A. chirripoensis, and neither A. niveus nor A. rugioperculatus possesses lingular spinules
arranged in clusters. A. niveus (like A. chirripoensis an orchid-feeding species) is characterised by its lingular
apex being developed into an acute, usually bifid, process, and also by the inner boundary of its very wide
submarginal zone of simple pores being concentric with its puparial margin. A. rugioperculatus, from coconut
palms and unidentified broad-leaved hosts, has its entire lingula head acute-triangular, and the inner boundary
of its submarginal band of simple pores lobe-like.
Aleurodicus clusiae sp. nov.

(Figs 41, 42, 135)
PUPARIUM. Habitus. Small, loose, groups of puparia occur under leaves (Fig. 135), each group in the middle
of a larger patch of mealy wax; dorsal surface covered by opaque white wax that appears almost carapace-
like; each puparium surrounded by thickly-flocculent wax such that it appears to nestle in a woolly “nest”.
Margin. Outline almost perfectly oval, 1.09–1.29 mm long, 0.77–0.93 mm wide, generally widest at abdomi-
nal segment I/II (n=8). Margin smooth. Dorsum. Cuticle rather sclerotic, brownish-pigmented in more mature
individuals. Longitudinal moulting suture reaching puparial margin; transverse moulting sutures reaching
almost to margin. Anterior to each cephalic compound pore is a tubercular structure, usually with a shallow
trough-like depression at its base (Fig. 41). Subcircular cephalothoracic depressions present (Fig. 41), but
abdominal equivalents are merely widened subdorsal extensions of segmental divisions (Fig. 42); abdominal
segmentation furrow-like submedially, less well defined subdorsally; segments II–VI each with a short trans-
verse furrow at about mid-length of segment; meso- and metathoracic segments indicated by thickened shal-
low folds (Fig. 41); abdominal segment VII not evident medially, pockets touching segment VI boundary,
segment (VII+)VIII similar in length to segment VI medially. Vasiform orifice rounded-cordate, wider than
long; operculum rounded-trapezoidal, smooth dorsally but finely spinulose along posterior edge, posterior
margin faintly concave and bearing a pair of setae, operculum occupies about three-quarters of the orifice; lin-
gula head rather straight-sided but rounded apically, densely covered by seta-like spinules, bearing the normal
4 stout setae; lingula extending only a little beyond edge of vasiform orifice when in its natural position, but
many specimens have the lingula further extended. Chaetotaxy. A pair of posterior marginal setae present, a
little shorter and more robust than submarginal pairs; eighth abdominal setae present, anterior to vasiform ori-
fice (Fig. 42); submargin with 12 pairs of very fine and long setae (including the nominal caudal pair), most
extending beyond puparial margin (Fig. 41), longest pairs (posterior-abdominally) up to 120μ long; single
pairs of pro-, meso- and metathoracic submedian setae present, similar to posterior marginal setae, but cepha-
lic submedian pair absent. Pores. Cephalic and anterior 4 pairs of abdominal compound pores all similar in
size, 25–35 μ in outer diameter, usually seen in lateral view on slides, distinctly cylindrical in shape and each
with a short, dagger-like axial process extending from the pore mouth (Figs 41, 42); posterior 2 pairs of
abdominal compound pores significantly smaller than remainder (Fig. 42), with axial processes internal only,
pair of pores on segment VII cylindrical in shape as seen on slides, pair on segment VIII shorter, appearing
more ovoid. 2 pairs of large robust cicatrices present thoracically (scars of large compound pores in third-
instar – see below). Immediately within puparial margin is a narrow pore-free zone mesad of which is a broad
band of wide-rimmed simple pores, the mesal boundary of this zone not interdigitating with compound pores;
dorsal disc with simple pores of apparently similar structure, very variable in size, some of them actually
larger than those of the submarginal band and appearing rather tubercle-like, these largest simple pores often

loosely clustered (Figs 41, 42). Venter. Ventral abdominal setae placed posterior to vasiform orifice, similar to
longest dorsal setae. Legs typically robust and two-segmented, smooth, each with a pronounced apical claw.
Antennal apices extending to articulation of hind legs, their bases anterolateral to fore legs, pointed apically.
Tracheal folds absent.
THIRD-INSTAR NYMPH. Outline broadly oval, 0.73–0.81 mm long, 0.53–0.65 mm wide, widest a mid-
length (n=12). Margin smooth. Vasiform orifice and operculum as in puparium, but lingula surprisingly vari-
able, measuring 70μ (head rather truncate) to 100μ (head elongate tongue-shaped). 2 pairs of large compound
pores present in cephalothorax, rounded bell-shaped, up to 40μ in width, each with an axial process that just
protrudes beyond pore mouth. 3 pairs of small cicatrices present, two pairs in cephalothoracic area and third
pair posterolateral to vasiform orifice, these being the scars of compound pores in second-instar. Chaetotaxy
as in puparium, setae as long as in puparium. Dorsum with pores generally distributed as in puparium, but
submarginal band interrupted by submarginal setae. Legs triangular, evidently 3-segmented, each with a
curved apical claw; antennae small, about same size as apical segments of legs; ventral abdominal setae simi-
lar to dorsal setae.
ADULTS. Study material is available as listed below. Basic parameters typical for Aleurodicus.
MATERIAL EXAMINED. Holotype puparium, GUYANA, Mazaruní-Potaro, plateau surrounding Kai-
eteur Falls, on Clusia sp. (Clusiaceae), 14.vi.2006 (J.H.Martin #8348) (BMNH). Paratypes: 12 puparia, 12
third-instar nymphs, 7 adult males, same locality and host(s) as holotype, 08.x.1991 (Martin #5961),
11.vi.2006 (Martin #8314) and 14.vi.2006 (Martin #8348) (BMNH, UG, USNM); 6 puparia dry on leaf sam-
ple, same data as holotype (BMNH). Non-type specimen, single puparium, unidentified host, Tukeit forest
trail near Kaieteur Falls, 13.vi.2006 (Martin #8341) (BMNH).
ETYMOLOGY. The specific name reflects the host plant genus / family, Clusia, Clusiaceae.
COMMENTS. Within Aleurodicus, A. clusiae is unusual in apparently having only one basic type of sim-
ple pore—the wide-rimmed variety as defined by Russell (1965). With its relatively small lingula, and a broad
submarginal pore band only comprising wide-rimmed pores, this species is here treated as a member of the
pulvinatus group of species: however, the rather sclerotic cuticle and apparently non-septate form of the dorsal
disc pores tend to indicate possible other affinities. The variation in shape and, hence, length of the third-instar
lingula is not understood.
A. clusiae is represented by type material from several separate samples, all collected from species of Clu-
sia in the vicinity of the Kaieteur Falls, Guyana. At least two species of Clusia are thought to be involved,
including C. grandiflora, both of them with extremely smooth, leathery, leaves. In all samples the nymphal
colonies comprised only very small numbers of individuals. A single puparium from an unidentified host, in
forest in the vicinity of Kaieteur Falls, is extremely similar to those in the type series but displays detail differ-
ences that lead to it being excluded from the paratype series.
Aleurodicus coccolobae Quaintance & Baker

(Figs 2, 13, 123)
Aleurodicus coccolobae Quaintance & Baker, 1913: 46. Lectotype puparium here designated, Mexico [USNM].
DISTRIBUTION. Neotropical Region - Belize, Brazil, Costa Rica, Ecuador, Honduras, Mexico, Panamá.
MATERIAL EXAMINED. Type material as detailed and discussed below (USNM); Belize – 4 samples,
Chiquibul Forest Reserve and Mountain Pine Ridge, on Cassia, ?Simarouba, Persea and Psidium (BMNH);
Costa Rica – 3 samples, Cahuita and Turrialba, on ?Bixa and palms (BMNH); Ecuador, 3 samples, Manta,
Coca and San Vicente, on Annona, Cocos and Inga (BMNH); Mexico – 3 samples, Veracruz and Progreso, on
Hibiscus, Manihot and “croton” (BMNH).

COMMENTS. Only a single slide of syntype material was available for study, housing three parasitised
puparia. This is clearly the slide mentioned by Quaintance & Baker in their original description – “Type
no.14765 ….. three parasitized specimens in balsam mount”. This slide had a third label affixed on top of one
of the original two, with the data “Q.2975, Yucatan, 1904”, and this label has now been transferred to the
obverse surface of the slide, to reveal the earlier label. The original two labels bear the data “Aleurodicus coc-
colobae Quaintance & Baker, Type no. 14765 USNM” and “2975 Q., Aleurodicus n. sp.?, 1/28/04, Progreso,
Yucatan”. A fourth label has now been added, on the obverse, with details of the lectotype designation. The
specimens on the lectotype slide had clearly not been treated for removal of waxy secretions, and all are still
occupied by parasitoid pupal material, but most of the main characteristics are visible on one specimen which
is here designated the lectotype and is indicated as such on a new slide label. A slight advantage of these
specimens not having been treated for wax removal is that the emergence of secretion from the main types of
simple pore is clearly visible, a character not seen in fully-prepared specimens on slides.
A. coccolobae is one of the species that form the dispersus-group (Russell, 1965), a natural grouping
within Aleurodicus, whose puparia either lack small compound pores entirely, or have only one pair present
(on abdominal segment VII) along with particularly dense provision of dorsal disc pores of four types, and
submedian thoracic setae much reduced in size. This small species-group was discussed by Russell as part of
her description of A. dispersus, the spiralling whitefly. The appearance of A. coccolobae and A. dispersus
puparia in life may be compared in figures 123 and 122 respectively.
Aleurodicus cocois Curtis

(Figs 65, 66)
Aleyrodes cocois Curtis, 1846: 284. Lectotype puparium, Barbados [designated by Martin, 1997: 125, MOV, examined].
Aleurodicus cocois (Curtis) Morgan, 1892: 32.
Aleurodicus anonae Morgan, 1892: 32. Lectotype, Guyana [designated by Mound & Halsey, 1978: 229, BMNH, exam-
ined]. Synonymised by Mound & Halsey, 1978: 229.
DISTRIBUTION. Neotropical Region – Brazil, Guyana, Venezuela and widely across the Caribbean islands
including Puerto Rico.
MATERIAL EXAMINED. Type material of Aleyrodes cocois and Aleurodicus anonae as detailed and
discussed by Martin (1997); numerous samples from the Caribbean and the countries listed above (BMNH,
USNM).
COMMENTS. A question had arisen about the status of material of Aleyrodes cocois Curtis [the type for
the entire subfamily Aleurodicinae, through its seniority over A. anonae Morgan] in BMNH, marked as hav-
ing “type” status. Investigations revealed that the true type sample of A. cocois resided in the MOV collection
in Melbourne, not in BMNH as indicated by Mound & Halsey (1978: 229). However, the synonymy of A.
anonae with A. cocois was found to be correct (Martin, 1997).
A. cocois is very similar to A. pulvinatus and A. juleikae (q.v.) and is here treated as a member of the pulv-
inatus-group of species in the key to species. However, the species limits of the cocois / pulvinatus / juleikae
assemblage are not clear and may change with future work. At present, the key (couplets 27-28, p. 19) may be
used to distinguish the species as here understood.
Aleurodicus darienpalmae sp. nov.

(Fig. 57)
PUPARIUM. Habitus. Not noted. Morphology. Most characters are shared with A. talamancensis (q.v.)
and only differences are described below. Outline. 0.92–0.97 mm long, 0.67–0.71 mm wide, generally

widest at abdominal segment I/II (n=26). Dorsum. Mesal boundary of submarginal wide-rimmed pore band
conspicuously lobulate, interdigitating with bases of submarginal setae (Fig. 57); division between abdominal
segment VI and VII/VIII marked medially by a distinct suture-like fold linking anterior edges of pockets, but
only 7 segments discernible on median line anterior to vasiform orifice; cephalic pair and anterior 4 pairs of
abdominal compound pores 30–40 μ maximum diameter, their axial processes short; operculum transversely
rounded-trapezoidal, its surface with fine punctae and subtly rugose, its posterior edge not conspicuously
emarginate, almost straight and without a pair of setae on its posterior margin; clusters of surface spinules on
lingula head more subtly defined (Fig. 57 expanded detail); most dorsal disc simple pores similar in size to
those in submarginal wide-rimmed pore band, a few a little larger.
MATERIAL EXAMINED. Holotype puparium, PANAMÁ, Darién Province, Enseñada del Guayabo, 16–
19 km S.E. of Jaqué, 7º 20´ N, 78º 05´ W, on unidentified Palmae, 13.i.1983 (J.H.Martin #3615) (BMNH).
Paratypes: 1 puparium, 1 third-instar nymph, same data as holotype (BMNH).
ETYMOLOGY. The species name is formed from two nouns in aposition, reflecting its type locality
(Darién – Province of Panamá) and its incomplete host plant data (Palmae—a family name now more usually
replaced by Arecaceae).
COMMENTS. This species differs from the very similar A. talamancensis principally in the nature of the
submarginal pore band, whose mesal boundary interdigitates with the bases of the submarginal setae: this
characteristic is also seen in the single third-instar nymph and it is not considered to be a variation of A. tala-
mancensis. Also, the puparial operculum of A. darienpalmae is not posteriorly emarginate as it is in A. tala-
mancensis.
Aleurodicus decemensis sp. nov.

(Fig. 72)
PUPARIUM. Habitus. Not noted. Margin. Outline oval, 1.05 mm long, 0.73 mm wide, widest at abdominal
segment II (n=1). Cuticle pale but very slightly dusky in cephalothoracic subdorsum, true margin smooth,
almost entirely deflexed; edge of puparium (apparent margin) rather uneven, resulting from submarginal pores
observed in profile; short folds leading mesad from true margin can give a slight appearance of laterally-con-
tiguous teeth. Dorsum. Longitudinal moulting suture only faintly marked, apparently reaching puparial mar-
gin; transverse moulting sutures slightly easier to discern, reaching submargin. Abdominal segment VII much
reduced medially, about same length as pockets; abdominal segmentation, meso- / metathoracic and pro-
mesothoracic segmental divisions clear but cephalic / prothoracic division less marked. Vasiform orifice
broadly-cordate, slightly wider than long, smooth; operculum trapezoidal, almost twice as wide as long, sur-
face with many minute dots, its posterior margin slightly convex, bearing a pair of stout setae; lingula head
proportionately very large, tongue-shaped, densely covered by seta-like spinules, bearing the normal 4 stout
setae, its apex almost reaching puparial edge; Chaetotaxy. A pair of posterior marginal setae present, about
90μ long, fine; eighth abdominal setae present, anterior to vasiform orifice, 30μ long, similar to single pairs of
submedian pro-, meso- and metathoracic setae; a pair of minute cephalic submedian setae also present, only
8μ long; submargin with 12 pairs of setae, most of them similar to posterior marginals but posteriormost 2
pairs very much longer and more robust, 120-160μ long. Pores. Cephalic compound pores 32μ in outer diam-
eter, anterior 4 pairs of abdominal compound pores almost identical to cephalics but 36μ in outer diameter;
axial processes of the 5 pairs of large compound pores extremely elongate, apically acute and sword-like, up
to 170μ long; length of axial processes leads to large compound pores being observed only laterally, and no
basal cells can be discerned; abdominal segments VII and VIII each with a pair of tiny compound pores about
16μ in diameter, each with a tiny, truncate, axial process that protrudes slightly beyond pore mouth. Immedi-
ately within puparial margin lies a single row of curious D-shaped pores (Fig. 72, detail of deflexed submar-

gin); further mesad, but still in submargin, is a row of much larger 8-shaped septate pores that seem not to be
the double-rimmed type that are common in the submargin of several other species of Aleurodicus; remainder
of dorsal disc with small 8-shaped pores and even smaller wide-rimmed pores; submedian area, particularly of
abdomen, is effectively defined by asymmetric geminate pores – curious modified 8-shaped pores in which
one half is much smaller than the other (Figs B xi, 72 expanded detail), with many having an apparent faint
double rim - this type of pore is also present elsewhere, particularly adjacent to bases of large compound
pores. Venter. Ventral abdominal setae 90μ long, fine, underlying lingular base. Legs and antennae as figured,
robust, typical for Aleurodicus. Tracheal folds absent.
THIRD-INSTAR NYMPH. Cuticle slightly pigmented, outline broadly oval, 0.58–0.59 mm long, 0.44–
0.45 mm wide, widest opposite posterior pair of compound pores (n=2). Margin down-curled but not
deflexed, rather uneven. Vasiform orifice and operculum as in puparium, but lingula just reaching beyond
body margin. 2 pairs of large compound pores present in cephalothorax, bell-shaped, 25-30μ wide, each with
a long and sword-like axial process that extends much beyond pore mouth and is up to 70μ long, measured
from pore-base. 3 pairs of small cicatrices present, two pairs in cephalothoracic area and third pair posterolat-
eral to vasiform orifice, these indicating presence of 3 pairs compound pores in second-instar. Chaetotaxy as
in puparium, posteriormost 2 submarginal pairs very long and robust, up to 150μ long; 4 pairs of cephalotho-
racic setae present but extremely small, especially cephalic pair, less than 5μ. Dorsum with 8-shaped pores of
differing sizes, outer (submarginal) row the largest, appearing more like double-rimmed pores than in pupar-
ium; interspersed with 8-shaped pores on dorsal disc are tiny wide-rimmed pores. Legs triangular, evidently 3-
segmented, each with a curved apical claw; antennae small, about same size as apical segments of legs; ven-
tral abdominal setae similar to posterior marginal setal pair.
MATERIAL EXAMINED. Holotype puparium, PANAMÁ, Canal Zone, Gamboa Hill, on Machaerium
sp., cf. milleflorum or purpurascens (Fabaceae: Papilionoideae), 20.iii.1983 (J.H.Martin #4091) (BMNH).
Paratypes: 2 third-instar nymphs, same data as holotype (BMNH).
ETYMOLOGY. The specific epithet is created from the latin words decem (ten) and ensis (sword),
describing the ten sword-like axial processes of the large compound pores in the puparium of this species.
COMMENTS. Although this species is known from just a single puparium and two third-instar nymphs,
its most unusual characteristics lead to its description being well justified. The extreme length and robust
nature of the compound pore axial processes is highly unusual (more reminiscent of Palaealeurodicus spe-
cies), and the form of the 8-shaped pores also differs consderably from the double-rimmed septate type more
often seen in Aleurodicus species. The asymmetric 8-shaped pores that line the submedian area of the pupar-
ium, with one “loop” only half the size of the other, may be unique. A. decemensis does not have obvious
close affinities with any other Aleurodicus species.
Aleurodicus dispersus Russell

(Figs 3, 71, 122)
Aleurodicus dispersus Russell, 1965: 49–54. Holotype puparium, U.S.A. (Florida) [USNM, paratypes examined].
DISTRIBUTION. Neotropical Region - widely distributed; Nearctic Region - U.S.A. (Florida); Palaearctic
Region - Canary Islands, Madeira; Ethiopian Region - Cameroun, Congo, Benin, Ghana, Guinea-Bissau,
Kenya, Nigeria, Sao Tomé, Tanzania, Togo; Malagasian Region - Mauritius; Oriental Region - Hainan
(China), India, Maldives, Sri Lanka, Thailand, Vietnam; Austro-oriental and Pacific Regions - widely distrib-
uted; Australia - Queensland.
MATERIAL EXAMINED. Paratype puparia, U.S.A., Florida, Key West, on Cocos nucifera, 12.vi.1964
(Weems) (BMNH); numerous samples from most countries listed above (BMNH).

COMMENTS. A. dispersus is one of only three or four known Aleurodicus species whose puparia possess
only 4 abdominal pairs of compound pores on the puparial abdomen, being entirely without pairs of smaller
compound pores on abdominal segment VII or VIII: the others are A. mirabilis and some populations of A.
floccissimus (q.v.) [which belong to a different species-group, see key], and A. coccolobae which may be the
closest relative of dispersus (living puparia of dispersus and coccolobae may be compared in figures 122 and
123). A third described member of the dispersus-group (A. flavus Hempel, q.v.) has additionally just one pair
of tiny compound pores, situated on abdominal segment VII, but otherwise strongly resembles A. dispersus.
Two species here described also appear to belong to the small dispersus assemblage of species – see A. char-
lesi (p. 22) and A. spectabilis (p. 42).
A. dispersus was unusual in remaining undescribed for many years, despite the accumulation of numerous
samples in the USNM collection, most of them interceptions at United States port quarantine facilities (see
also discussion of A. niveus, p. 38); yet A. dispersus is now the best-known of all members of the Aleurodici-
nae. It is probable that A. dispersus (the so-called “spiralling whitefly”) is native to the Caribbean and north-
ern South America. Although many of the paratypes are from these native areas, Russell chose a sample from
Florida from which to select the holotype. This curious technicality qualifies A. dispersus as one of several
neotropical whiteflies to have been described from material sampled in an area of probable introduction. This
choice of holotype may have been the result of most other specimens, then available to Russell, being isolated
quarantine interceptions with imprecise originating locality data. Since its description, the spiralling whitefly
has become almost pan-tropical in its distribution, its rapid radiation having begun in the 1980s (Martin &
Lucas, 1984; Martin, 1990). Winter temperatures appear to have, for the present, prevented its invasion
extending into the Middle East and Mediterranean regions, and limited its expansion in Australia.
Aleurodicus dugesii Cockerell

(Figs 4, 22, 131)
Aleurodicus dugesii Cockerell, 1896: 302. Lectotype here designated, Mexico [USNM].
Aleurodicus poriferus Sampson & Drews, 1941: 149. Syntypes, Mexico [UCD, examined]. Synonymised by Martin,
2004: 22.
DISTRIBUTION. Neotropical Region – Belize, Guatemala, Mexico, Nicaragua, Venezuela; Nearctic Region
– U.S.A. (Arizona, California, Florida, Louisiana, Texas); Pacific Region – Hawaii; Austro-oriental Region -
Java.
MATERIAL EXAMINED. Lectotype and paralectotypes of A. dugesii, and other probable former syntype
specimens as detailed and discussed below (USNM); numerous probable former syntype puparia of Aleurodi-
cus dugesii, as detailed and discussed below (BMNH); 2 syntype puparia of Aleurodicus poriferus, Mexico,
Sinaloa State, on Bumelia laetivirens, viii.1925 (UCD); Hawaii, Honolulu airport, on Hibiscus sp., x.2002
(BMNH); several samples Belize, Chiquibul Forest Reserve, host records tabulated in Martin (2004) (BMNH,
USMN); Nicaragua, near Nandaime, on Malvaviscus arboreus, 13.vi.2004 (BMNH); California, Catalina
Island, on Hibiscus sp., 09.xii.2003 (BMNH).
COMMENTS. Six slides in USNM purported to be “types” of A. dugesii. One slide of puparia and one
slide of adults bear the data “on Hibiscus rosa-sinensis, Guanajuato, Mexico, T.D.A.Cockerell colr., Sept. 25,
1896, Q. 3273, Bur. Ent. 7233” on one label, the data matching that quoted in the description except that Dr
Alfred Dugès was the actual collector, who sent copious material to Cockerell. Three more of the USNM
slides bear the data “Q. 3273, Mexico, on guava, June 21, 1892, part of type” on one label. However, four of
these five slides have a second, red, label with the data “Aleurodicus dugesii Ckll Type no. 14764 U.S.N.M.”.
The adult preparation from Hibiscus has no second label. Whilst the date of the three guava slides would be
consistent with this material also being part of that sent by Dugès, and the Quaintance number 3273 is the

same, guava is not a host discussed by Cockerell: possibly “guava” may have been a misinterpretation of the
locality “Guanajuato” by whoever made these slides, and they are thought likely to be part of the same Dugès
material. The puparia on the Hibiscus slide from Guanajuato are in good condition, as well as being those with
the data as quoted by Cockerell – and one puparium has been here designated as the lectotype; the three
adults on the other slide bearing this data become paralectotypes.
The sixth USNM slide bears the data “TYPE” on one label and “Q.3274, Aleurodicus dugesii Ckll., Sid-
ney Lang, 1/29/99 [29.i.1899]: with its collection date three years subsequent to Cockerell’s description, this
slide clearly is not part of the type material.
Three slides in BMNH were made by the author from dry material with the following data: “Aleurodicus
dugesii Ckll, on ?, Guanajuato, Mexico (Dugès), Pres. T.D.A. Cockerell, Det. Quaintance, B.M. 1923-178”. A
note in the BMNH Register for this accession number notes “Pres. by T.D.A. Cockerell. Aleurodidae det. by
Quaintance and part of original [i.e. type] lots”. Being part of Cockerell’s original material this is possibly
syntypic, but the absence of host and date raises a slight doubt and they are not regarded as paralectotypes.
A. dugesii has recently gained similar notoriety to A. dispersus although not yet increasing its range as
extensively. It causes direct feeding damage to crops, and also cosmetic problems on ornamental plants
through the unsightly appearance of its own woolly-waxy secretions (Fig. 131 graphically illustrates this), its
excreted honeydew and the sooty mould that grows on the honeydew. Although described from Mexico, and
almost certainly a native of Central America, it is the recent movement of this species into the southern states
of the U.S.A. and Hawaii that has raised its profile, provided it with a vernacular name, “giant whitefly”, and
gained it much coverage in contempory Internet reports. It has now been found in the Austro-oriental Region,
a photomicrograph of a slide-mounted specimen from the vicinity of Bogor, Java, being sent to the author for
confirmation of the determination.
Aleurodicus etiennei sp. nov.

(Fig. 54)
PUPARIUM. Habitus. Not noted. Margin. Outline oval, 1.12–1.31 mm long, 0.86–1.00 mm wide, generally
widest at abdominal segment II (n=7). Margin rather irregular, usually with submarginal wide-rimmed pores
seen in profile on puparial edge. Dorsum (Fig. 54). Longitudinal moulting suture reaching puparial margin;
transverse moulting sutures terminating anterior to compound pores on abdominal segment III. Abdominal
segment VII medially about half length of segment VI. Vasiform orifice cordate, about as long as wide, situ-
ated on a shallow elevation that extends posteriorly from pockets; operculum trapezoidal, its posterior margin
shallowly “m”-shaped, bearing a pair of stout setae similar to lingular setae; lingula head slightly acuminate,
densely covered by seta-like spinules, bearing the normal 4 stout setae; lingular apex extends to midway
between vasiform orifice and puparial margin. Chaetotaxy. A pair of long posterior marginal setae present;
single submedian pairs of eighth abdominal, cephalic, pro-, meso- and metathoracic setae present, all long and
hair-like, similar to 12 pairs of submarginal setae (including the nominal caudal pair) which extend beyond
puparial margin. Pores. Cephalic compound pores slightly smaller than anteriormost 4 pairs of abdominal
compound pores which are 50–60 μ in outer diameter, each pore with a dagger-like axial process; posterior-
most 2 pairs of abdominal compound pores relatively large, broadly bell-shaped, each with a blunt axial pro-
cess that does not protrude, pair on abdominal segment VII presenting laterally to viewer, 35–50 μ wide, pair
on segment VIII presenting obliquely with basal cells clearly visible. Immediately within puparial margin is a
very narrow band of tiny wide-rimmed simple pores, the margin usually slightly down-curled; mesal to the
wide-rimmed pore band is a widely-separated irregular double row of large double-rimmed septate pores,
with some of the submarginal setal bases situated between the two rows of pores; remainder of dorsal disc
with small septate pores with those on abdominal segments II–VI in single rows across the middle of each

segment. Venter. Ventral abdominal setae finer than dorsal setae. 2 pairs of large “sacs” of delicate ventral
cuticle present anterolateral to vasiform orifice. Legs typically robust and two-segmented, smooth, each with
a pronounced apical claw. Antennal apices extending to mid length of hind legs, their bases anterolateral to
fore legs, pointed apically. Tracheal folds absent.
ADULTS. Study material is available as listed below. Basic parameters typical for Aleurodicus.
MATERIAL EXAMINED. Holotype puparium, FRENCH GUIANA, Roura, on Annona sp. (Annon-
aceae), 20.v.1999 (J. Etienne, LNPV ref.0500080) (MNHN). Paratypes: 33 puparia, 3 adult females, 2 adult
males, same data as holotype (all on slides, BMNH, LNPV, MNHN); further adults in alcohol, same data as
holotype (LNPV); 3 puparia, SURINAM, on Citrus sp., “letter of 31.x.1968” (van Brussel 1747; USDA 68-
24068) (USNM); 2 puparia, BOLIVIA, on Papilionaceae, ii.1922 (Mann; Quaintance 23358) (USNM); 5
puparia, BOLIVIA, on Citrus (orange), vii.1921 (Mann; Quaintance 23357) (USNM). Other material: 7 third-
instar nymphs, TRINIDAD (intercepted at John F. Kennedy International Airport, U.S.A.), on Annona sp.,
02.v.1973, (Walters; USDA 73-18346) (USNM).
ETYMOLOGY. This species is named for the collector of the holotype sample, Jean Etienne, Guade-
loupe.
COMMENTS. The most characteristic feature of the puparia of A. etiennei is the submarginal widely-sep-
arated double row of large double-rimmed septate pores. Whilst the characteristics of the vasiform orifice and
lingula suggest affinities with A. neglectus and A. magnificus (compare figures 25 and 54), both of those spe-
cies have smaller double-rimmed pores that are situated within the submarginal wide-rimmed pore band (Fig.
59), and they also have many more septate pores on the dorsal disc: A. etiennei is therefore best placed within
the capiangae / dugesii group of species.
Seven third-instar nymphs, intercepted by the US quarantine service (see Material Examined, above)
appear likely to belong to A. etiennei but no puparia-associated third-instars are available for comparison.
Aleurodicus flavus Hempel

(Fig. 45)
Aleurodicus flavus Hempel, 1922a: 4. Syntypes, Brazil [MZUSP, not examined].
DISTRIBUTION. Neotropical Region – Brazil.

MATERIAL EXAMINED. 1 puparium, Brazil, 14.xii.1922 (Bondar #359, Q. 23020) (USNM); 5 puparia,
Brazil, Bahia, received 15.viii.1923 (Bondar #491, Q. 23301) (USNM); 4 puparia, Brazil, Olinda, Pernam-
buco, on Cocos nucifera, i.1972 (Arruda) (BMNH ex-USNM, B.M.1998-131).
COMMENTS. Mound & Halsey (1978) listed both MZUSP and USNM as depositories for syntypes of A.
flavus. However, the USNM collection does not contain any type material (Debra Creel, personal communica-
tion). Two slides in USNM were donated by Bondar (see above). In the species description Hempel had noted
that the original material had been sent to him for classification by Bondar, and it is therefore probable that
these are part of the original material. However, the only certain type material resides in MZUSP, quoted as
type #20,546 by Hempel. Other material has therefore been compared with the Bondar donation to USNM for
determination purposes.
This species was discussed by Russell (1965), as part of her description of A. dispersus. A. flavus is very
similar to A. dispersus, but A. flavus possesses a pair of tiny compound pores on abdominal segment VII (not
mentioned by Hempel), and has notched simple pores extending across the median line on segment VII (Fig.
45), and on segment VIII posterior to the vasiform orifice. A. charlesi, described here, is also a member of the
dispersus species-group, similarly with one pair of smaller compound pores present on abdominal segment
VII, but these are considerably larger in charlesi than in A. flavus.

Aleurodicus floccissimus (Martin, Hérnandez-Suarez & Carnero) comb. nov.
(Figs 69, 127, 140–142)
Lecanoideus floccissimus Martin, Hérnandez-Suarez & Carnero, 1997: 1262. Holotype puparium, Canary Is [BMNH,
examined].
DISTRIBUTION. Neotropical Region – Colombia, Ecuador, French Guiana, Peru, Trinidad; Palaearctic
Region – Canary Islands (La Gomera, Tenerife).
MATERIAL EXAMINED. Type material listed in description, Canary Islands and Ecuador (BMNH,
USNM); numerous specimens from all countries listed above (BMNH).
COMMENTS. Lecanoideus is here regarded as a junior synonym of Aleurodicus for reasons discussed on
p. 15. This species was described from an introduced population on Tenerife, Canary Islands. It had probably
escaped detection there for several years through coexisting with A. dispersus, which had been established
since the 1960s. At the time of its description, material from Ecuador was discovered in the BMNH collection,
confirming that this was indeed an introduction from the neotropics. More New World specimens have since
been found amongst unidentified USNM quarantine-intercepted material examined for the present study, and
it has more recently been field-collected from several other neotropical locations (BMNH).
The puparia of some populations are devoid of small compound pores on abdominal segment VIII, but
others have a pair of pores (sometimes a single pore on one side only) underneath the lingula (Fig. 69). With
specimens of both types occurring in some samples, this does appear to be intraspecific variation. A. floccissi-
mus may be distinguished from A. mirabilis, the other species formerly included in Lecanoideus, by reference
to couplet 23 of the key to species (p. 18), and comparison of figures 69 and 70.
Aleurodicus guppyii Quaintance & Baker

(Figs 62, 138)
Aleurodicus guppyii Quaintance & Baker, 1913: 59. Syntypes, Trinidad [BMNH, examined].
Aleurodicus guppyi Quaintance & Baker: Martin & Watson, 1998: 91; Martin & Mound, 2007: 50 [unjustified emenda-
tion].
DISTRIBUTION. Neotropical Region – Trinidad.

MATERIAL EXAMINED. 2 syntype puparia, Trinidad, Port of Spain, on Rheedia latiflora, received
25.v.1911 (F.W.Urich, Q. 6717) (BMNH ex-USNM, B.M. 1998-131).
COMMENTS. This species is only known from its original material. Two syntype puparia are present in
BMNH, but are not in ideal condition, the nature of the simple pores not being clear. However, it is clear that
there is no band of modified 8-shaped pores between the puparial margin and the submarginal band of wide-
rimmed pores (see couplet 26 of key to Aleurodicus species, p. 18, and fig. 62), and this character distin-
guishes A. guppyii from the similar A. juleikae. The distinct pigmented patch in the middle of the forewing
(Fig. 138) is a useful diagnostic feature.
Aleurodicus inversus Martin

(Figs 5, 23, 24)
Aleurodicus inversus Martin, 2004: 22. Holotype puparium, Belize [BMNH, examined].
DISTRIBUTION. Neotropical Region – Belize.

MATERIAL EXAMINED. Type material listed in description, Belize (BMNH, USNM).

COMMENTS. The puparial characters of A. inversus indicate that it is a close relative of A. dugesii Cock-
erell, sharing the characters of a submarginal zone of wide-rimmed and double-rimmed simple pores, the
arrangement of its six pairs of abdominal compound pores, and submedian area having only sparse septate and
minute bright pores. However, A. inversus is immediately distinguished by the zone of wide-rimmed pores
occupying the subdorsum as well as the submargin (encompassing abdominal compound pore pairs 1–4), by
the pronounced lobulate inner margin of this zone (Fig. 23), and by the presence of septate double-rimmed
pores mesal to abdominal compound pore pairs 1–4 and between the 4th and 5th/6th pairs of abdominal com-
pound pores; in contrast, the wide-rimmed pores in A. dugesii are not obviously septate, and are only present
in the submargin (Fig. 22). The possession of a broad zone of wide-rimmed pores and a submarginal row of
double-rimmed septate pores by A. inversus also indicates similarity to the spiralling whitefly, A. dispersus
Russell, but A. dispersus has only 4 pairs of abdominal compound pores (all large), its double-rimmed septate
pores are notched rather than septate, these are in a single outer submarginal row only, and its submedian area
is very densely provided with smaller notched pores, in contrast to the sparse submedian septate pores in A.
inversus.
All the recorded host plants for A. inversus are woody forest vines in Belize, and this whitefly species dis-
plays a clear preference for such plants.
Aleurodicus jamaicensis Cockerell

(Fig. 61)
Aleurodicus jamaicensis Cockerell, 1902: 280. Holotype puparium, Jamaica [USNM, examined].
DISTRIBUTION. Neotropical Region – Jamaica.

MATERIAL EXAMINED. Holotype puparium, Jamaica, Kingston, “postal May 26 1903 from
T.D.A.Cockerell”, Q.3276, Type no. 14771 (USNM, 8999); numerous puparia, Jamaica, on Cocos nucifera
(BMNH).
COMMENTS. A. jamaicensis was described from a single specimen and the type specimen is therefore
properly a holotype, not a syntype as listed in Mound & Halsey (1978). The holotype has a very large thoracic
parasitoid emergence hole, and was not cleared of debris or otherwise treated prior to mounting. When
redescribing it, Quaintance & Baker (1913) readily admitted that their own redescription was wanting,
because of the condition of the specimen. Through the kindness of USNM, the author was permitted to
remount this specimen. Remounting has allowed the dorsum and venter to be separated, the parasitoid’s pupal
remains to be removed and mounted adjacent to the puparium, maceration of remaining body contents, de-
waxing and staining. The result is a great improvement, and it can be seen that parasitoid emergence occurred
when the adult whitefly, a male (its claspers are still present), was forming within its puparium. The type prep-
aration retains its original labels, with the addition of a circular “holotype” label.
Submarginal chaetotaxy appears to comprise the 12 setal pairs typical for Aleurodicus, these setae rather
short and lining the mesal boundary of the wide-rimmed pore band (Fig. 61): however, cephalothoracic sub-
median setae have been destroyed by the emergence of the parasitoid. A. jamaicensis belongs to the pulvina-
tus-group of species, with a submarginal band of wide-rimmed pores, and its dorsal disc with scattered septate
pores. However, there are small numbers of bright pores anterior to the small compound pores on abdominal
segment VII but none adjacent to the compound pores on abdominal segment III. Most distinctive of all,
within the pulvinatus group, is the absence of small compound pores on abdominal segment VIII.

Aleurodicus juleikae Bondar
(Figs 63, 64)
Aleurodicus juleikae Bondar, 1923: 78. Lectotype puparium here designated, Brazil [BMNH].
DISTRIBUTION. Neotropical Region – Brazil, Colombia, Dominican Republic, Ecuador, Guyana, Peru,
Venezuela.
MATERIAL EXAMINED. Type material as detailed and discussed below (BMNH, USNM); 1 puparium,
presumed Brazil, on Psidium sp., (Bondar #764, Q.23007), labelled “Aleurodicus candidus” [a manuscript
name only] (USNM); puparia with the following country / host combinations – Brazil: Eucalyptus, Tri-
umpheta, Persea, Psidium, Casearia, “Psytachantus”; Colombia: Matisia, Tectona; Dominican Republic:
Annona; Ecuador: Anthurium, palm; Guyana: Avicennia; Peru: Citrus, Cocos, Mangifera, Persea, Tricho-
philia, palm; Venezuela: “Melicoccus” (BMNH, USNM).
COMMENTS. A. juleikae is one of his then newly-described whitefly species of which Bondar donated
material to BMNH and USNM, the material regarded as syntypic (see p. 9). A single slide of Bondar material
in BMNH, probably made in the 1920s, contains 4 puparia in excellent condition, and bears the data “Aleuro-
dicus juleikae Bondar, Brazil, Bahia, on Loranthaceae, Pres. Dr. G. Bondar, 7.xii.23, B.M.1924/44, 671”. As
part of this study, four additional slides (containing 14 puparia and 3 third-instar nymphs) have been made
from the dry residue of Bondar’s donation to BMNH, but the intervening eighty years have led to the speci-
mens being rather abraded, with many setae having been lost. One slide in USNM contains 7 puparia, but the
mountant is thicker than optimum and detail is not so readily visible as in the 1920s BMNH slide, the data
being “Q[uaintance] 23300, Aleurodicus juleikae, Bahia, Brazil, G. Bondar, #671”. Another USNM slide
bears the data “”Q.23311, 672 - B. phrygilank [sic], 671 – A. juliskae [sic], Bahia, Brazil, G. Bondar”, and
contains 10 adults along with one puparium that appears to be A. juleikae. From the Bondar number, 671, the
material in both the American and British samples appears to be from the same original sample. Material dis-
playing a range of characters, and collected from a number of hosts, has been determined as “A. juleikae, or
similar” and it is important to fix a type against which other samples can be compared: a lectotype is therefore
here designated for A. juleikae, and is one of the 4 specimens on the 1920s BMNH slide, on which it is
clearly indicated. The remaining specimens on the BMNH slides, and all on the 7-puparia USNM slide, are
paralectotypes: the status of the mixed slide (Q. 23311) is uncertain.
Only two characters distinguish the type material of A. juleikae from the great majority of specimens of A.
pulvinatus (Maskell), p. 40. The largest compound pores of the type sample of A. juleikae are up to 50μ in
outer diameter, somewhat larger than is typical for A. pulvinatus: however, Martin & Watson (1998: 94) rede-
fined A. pulvinatus with the size of the largest pair of compound pores (anteriormost abdominal pair) ranging
from 25–48μ. Further, a sample from Poraqueiba sericea (Icacinaceae) in Peru has compound pores up to
60μ in diameter but is otherwise exactly as the lectotype puparium of A. pulvinatus, including the outline
being flattened anteriorly and each puparium having a darkly-pigmented longitudinal subdorsal band on each
side of the puparium. It is concluded that compound pore size is not significant in distinguishing these two
species.
The second character used to separate A. juleikae from A. pulvinatus is the nature of the submarginal band
of wide-rimmed simple pores. In A. pulvinatus the mesal boundary of this pore band does not closely
approach the abdominal compound pores, and the boundary is not therefore affected by their proximity (Figs
67, 68): in contrast, the mesal boundary of the pore band in A. juleikae is much closer to the compound pores,
and is sinuous, often almost interdigitating with the compound pores (Figs 63, 64). This character does appear
to be more reliable and specimens with this character, but otherwise answering the diagnosis of A. pulvinatus,
are determined as A. juleikae. Figures 124 and 125 are photographs of living puparial colonies that have been
cautiously determined as A. juleikae.

A. cocois (Curtis) (see p. 26 and figure 65) clearly belongs to this same group of species, differing from
both A. juleikae and A. pulvinatus in having the mesal boundary of the submarginal pore band actually inter-
digitating with the abdominal compound pores. Also, in A. cocois the mesal boundary of the pore band in the
meso- and metathorax is almost straight, parallel to the longitudinal moulting suture, emphasised by a slight
cuticular fold. The species limits of A. cocois, A. pulvinatus and A. juleikae are not satisfactorily understood,
however, with intermediate forms occurring.
Aleurodicus magnificus Costa Lima

(Figs 6, 25, 58)
Aleurodicus magnificus Costa Lima, 1928: 129–131. Lectotype puparium here designated, Brazil [USNM].
DISTRIBUTION. Neotropical Region - Belize, Brazil, Ecuador, Panamá.

MATERIAL EXAMINED. Type material detailed and discussed below (USNM); puparia with the fol-
lowing country / host combinations – Belize: Tetracera; Brazil: ?Annona; Ecuador; Annona; Panamá: Tetrac-
era, Nectandra, Unonopsis, Vismia (BMNH).
COMMENTS. A slide bearing type data was loaned to the author by USNM as part of this study, although
no depository was quoted by Mound & Halsey (1978). The data are “Aleurodicus magnificus Costa Lima, on
orange, S. Lourenço, Sul de Minas, Brazil, 1921, from Costa Lima 1949”. The slide contains seven puparia,
two of them noticeably smaller than the other five and perhaps representing sexual dimorphism. For the rea-
sons stated here on p. 9, a lectotype has been here designated from amongst the specimens on this USNM
slide, and is clearly indicated.
This species secretes a dense, woolly, tangle with particularly coarse strands of secretion. Colonies of
puparia often appear dirty, greyish. Most commonly, puparia are aligned along the undersides of leaf midribs,
rather than in broader aggregations - reminiscent of the species formerly included in Lecanoideus, particularly
A. mirabilis (see below).
Morphologically, A. magnificus is extremely similar to A. neglectus, and the two species’ general appear-
ance is as in figures 6 and 25; the two species comprise a small species-group, differing in the distribution of
simple pores on the dorsal disc – see key to species, couplets 4 & 5 (p. 16) and figures 58 and 59. Both species
lack notched simple pores on abdominal segments I and VII.
Aleurodicus maritimus Hempel

(Figs 7, 26)
Aleurodicus maritimus Hempel, 1922b: 1160–1161. Syntypes, Brazil [MZUSP, examined].

Aleurodicus linguosus Bondar, 1923: 76–78. Syntypes, Brazil [USNM, examined]. Synonymised by Costa Lima, 1928:
133.
DISTRIBUTION. Neotropical Region - Belize, Brazil, Colombia, Ecuador, French Guiana, Guyana, Mexico,
Nicaragua, Panamá, Surinam, Trinidad.
MATERIAL EXAMINED. Syntypes of A. maritimus and A. linguous as discussed below (MZUSP,
USNM); numerous samples from all countries listed above (BMNH).
COMMENTS. The author was able to examine syntypes of both A. linguosus and A. maritimus through
the kindness of USNM and MZUSP, respectively, in connection with an earlier study of aleurodicine white-
flies in Belize (Martin, 2004). Costa Lima (1928) considered A. linguosus to be a junior synonym of A. marit-
imus, and his opinion is considered sound by the present author.
A. maritimus is a member of the capiangae / dugesii species-group.

Aleurodicus mirabilis (Cockerell), revived combination
(Figs 70, 128, 130, 143, 144)
Aleurodes mirabilis Cockerell, 1898: 225. Syntypes, Mexico [USNM, not examined].
Aleurodicus mirabilis (Cockerell) Cockerell, 1899: 360.
Aleurodicus (Lecanoideus) giganteus Quaintance & Baker, 1913: 70. Syntypes, Brazil [USNM, not examined]. [Syn-
onymised by Martin, 2004: 18]
Lecanoideus mirabilis (Cockerell) Mound & Halsey, 1978: 243.
DISTRIBUTION. Neotropical Region – Brazil, Colombia, Ecuador, Guyana, Mexico, Trinidad.

MATERIAL EXAMINED. Samples from all countries listed above (BMNH).
COMMENTS. Lecanoideus is here regarded as a junior synonym of Aleurodicus for reasons discussed on
p. 15. The decision to regard L. giganteus as a junior synonym of L. mirabilis was based upon an examination
of syntype material at USNM by Steve Nakahara (personal communication), and was discussed by Martin
(2004).
The immature stages of A. mirabilis secrete particularly thickly flocculent waxy material (Fig. 143) that
can obscure the adults completely unless lifted clear (Fig. 144), and colonies frequently line major leaf veins
(Fig. 128), a characteristic also seen in A. floccissimus, and in A. magnificus (see above). A. mirabilis differs
from A. floccissimus as detailed in key couplet 23 (p. 18) and figures 69 and 70. Adults leave particularly
broad, white, trails under leaves whilst ovipositing (Fig. 130).
Aleurodicus neglectus Quaintance & Baker

(Fig. 59)
Aleurodicus neglectus Quaintance & Baker, 1913: 63. Lectotype puparium here designated, Guyana [BMNH].
Aleurodicus flumineus Hempel, 1918: 211. Syntypes, Brazil [BMNH, USNM, examined]. Synonymised by Bondar,
1923: 61.
DISTRIBUTION. Neotropical Region – Barbados, Brazil, Guyana, Trinidad.

MATERIAL EXAMINED. Type material of A. neglectus detailed and discussed below (BMNH, USNM);
7 puparia, syntypes of A. flumineus Hempel, Brazil, Rio de Janeiro State, on Moquilea tomentosa, i.1916
(Moura via Costa Lima, USNM #10075) (BMNH ex-USNM, B.M.1998-131); puparia with the following
country / host combinations – Barbados: Annona; Brazil: Moquilea, Eriotheca; Guyana: Ceiba, Vismia, Tet-
racera, Coussapoa; Trinidad: Annona, Cocos (BMNH).
COMMENTS. A. neglectus was described from a mixture of material from several hosts in Brazil, Guy-
ana and Trinidad. Syntypic material of A. neglectus available to the author comprised one USNM slide with
only a forewing [guava, Brazil, 1882, Koebele, no. 3160], one USNM slide containing 4 much-distorted post-
emergence pupal cases [Annona reticulata, Trinidad, 28.ix.(18)96, 2/3166], and a series of BMNH slides from
Guyana [Annona squamosa, Demerara, Sept. 1892, no.126 BWIs 112, ex- Newstead collection]. The BMNH
Demerara slides are mostly preparations of adults, but one slide contains post-emergence pupal cases of which
one is in good condition and displays most characters well: this best specimen has been here designated the
lectotype and is clearly indicated on its slide.
A. neglectus and A. magnificus are extremely similar and together form a small species-group. They may
be distinguished by use of the key to species, couplets 4 & 5 (p. 16), figures 58 and 59, and comments on A.
magnificus (p. 35).

Aleurodicus nicaraguensis sp. nov.
(Fig. 47)
PUPARIUM. Habitus. Not noted. Margin. Outline broadly oval, 1.08–1.25 mm long, 0.81–0.93 mm wide,
generally widest at abdominal segment II (n=8). Dorsum. Longitudinal moulting suture reaching puparial;
transverse moulting sutures terminating in outer subdorsum. Abdominal segment VII much reduced medially,
but pockets not reaching into segment VI. Vasiform orifice rounded-cordate, anterior margin poorly defined,
situated on a shallow elevation; operculum trapezoidal, its posterior margin straight to faintly emarginate,
bearing a pair of fine setae similar to eighth abdominal pair; lingula head tongue-shaped, densely covered by
seta-like spinules, bearing the normal 4 stout setae, its apex almost reaching bases of caudal setae. Chaetotaxy.
A pair of posterior marginal setae present, similar to 12 pairs of submarginal setae (including nominal caudal
pair) whose bases are mesal to submarginal simple pores; single pairs of eighth abdominal and cephalic sub-
median setae present, both pairs about half length of submarginal setae but longer than submedian pairs of
pro-, meso- and metathoracic setae. Pores. Cephalic compound pores slightly smaller than anterior 4 pairs of
abdominal pairs, most of which are 60–75μ in outer diameter and with broad, dagger-like axial processes;
small pairs of compound pores on abdominal segments VII and VIII are about 25μ in diameter. Submargin
defined by a band of crowded wide-rimmed simple pores, mesal boundary of the band interdigitating with a
single row of large double-rimmed septate pores 20–25 μ in outer diameter, their spacing a little uneven (Fig.
47), 23 on each side in holotype; remainder of dorsal disc very sparsely provided with extremely small septate
pores. Venter. Ventral abdominal setae similar to eighth abdominal dorsal setae, underlying vasiform orifice.
Legs typically robust and two-segmented, smooth, each with a pronounced apical claw. Antennae long, corru-
gate-sided, apically acute, apices extending to slightly beyond articulation of hind legs, bases anterolateral to
fore legs. Tracheal folds absent.
MATERIAL EXAMINED. Holotype puparium, NICARAGUA, Rio San Juan / Rio Bartola confluence,
on unidentified woody broadleaf host, 21.vi.2004 (J.H.Martin #8070) (BMNH). Paratypes: 6 puparia, 1 adult
female, same data as holotype (BMNH); 3 puparia, 1 adult male, same locality as holotype, on unidentified
woody broadleaf host, 20.vi.2004 (Martin #8059) (BMNH).
ETYMOLOGY. This species is named for its country of collection, Nicaragua.
COMMENTS. The presence of very large double-rimmed septate pores in the submargin is most reminis-
cent of A. maritimus but possessing a broad submarginal band of wide-rimmed pores suggests possible closer
affinities with other members of the capiangae / dugesii species-group, to which A. nicaraguensis belongs.
Aleurodicus niveus Martin

(Figs 8, 33, 121)
Aleurodicus niveus Martin, 2004: 25. Holotype puparium, Belize [BMNH, examined].
DISTRIBUTION. Neotropical Region – Belize, Brazil, El Salvador, Guatemala, Honduras, Mexico, Nicara-
gua, Panamá (including Canal Zone), Venezuela – see discussion, below.
MATERIAL EXAMINED. Holotype puparium, BELIZE, CFR, Las Cuevas forest, on Encyclia cochleata
(Orchidaceae), 04.vi.2004, (J. H. Martin #7986) (BMNH). Paratypes: 17 puparia (7 dry on leaf), same data as
holotype (BMNH, USNM). Non-type material: 1 puparium, VENEZUELA (intercepted at Miami, U.S.A.), on
Oncidium sp., 14.iv.1981 (USNM); 3 puparia, BRAZIL (intercepted at Washington DC, U.S.A.), on Oncidium
sp., 14.iv.1938 (USNM); numerous specimens from orchids, intercepted in U.S.A. from Central American
countries (see below) (BMNH, USNM).
COMMENTS. This species was described from a single colony, a group of puparia under one leaf of their
epiphytic orchid host in secondary rainforest in the Chiquibul Forest Reserve in Belize: a colour photograph

of this colony (Martin, 2004: 119) is reproduced here, fig. 121). Most individuals of the type sample proved to
be parasitised, but five undamaged specimens were preserved in alcohol as soon as this became evident: three
of these show no evident signs of parasitism under high magnification, and it is thought unlikely that the habi-
tus appearance of the colony was affected by the development of parasitoids (but see below).
The features of the vasiform orifice, operculum and lingula (Fig. 33) are highly characteristic, with the
very large operculum and acute, often bifid, apical lingular process of particular note. In other characters, A.
niveus most closely resembles A. rugioperculatus, also described from Belize.
Only the small type sample was known to the author at the time of its description. However, much un-
named slide-mounted aleurodicine material was made available for the present study, through the kindness of
USNM. Amongst this material were numerous slides that had been categorised as “species 1” (120 slides +) or
“species 2” (250 slides +) of a “new genus close to Aleurodicus”. All of these slides represented quarantine
interceptions at US ports, and almost all had been taken from the foliage of orchids imported into the U.S.A.
from Central American countries. Although displaying a degree of variability, particularly in overall puparial
size, apparent degree of development of axial processes in the compound pores, and presence / absence of
small compound pores on abdominal segment VIII, all of this material has been here determined as A. niveus,
which may be either a somewhat variable species or a species-complex. The original, pencilled assignment to
“species 1” or “species 2” has been left visible on each slide, to aid any future investigation of the nature of
variation in this species or species-group.
Listing the examined quarantine-intercepted material in full has not been practical, with so many samples
involved, and only the countries of distribution are listed above.
Aleurodicus ornatus Cockerell

(Fig. 53)
Aleurodicus ornatus Cockerell, 1893: 105. Lectotype puparium here designated, Jamaica [USNM].
DISTRIBUTION. Neotropical Region – Haiti, Jamaica, Puerto Rico.

MATERIAL EXAMINED. Type material detailed and discussed below (USNM); 3 puparia, Jamaica,
Bath, on Cocos nucifera, 21.v.1969 (Heinze) (BMNH).
COMMENTS. A slide of a single puparium of this species, bearing a USNM type label, has one label
bearing the data “Aleurodicus ornatus Ckll, type no.14773, USNM”, and other with “5673, Box 2, Q 6782,
Cotype”. With such sparse data on this slide, Debra Creel (USNM) very kindly investigated, with the follow-
ing report that I quote: “……another slide [with adults only] ….. says ‘Aleurodicus ornatus Ckll. Q6782.
Remounted from slide with pupa [the slide whose data are quoted above] Type No. 14773’ with “capsicum”
and “Jamaica” in pencil. …….an envelope of dry material says ‘A. ornatus Ckll on Capsicum, Kingston
Jamaica, from T D A Cockerell 1903, Q3284’ on the outside, and inside are a pupa, adult and leaf material
with data ‘5673 on Capsicum Kingston Jamaica’. The year, 1903, may be a year of transfer from Jamaica (as
with Bondar material sent from Brazil and discussed on p. 9), thus not precluding the material being part of
the original sample that was described in 1893. The data for “5673” are also consistent with the quoted type
data but no collection date is given. It is considered that the dry material, and single-puparium slide, are part
of Cockerell’s original sample, sent from Jamaica, presumably to Quaintance. The single puparium from the
dry envelope has been slide-mounted as part of this study and is here designated as the lectotype, which is
clearly indicated; on the same slide is a single adult forewing, the only other insect material that was contained
in the envelope.
A. ornatus is the only known member of the capiangae / dugesii species-group to possess but a single pair
of small compound pores, on abdominal segment VII (Fig. 53). Three puparia in BMNH (data above) closely

resemble the lectotype of A. ornatus in having small compound pores only on abdominal segment VII, but
they have more numerous submarginal double-rimmed pores which are, thus, closer together: these specimens
(Fig. 53) are determined as A. ornatus with caution.
Aleurodicus pauciporus Martin

(Figs 9, 27, 28)
Aleurodicus pauciporus Martin, 2004: 26. Holotype puparium, Belize [BMNH, examined].
DISTRIBUTION. Neotropical Region – Belize, Costa Rica, [Panamá], [Venezuela].

MATERIAL EXAMINED. Type material listed in description, Belize (BMNH, USNM); 9 puparia, Costa
Rica, Guanacaste Province, Santa Rosa NP, on undetermined host, 06.iii.1990 (Cox) (BMNH); 7 puparia,
Costa Rica, northern Heredia Province, 10 km north of Puerto Viejo, on undetermined host, 02.ii.1983 (Mar-
tin) (BMNH); 2 puparia, Ecuador, Manabi Province, Pedernales, on Psidium guajava, 11.ii.2005 (Martin)
(BMNH); 1 puparium, Panamá, v.1912 (Busck, Q. 8801) (USNM), 5 puparia, Panamá, Canal Zone, Barro
Colorado Island, 20.iii.1923 (Wheeler-Zetek, Q. 23110) (USNM); 3 puparia, 2 third-instar nymphs, Venezu-
ela, Turmero, Aragua, 01.iii.1982 (Arnal / Debrot #5) (USNM).
COMMENTS. The most notable feature of the puparia of this species is the paucity of dorsal simple
pores, which are more numerous, and of more types, in most species of Aleurodicus. The lack of simple pores
is reflected in similarly sparse secretion by the immature stages, with the puparia of some colonies distinctly
brownish submarginally. Nevertheless, the nature and distribution of the compound pores, and the puparial
chaetotaxy, meet the generic definition of Aleurodicus. Within the genus, A. pauciporus is the only described
species with such scarcity of simple pores. The puparia from Costa Rica, Ecuador, Panamá and Venezuela
(listed above) are tentatively determined as A. pauciporus but have a higher density of submarginal pores, but
this pore band is still poorly defined mesally.
This species is cautiously treated as a member of the talamancensis species-group.
Aleurodicus pseudugesii sp. nov.

(Figs 43, 44)
PUPARIUM. Habitus. Notebook observation stated only that waxy secretions were present, opaque and rather
plain – possibly through being worn by leaf movement. Colony was present under one leaflet of its coconut
palm host. Margin. Outline oval, 0.98–1.20 mm long, 0.67–0.82 mm wide, generally widest at abdominal seg-
ment I/II (n=15). Margin. True margin almost smooth, but margin usually slightly down-curled and submar-
ginal pores in relief lend visible edge of puparium a roughened appearance (Fig. 43). Dorsum. Longitudinal
moulting suture reaching puparial margin; transverse moulting sutures almost reaching as far as row of com-
pound pores. Abdominal segment VII extremely reduced medially, such that only 7 abdominal segments dis-
cernible medially; abdominal segmentation suture-like as far laterad as row of compound pores; meso- /
metathoracic division marked as a thickened furrow, pro- / mesothoracic division rather more subtly indicated
submedially only. Vasiform orifice (Fig. 44) cordate, about as long as wide, smooth; operculum trapezoidal,
dorsally smooth, its posterior margin very shallowly “m”-shaped, bearing a pair of long setae; lingula head
elongate, acute, only its extreme apex rounded, densely covered by seta-like spinules, bearing the normal 4
stout setae near its apex; lingular apex closely approaches, but does not overlap, puparial margin. Chaetotaxy.
A pair of posterior marginal setae, pair of eighth abdominal setae, 4 pairs of submedian cephalothoracic setae,
and 12 pairs of outer submarginal setae present, all long and fine, the submarginals with most of their lengths

extending beyond puparial margin. Pores. Anterior 4 pairs of abdominal compound pores all similar in size,
most 50–60μ in outer diameter, each with a broad, dagger-shaped axial process extending beyond pore mouth;
cephalic compound pores a little smaller, with axial processes proportionately narrower; posterior 2 pairs of
small abdominal compound pores rather large, 30–40μ in diameter, both pairs appearing broadly bell- or
medusa-shaped in lateral view (Fig. 44). Immediately within puparial margin are a few ranks of tiny wide-
rimmed pores (Fig. 43); lining mesal boundary of wide-rimmed pore band are widely-spaced small double-
rimmed septate pores (Figs 43, 44 ), apparently 13 on one side and 16 on the other in holotype (mesal to these
small double-rimmed pores boundary of narrow submargin is effectively defined by submarginal setal bases).
Abdominal segments VII and VIII each with a loose array of large bright simple pores anterior to the small
compound pore on the respective segment (Fig. 44); remainder of dorsal disc sparsely provided with a mixture
of tiny septate pores and minute porettes. Venter. Ventral abdominal setae similar to dorsal setae; other charac-
ters typical for Aleurodicus.
THIRD-INSTAR NYMPH. Outline oval, 0.68–0.77 mm long, 0.42–0.49 mm wide, widest slightly poste-
rior to mid-length (n=6). Margin smooth. Vasiform orifice, operculum and lingula as in puparium, lingula
slightly overlapping edge of body. 2 pairs of large compound pores present in cephalothorax, broadly medusa-
shaped, 30–40μ in width, each with an axial process that just reaches pore mouth. 3 pairs of small cicatrices
present, two pairs in cephalothoracic area and third pair posterolateral to vasiform orifice, these being the
scars of compound pores in second-instar; positions of puparial compound pores clearly visible in all speci-
mens, as pale circular patches, but no specimen apparently close to ecdysis. Chaetotaxy as in puparium, setae
as long as in puparium. Dorsum with pores generally distributed as in puparium, but double-rimmed pores
absent and bright pores on abdominal segments VII/VIII much less numerous. Legs triangular, evidently 3-
segmented, each with a curved apical claw; antennae small, about same size as apical segments of legs; ven-
tral abdominal setae similar to dorsal setae.
ADULTS. Study material available as listed below. Basic parameters typical for Aleurodicus.
MATERIAL EXAMINED. Holotype puparium, ECUADOR, Orellana Province, military campus near
Coca, on Cocos nucifera (Arecaceae), 16.ii.2005 (J.H.Martin #8146) (BMNH). Paratypes: 28 puparia, 6 third-
instar nymphs, 9 adult males, 9 adult females, same data as holotype (BMNH, USNM); 1 puparium, PERU,
Junin, on Cocos nucifera, 19.viii.2005 (Lozada) (BMNH).
ETYMOLOGY. The specific epithet refers to the similarities between this species and A. dugesii.
COMMENTS. A. pseudugesii most closely resembles A. dugesii Cockerell and A. araujoi Sampson &
Drews. The presence of a single row of widely spaced double-rimmed pores, marking the inner boundary of a
submarginal band of wide-rimmed pores suggests closest affinity with A. dugesii, but A. dugesii has larger but
non-septate double-rimmed pores (inner part of pore merely notched on one side), larger wide-rimmed pores
in a broader band, does not have arrays of bright pores on abdominal segments VII and VIII, and lacks a pair
of cephalic submedian setae. A. araujoi, also one of the capiangae / dugesii species-group, shares with A.
pseudugesii the array of bright pores on each side of abdominal segment VII (but not also on segment VIII as
in pseudugesii), and a rather acuminate lingula; however, A. araujoi (Fig. 21) has many more, much larger,
double-rimmed septate pores, in an uneven row, these surrounded by sparse tiny simple pores that may be of
the wide-rimmed variety but which do not form a distinct band and some of which extend mesal to compound
pores and submarginal setal bases.
Aleurodicus pulvinatus (Maskell)

(Figs A, 10, 29, 67, 68, 139)
Aleurodes pulvinata Maskell, 1896: 439. Lectotype puparium, Trinidad [designated by Martin & Watson, 1998: 95,
USNM, examined].
Aleurodicus pulvinatus (Maskell); Cockerell, 1902: 280.

Aleurodicus iridescens Cockerell, 1898: 225. Lectotype, Mexico [designated by Martin & Watson, 1998: 95, USNM,
examined]. A. iridescens was erroneously synonymised with A. cocois (Curtis) by Quaintance & Baker, 1913: 47,
but subsequently synonymised with A. pulvinatus by Martin & Watson, 1998: 93.
Aleurodicus bifasciatus Bondar, 1922: 85. Synonymised by Bondar, 1923: 66.
DISTRIBUTION. Neotropical Region - widely distributed, especially amongst the Caribbean islands.
MATERIAL EXAMINED. Type material as detailed by Martin & Watson (1998); numerous samples,
from many South and Central American countries (BMNH, CDFA, USNM).
COMMENTS. A. pulvinatus was investigated by Martin & Watson (1998), who redefined this species,
concluding that puparial pigmentation, and the size of the large compound pores (25–48 μ in diameter) varies
considerably. Puparia may be completely pale, have pronounced subdorsal pigmentation in a longitudinal
stripe on each side of the puparium, or possess more limited marking in the form of a pair of thoracic patches
or, occasionally, a median thoracic patch (Figs 67, 68). This species is widespead in the Caribbean area, where
it sometimes causes considerable damage to coconut palms.
As discussed in the comments on A. juleikae (q.v.), the species limits of A. pulvinatus, A. cocois and A.
juleikae will probably require further detailed investigation to resolve satisfactorily. As currently understood,
these three species may be distinguished by use of couplets 27 and 28 of the species key, p. 19.
Aleurodicus rugioperculatus Martin

(Figs 12, 14–16, 32)
Aleurodicus rugioperculatus Martin, 2004: 29. Holotype puparium, Belize [BMNH, examined].
DISTRIBUTION. Neotropical Region – Belize, Guatemala, Mexico.

MATERIAL EXAMINED. Type material as detailed in description, Belize, Mexico (BMNH, USNM);
non-type material – 2 puparia, 3 adult females, Guatemala, on palm, 18.iii.1913 (E. Bethel / Quaintance
#8826) (USNM); several samples, Mexico, on Caryocar amygdaliferon, Cocos nucifera, Melia sp, Musa sp.,
Persea americana and Psidium guajava, various dates and collectors (USNM).
COMMENTS. The lingular apex of this species is atypical for Aleurodicus, being narrowly acute, its 4
setae situated very close to the apex (Fig. 16). The subtle reticulate patterning of the cuticle in the cephalotho-
rax and abdominal subdorsum (Figs 14, 15), seen in many specimens, is also unusual as is the sometimes
extensive dark, cloudy cuticular pigmentation seen in some individual puparia. A. rugioperculatus is most
similar to A. niveus (also described from Belize) and may be distinguished from it by use of the key to species
of Aleurodicus, couplets 30-31, p. 19.
Aleurodicus septatissimus sp. nov.

(Figs. 48, 49)
PUPARIUM. Habitus. Unknown. Margin. Outline oval, 1.61 mm long, 1.14–1.18 mm wide, generally widest
at abdominal segment II/III (n=3). Margin rather variably crenulate, caused by proximity of outer-submar-
ginal double-rimmed septate pores. Dorsum. Cuticle slightly pigmented, brownish, in dorsal disc area of tho-
rax with pigmentation rhachis-like in appearance on abdominal segments II–VI. Longitudinal moulting suture
reaching puparial margin; transverse moulting sutures faintly marked almost to puparial margin. Abdominal
segment VII much reduced medially, less than half median length of segment VI. Vasiform orifice broadly
cordate, wider than long, smooth; operculum trapezoidal, surface roughened-granular, its posterior margin
faintly convex and bearing a pair of stout setae; lingula head elongate-oval, densely covered by seta-like

spinules, bearing the normal 4 stout setae; lingula extending about half way from posterior margin of vasiform
orifice to puparial margin. Chaetotaxy. A pair of posterior marginal setae the longest on puparium, up to 120μ
long; eighth abdominal setae present anterior to vasiform orifice, similar to 12 pairs of submarginal setae, all
very fine (Fig. 48) except for nominal caudal pair which are more robust and similar to posterior marginals;
submarginals not, or hardly, reaching puparial margin; 3 pairs of submedian thoracic setae present, very small
and fine, cephalic pair wanting. Pores. Cephalic and anterior 4 pairs of abdominal compound pores all similar,
largest (usually on abdominal segment III) up to 55μ in outer diameter; axial processes of large compound
pores, faintly indicated, apically rounded and extending only slightly beyond pore mouth. 2 pairs of very
small compound pores present, 1 pair each on abdominal segments VII and VIII, only about 20μ in diameter.
Immediately within puparial margin are 2 ranks of large double-rimmed septate pores, outer rim of each pore
contiguous with pores in adjacent rank and with neighbouring pores in same rank (Figs 48, 49); remainder of
dorsal disc with numerous small septate pores; subdorsal region of abdominal segments III–VII (ends of the
pigmented apparent rhachis arms) with loose groupings of tiny wide-rimmed bright pores, these more sparse
on segment VII. Venter. Ventral characteristics typical for Aleurodicus.
MATERIAL EXAMINED. Holotype puparium, PERU (intercepted at Miami, U.S.A.), on unidentified
Orchidaceae, 11.viii.1965 (Buff, 35932, 65-16856) (USNM). Paratypes: 2 puparia, same data as holotype, on
same slide (USNM).
ETYMOLOGY. The specific epithet a noun in aposition, created to reflect the extreme density of septate
pores.
COMMENTS. The three specimens of A. septatissimus are unlike those of any other known species of
Aleurodicus, possessing a complete double-ranked outer-submarginal row of large double-rimmed septate
pores that are almost contiguous. Its affinities appear to lie with the members of the capiangae / dugesii spe-
cies-group. As with any species known only from quarantine interceptions, the precise geographical location
of its domicile remains unknown.
Aleurodicus spectabilis sp. nov.

(Figs 50, 51)
PUPARIUM. Habitus. Not known. Margin. Outline oval, 1.42–1.45 mm long, 1.07–1.14 mm wide, generally
widest at abdominal segment II (n=3). Margin smooth but immediately within margin lies a single row of
almost contiguous crescent-shaped glands (Fig. 50). Dorsum. Longitudinal moulting suture reaching puparial
margin; transverse moulting sutures terminating in subdorsum anterior to abdominal segment III compound
pores. Abdominal segment VII extremely reduced medially such that only 7 abdominal segments readily
apparent anterior to vasiform orifice. Vasiform orifice cordate, wider than long; operculum rounded-trapezoi-
dal, its posterior margin very slightly convex, bearing a pair of fine setae which are not obscured by sides of
lingula; lingula head tongue-shaped, densely covered by seta-like spinules, bearing the normal 4 stout setae;
lingula (Fig. 51) extending about two-thirds of distance from posterior edge of vasiform orifice towards
puparial margin. Chaetotaxy. A pair of posterior marginal setae and pair of eighth abdominal setae present,
similar to 12 pairs of submarginal setae (including the nominal caudal pair), all long and fine, caudal pair up to
130 μ long; single submedian pairs of pro-, meso- and metathoracic setae present, minute, no more than 12 μ
in length, but cephalic submedian pair absent. Pores. Cephalic compound pores similar to anteriormost 4 pairs
of abdominal compound pores which are maximally 40–50 μ in outer diameter, each with a dagger-shaped
axial process extending beyond pore mouth; posteriormost 2 pairs of abdominal compound pores, on seg-
ments VII and VIII (Fig. 51), extremely small, 12–18 μ in maximum diameter. Mesal to marginal crescent-
shaped glands, submargin is punctuated by a broad band of very large double-rimmed 8-shaped septate pores
(20–25 μ in longest dimension), these pores mostly in 3 distinct ranks (Fig. 50); remainder of dorsal disc

(except area immediately surrounding vasiform orifice, the median part of posterior abdominal segments, and
areas surrounding cicatrices) crowded with large notched pores (Fig. 50), with the notched nature of the pores
most evident in more median areas; a few minute bright simple pores present adjacent to small compound
pores on abdominal segments VII and VIII. Venter. Ventral abdominal setae similar to dorsal submarginals.
Legs typically robust and two-segmented, smooth, each with a pronounced apical claw. Antennal apices
extending to, their bases anterolateral to fore legs, pointed apically. Tracheal folds absent.
MATERIAL EXAMINED. Holotype puparium, BRAZIL, Minas Gerais, Vicosa, on satsuma citrus (Ruta-
ceae), 10.v.1932 (E. J. Hambleton #858) (USNM). Paratypes: 2 puparia, same data as holotype, on same slide.
ETYMOLOGY. The specific epithet is the Latin word spectabilis, the dictionary meaning given as “worth
looking at”.
COMMENTS. The puparia of A. spectabilis are large for a species of Aleurodicus and the submarginal
ranks of large double-rimmed 8-shaped septate pores, combined with very crowded notched pores on the dor-
sal disc, render the species readily recognisable. Despite the large size, and despite the presence of a pair of
submedian prothoracic setae, this species may belong to the dispersus-group, with many large notched pores
on the dorsal disc, extremely small compound pores on abdominal segments VII and VIII (reminiscent of the
single tiny pair on segment VII of flavus), and with the cephalothoracic submedian setae very much smaller
than remainder of dorsal setae.
Aleurodicus talamancensis Martin

(Figs 19, 20, 34–36, 132)
Aleurodicus talamancensis Martin, 2005: 3. Holotype puparium, Costa Rica [BMNH, examined].
Redescription, based on reappraisal of species limits

As originally defined, this species included material that is here described as a distinct species, A. chirri-
poensis (p. 23). The descriptions of A. chirripoensis and another new species, A. darienpalmae (p. 26), are
based on comparison with A. talamancensis and, accordingly, the description of A. talamancensis is repro-
duced here, slightly modified to include data not available at the time of original description.
PUPARIUM. Margin. Outline broadly ovoid, 1.05–1.25 mm long, 0.75–0.91 mm wide, generally widest
at abdominal segment II/III (n=25), but specimens of only 0.90 mm long have been examined subsequent to
original description. Margin usually apparently irregular as a result of protuberant submarginal pores viewed
in profile, sometimes broadly lobulate when completely flattened, with 4–7 shallow lobules per 0.1 mm of
abdominal margin. Dorsum. Cuticle often brownish and distinctly paler in submedian area and inner subdor-
sum (Figs 19, 132). Longitudinal moulting suture reaching puparial margin; transverse moulting sutures dis-
tally curving abruptly anteriad, becoming almost tangential to puparial margin and terminating opposite
middle legs (Fig. 35). Cuticle of submedian area somewhat corrugate, becoming subtly reticulate subdorsally
and then more rugose towards submargin. Abdominal segmentation marked by folds most of which are
suture-like submedially but still evident in subdorsum; segment VI/VII boundary sometimes medially only
marked by presence of pockets, and not by any fold; median length of segment VII extremely reduced, only 7
segments discernible medially. Submedian abdominal depressions narrow and elongate, usually little more
than widenings of intersegmental divisions. Cephalothoracic segmentation only subtly marked. Abdominal
segments VII and VIII each bearing an ovoid raised area (Fig. 34) that is lateral to vasiform orifice on segment
VII and posterolateral to vasiform orifice on segment VIII; raised areas on segment VIII forming caudal fur-
row, within which lies excluded part of lingula. Vasiform orifice (Figs 34) broadly cordate, a little wider than
long, its rim smooth laterally, situated about its own length from posterior margin of puparium; operculum
wider than long, laterally rounded, its anterior edge straight but posterior edge distinctly emarginate where it

overlies lingula, opercular surface somewhat rugose-punctate, apparently without a posterior pair of setae; lin-
gula head spinules loosely clustered, rather than covering surface evenly; lingula protruding beyond vasiform
orifice, almost as wide as operculum at point of its emergence from under operculum, apically rounded, later-
ally emarginate in type series (but less so in coconut-feeding material from Ecuador – see Material Exam-
ined), its two pairs of setae situated close to apex, distal setal pair stouter and longer than proximal pair,
lingular apex closely approaching puparial margin. Chaetotaxy. Posterior marginal setae, 12 pairs of outer
submarginal setae (including nominal caudal pair), and single pairs of submedian pro-, meso- and metatho-
racic setae present, each long and hair-like; eighth abdominal setae present but much shorter and finer, situ-
ated anterior to vasiform orifice (Fig. 34). Pores. Cephalic pair and anterior 4 pairs of abdominal compound
pores (situated on segments III–VI) presenting laterally to viewer, 35 μ maximum diameter in holotype, each
with an axial process protruding beyond pore rim, apically acute when not damaged; a pair of much smaller
compound pores, about 16–20 μ in maximum diameter, present on each of abdominal segments VII & VIII,
each located on outer edge of its segment’s raised area (Fig. 34). Narrow submargin defined by band of
crowded, wide-rimmed pores that stand proud from puparial surface, inner boundary of zone confluent with
row of submarginal setal bases, pore band not interrupted at posterior extremity of puparium but often diffi-
cult to see there because of marginal down-curling; when margin completely flattened on slide, extreme outer
submargin can be seen to be devoid of pores (Fig. 36). Dorsal disc with scattered bright pores, most slightly
larger than those of submarginal band, usually 2–3 present in vicinity of each large compound pore, and 3–6
on each raised area of abdominal segments VII & VIII (Fig. 34). Venter. Ventral characters typical for Aleuro-
dicus.
MATERIAL EXAMINED. Type material from Musa as detailed in description, Costa Rica (BMNH,
InBIO, USNM); non-type material – several samples, various dates and collectors: Colombia, on Anthurium
sp. (USNM); Costa Rica, on Cocos nucifera, Ficus sp. and ?Maxillaria sp. (BMNH, USNM); Ecuador, on
Cocos nucifera and unidentified orchids (BMNH, USNM); Nicaragua, on palm (USNM); Panamá, on Cocos
nucifera, cultivated variety of Musa and an unidentified palm (BMNH, USNM).
DISTRIBUTION. Neotropical Region – Colombia, Costa Rica, Ecuador, Nicaragua, Panamá (including
Canal Zone).
COMMENTS. In 2005, severe damage to banana plantations in Costa Rica was reported, caused by enor-
mous populations of this species of Aleurodicus. A. talamancensis was described by Martin (2005), based
upon extensive material from bananas, and one sample from ?Maxillaria sp., all the material from Costa Rica.
Collecting by the author in Ecuador, and material kindly loaned to the author for the present study (courtesy of
USNM), has revealed more material of A. talamancensis from several, mostly monocotyledonous, hosts and
from other countries (see above). The additional material has indicated that the sample from ?Maxillaria is a
distinct species, here described (p. 23) as A. chirripoensis. Two BMNH specimens from a palm in Darién
Province, Panamá, are clearly closely related to A. talamancensis, although the mesal boundary of the sub-
marginal wide-rimmed pore band is interdigitated with the submarginal setal bases, and these two puparia
(and one third-instar nymph) are here described as A. darienpalmae (p. 26).
Aleurodicus trinidadensis Quaintance & Baker

(Fig. 56)
Aleurodicus trinidadensis Quaintance & Baker, 1913: 69. Lectotype puparium here designated, Trinidad [USNM].
DISTRIBUTION. Neotropical Region – Ecuador, Guyana, Trinidad.

MATERIAL EXAMINED. Type material detailed and discussed below (USNM); 1 puparium, Ecuador,
Orellana Province, Yasuní N.P., Napo Wildlife Center, on unidentified host, 18.ii.2005 (Martin) (BMNH); 4

puparia, Guyana, Kaieteur Falls, on Clusia sp. and unidentified hosts, vi.2006 (Martin) (BMNH). 1 puparium,
Trinidad, Erin, on Musa sp., 22.i.1919 (Williams) (BMNH); 1 puparium, Trinidad, Matelot, on Cocos
nucifera, 16.x.1996 (International Institute of Entomology #23532) (BMNH).
COMMENTS. Three slides of original material were loaned to the author by USNM: one comprises a few
damaged adults, and a single puparium in poor condition; another contains a single uncleared third-instar
nymph; the third (evidently made at a later date than the other two) contains a single post-emergence pupal
case in good condition. The data on these slides are “Aleurodicus trinidadensis Q & B, TYPE, on coconut,
F.W.Urich, March 27, 1912, Q 8066”, with “type no. 14777, USNM” on one red label on each of the two older
slides. Also lent to the author was an envelope with residual dry material from the original sample, containing
one further puparium which has now also been slide-mounted. The slide preparation made from the dry spec-
imen is the most complete specimen of all, and is here designated the lectotype: the slide bears the full data
on newly-printed labels, along with a small, printed, red label from the dry-collection envelope, bearing the
data “Type no. 14777, U.S.N.M.”.
A. trinidadensis is a member of the capiangae / dugesii species-group, bearing closest resemblance to A.
vinculus (seq.). As discussed in the comments on A. capiangae (p. 21), the number of double-rimmed septate
pores in the submarginal row varies significantly within the species-group, and the species limits may require
revision in the light of more detailed studies.
Aleurodicus vinculus Martin

(Figs 11, 30, 31)
Aleurodicus vinculus Martin, 2004: 30. Holotype puparium, Belize [BMNH, examined.]
DISTRIBUTION. Neotropical Region – Belize.

MATERIAL EXAMINED. Type material as detailed in description, Belize (BMNH).
COMMENTS. Only a single sample comprising three puparia, and a further single specimen, are known,
and these were initially tentatively identified as a variant of A. trinidadensis Quaintance & Baker. In compari-
son with study material of A. trinidadensis from Trinidad, the Belizean specimens have a much greater num-
ber of small septate pores on the dorsal disc, these arranged in distinct chain-like rows across the submedian
areas of the cephalothorax and abdominal segments II-VIII; on the meso- and metathorax, there are two such
chains of pores (Fig. 30). Without more study samples, the degree of variation in A. trinidadensis is uncertain,
but A. vinculus is here considered to be distinct, defined by these chains of septate pores, and the species is a
member of the capiangae / dugesii species-group.
PALAEALEURODICUS gen. nov.

Type species: Aleurodes holmesii Maskell, 1896: 435
DIAGNOSIS. Characters of most diagnostic importance are given in bold. As interpreted here, Palaealeurod-
icus comprises species with the following combination of characters.
PUPARIA (Figs 76-78, 80-83, 86, 136, 137) – submedian cephalothoracic, 8th abdominal and anterior
marginal setae, absent; submargin and/or dorsal disc usually with no evident simple pores, or with a
few of one type only; cephalic pair and 4–6 abdominal pairs of large subdorsal compound pores present, most
or all with a robust, acute, axial process even if this does not extend beyond pore mouth; cephalic and at
least 2 abdominal pairs with their axial processes dagger-like, extending well beyond the pore mouth,
and even beyond the puparial margin; anteriormost abdominal compound pore pair located on segment III

(but lateral to median part of segment II, as seen in Aleurodicus, see Fig. A), or on segment IV; two pairs of
cicatrices present on thoracic area (scars of third-instar compound pores); lingula always protrudes beyond
vasiform orifice, its four subapical setae on the protruding part; ventrally, each leg rounded apically, with-
out an apical claw, antennae very short and close to fore legs (Fig. 86); margin usually with 3–5 pro-
nounced teeth per 0.1 mm of margin; usually only 0–2 pairs of submarginal setae present (including the
nominal caudal pair).
THIRD-INSTAR NYMPHS (Fig. 84) – with same chaetotaxy as its associated puparia, or with an
increased number of setae; with 2 pairs of compound pores, thoracically situated; ventrally, legs and antennae
smaller than in puparia, sometimes legs inevident. Third-instar with 2 pairs of cicatrices, on cephalus and
last abdominal segment (Fig. 84); second-instar with 2 pairs of compound pores in same positions.
ADULTS (when known) (Figs 79, 85, 87–89, 98, 99) – with 7-segmented antennae; forewing vein R
forked (Figs 87, 98); antennae of males with many prominent sensoria that give the flagellar segments a
“knobbly” appearance (Fig. 99); abdominal wax plates numbering 4 pairs in males, and in females of cin-
namomi, holmesii and machili, but 5 pairs in females of borneensis (Fig. 85).
ETYMOLOGY. The chosen name is designed to indicate the palaeotropical domicile of the included spe-
cies, along with inclusion of this genus within the Aleurodicinae.
COMMENTS. The suite of characters given in bold, above, serves to define this new genus and to distin-
guish it from Aleurodicus, in which genus all previously described species were placed, and from Aleuroctar-
thrus gen. nov. whose sole species was also previously placed in Aleurodicus (see pp. 51, 52). Of particular
note are the absence of puparial leg claws (whose presence is normally diagnostic for the Aleurodicinae), very
short puparial antennae, almost complete absence of dorsal setae (there is one exception to this), and almost
complete absence of simple pores in the puparia. However, the presence of compound pores and four lingular
setae reliably indicate inclusion in the Aleurodicinae. The presence of five pairs of abdominal wax plates in
the females of P. borneensis (q.v.) is a character apparently unique to that species: females of two other Palae-
aleurodicus species are known at present, and they possess the more usual four pairs.
The species of Palaealeurodicus are poorly known, both taxonomically and biologically. A particular
characteristic of the genus is the tendency for the puparial dorsum to become significantly raised as the adult
develops inside, with the puparial sides covered by a waxy pallisade (Figs 136, 137). Probably P. machili has
been the most frequently collected, being very common on species of Machilus and some other lauraceous
hosts in Hong Kong and Taiwan, and also being known from India. P. holmesii has the widest known distribu-
tion, ranging from India to Australia and Fiji, but with no indication of whether it is native throughout its
whole range (probably unlikely) or, if not, from whence it originated.
Key to Palaealeurodicus species – puparia
1. Margin smooth; with 12 pairs of submarginal setae; penultimate pair of compound pores (abdominal seg-
ment VII) ring-shaped, without axial processes (Fig. 81) ...................................... indicus (Regu & David)
- Margin dentate (Figs 76–78, 80, 82, 83); without, or with only 1–3 (posterior abdominal) pairs of submar-
ginal setae; every compound pore with an acute axial process, even if not extending beyond pore mouth 2
2. With 4 pairs of abdominal compound pores only (Fig. 76). ...................................... antidesmae (Corbett)
- With 5 or 6 pairs of abdominal compound pores..........................................................................................3
3. With 5 pairs of abdominal compound pores (Figs 77, 80, 83) .....................................................................4
- With 6 pairs of abdominal compound pores (Figs 78, 82) ...........................................................................6
4. All compound pores similar in form and size (Fig. 83); abdominal segmentation very pronounced into
outer subdorsum............................................................................................................. borneensis sp. nov.
- Abdominal compound pores of two distinctly different sizes (Figs 77, 80); abdominal segmentation less

pronounced ...................................................................................................................................................5
5. With anteriormost 1 pair of abdominal compound pores much smaller than cephalic pair and other 4
abdominal pairs (Fig. 80).............................................................................................. machili (Takahashi)
- With anteriormost 3 pairs of abdominal compound pores much smaller than cephalic pair and posterior-
most 2 pairs (Fig. 77).............................................................................................. cinnamomi (Takahashi)
6. Marginal teeth fully separate, their sides not contiguous, margin itself coarsely and regularly dentate (Fig.
78) ................................................................................................................................... holmesii (Maskell)
- Marginal teeth with their sides contiguous, margin itself with shallow and rather irregular crenulations
(Fig. 82) .......................................................................................................................... wallaceus (Martin)
Palaealeurodicus antidesmae (Corbett) comb. nov.

(Fig. 76)
Aleurodicus antidesmae Corbett, 1926: 267. Holotype puparium, Sri Lanka [presumed lost].
DISTRIBUTION. Oriental Region – Sri Lanka.

MATERIAL EXAMINED. 1 first-instar nymph, Ceylon, Pundaluoya, on Antidesma bunius (Euphorbi-
aceae) (E. E. Green) (BMNH).
COMMENTS. This species was described from a solitary specimen but this holotype, collected by E. E.
Green and illustrated by Corbett (Fig. 76), has not been present in the BMNH collection for at least 30 years
(author’s personal observation). An examination of residual leaf material from the original sample, in the
BMNH dry collection, resulted in the discovery of a first-instar crawler which almost certainly belongs to the
same species and is now the sole known specimen. As described, with only four abdominal pairs of compound
pores, puparia of P. antidesmae should be immediately recognisable if collected again.
Palaealeurodicus borneensis sp. nov.

(Figs 79, 83–89, 136, 137)
PUPARIUM. Habitus. Immature stages feeding on leaf petioles and under leaf midribs. Maturing puparia
(with developing adults inside) become tall, with sides obscured (probably protected) by waxy pallisade (Figs
136, 137). Dorsum with glassy wax carapace (Fig. 136) that readily detaches from dried puparia and can be
slide-mounted directly into balsam. Adjacent to puparia, on leaf surface, can be seen fine pieces of wax fila-
ment, but the mature specimens of the type sample have no sign of filaments emerging from compound pores:
it is likely that the earlier (feeding) puparial stage would have these filaments visible, emerging from the com-
pound pores. Species strongly size-dimorphic, with male puparia much smaller than those of females. Margin.
Outline 1.15–1.21 mm long, 0.60–0.67 mm wide (female, n=14), 0.83–0.90 mm long, 0.45–0.48 mm wide
(male, n=14, Fig. 83), generally widest at abdominal segment III/IV. Marginal teeth very well-developed,
rather rounded-castellate when in perfect lateral view, 3.5–4 per 0.1 mm on lateral parts of thorax and abdo-
men in female, 4.5–5 per 0.1 mm in male. Dorsum. Cuticle brown, slightly mottled, and rather roughened by
granular markings and corrugations, especially submarginally. Outer submargin with wavy folding that gives
illusion of margin having double row of teeth (Fig. 83). Longitudinal moulting suture reaching puparial mar-
gin; transverse moulting sutures extending over half-way towards margin, often indicated almost to margin.
Abdomen with most segmental divisions well marked into submargin but with segments I and II only apparent
submedially and separated by a short fold only; segment II/III boundary marked by a “W”-shaped fold, and
remaining segmentation similarly indicated, with segment VII not reduced in length medially. Meso-/metatho-

racic boundary marked by transverse fold but other cephalothoracic segmentation only subtly indicated. A
pair of cephalic markings, anteromesal to compound pores, visible in some specimens (Fig. 83), possibly ves-
tigial “eyespots”. Vasiform orifice rounded-cordate; operculum transverse-ovoid, surface marked similarly to
lingula, its posterior margin very slightly convex and with pair of fine setae visible only sometimes; lingula
head tongue-shaped, slightly darker than rest of puparium, densely covered by fine dark blunt spinules, bear-
ing the normal 4 stout setae and with half its length extending beyond vasiform orifice. Chaetotaxy. A pair of
posterior marginal setae present, long and fine, up to 75μ in female puparia, a little shorter in males. Dorsum
entirely without setae except for 2 outer submarginal pairs, close to posterior end of puparium, similar to pos-
terior marginal pair (Fig. 83). Pores. Cephalic pair and 5 abdominal pairs of compound pores, all similar to
each other in size and form, up to 40µ in outer diameter (female) or 30μ (male), abdominal pairs situated on
segments IV–VIII. Axial processes of large compound pores long and sword-like, each with more than half its
length extending beyond pore mouth, longest pair (almost always on abdominal segment VIII) 0.13–0.155
mm in females or 0.09–0.115 mm in males. No simple pores discernible on the dorsum. Venter. Ventral
abdominal setae long and fine, underlying vasiform orifice, often longer than dorsal and posterior marginal
setae. Legs with two segments faintly indicated, almost straight-sided, smooth, apically rounded and without
apical claws (Fig. 86). Antennae very short, similar to legs but pointed apically and with no segmentation
(Fig. 86). Tracheal folds absent. 4 pairs of spiracles readily visible.
THIRD-INSTAR NYMPH (Fig. 84). Same brownish pigmentation as puparium. Ovoid, 0.5–0.64 mm
long (n=8). Margin smooth to slightly irregular. Chaetotaxy as in puparium. 2 pairs of compound pores situ-
ated on thorax, similar to those in puparia. 2 pairs of cicatrices present (scars of compound pores in second-
instar nymph), one pair cephalically and the other pair to either side of lingula. Ventral abdominal setal pair
and spiracles readily discernible; legs not evident at all, but antennae visible as tiny triangular points on two
specimens.
SECOND-INSTAR NYMPH. Brown-pigmented, ovoid, 0.40 X 0.21 mm (n=1). Margin smooth. Chaetot-
axy as in puparium. 2 pairs of compound pores present, 16μ in diameter, one pair cephalically, the other pair
on either side of vasiform orifice. Cicatrices absent. Vasiform orifice trapezoidal, operculum transversely
elliptical, lingula with 4 hairs discernible. Abdominal segmentation well marked medially. Ventral characters
mostly obscured by fungal mycelium but ventral abdominal setal pair present.
FIRST-INSTAR NYMPH. Cuticle slightly pigmented, outline ovoid, 0.28 X 0.17 mm (n=1). Margin
smooth. Chaetotaxy: single pairs of anterior and posterior marginal setae present; with a submarginal row of
setae, 18 on one side and 17 on the other; with an apparent pair of setal bases some distance anterior to vasi-
form orifice; ventral abdominal setal pair discernible. Compound pores and cicatrices absent. Vasiform orifice
triangular, operculum trapezoidal and occupying over half of vasiform orifice, detail of lingula not readily dis-
cernible. Much of ventral surface apparently lost, with no indication of antennae or legs.
ADULTS. Wings evenly dusky brownish, entire surface punctuated by minute black dots, venation as
shown (Figs 87, 88); forewing (female, n=2) 1.80-1.85 mm long, 0.91-0.95 mm wide, R1 / Rs branch in about
basal one-third of wing; hind wing 1.47 mm long, 0.70 mm wide (n=1). Abdominal wax plates numbering 4
pairs in males, but 5 pairs in females, posteriormost 2 pairs apparently on same segment (Fig. 85); dorsal pos-
teriormost pair of female wax plates subcircular and punctuated by faintly-marked polygonal reticulations
(Fig. 85, expanded detail), other wax plates with more typical finely sculptured surfaces visible with X400
magnification. Female antennae 7-segmented, densely spinulose, with sparse setae, apical segment rather
swollen and with two large hemi-annular sensoria (Fig. 89). Head with 2 discreet groups of ommatidia on
each side (Fig. 79), effectively with 4 compound eyes, with a distinct ocellus contiguous with each dorsal
compound eye, and a spinulose swelling between each dorsal and ventral compound eye (Fig. 79). Tarsi with
2 claws, the paronychium apparently spatulate with a few short spines. Apical rostral segment about 0.2 mm
long.

MATERIAL EXAMINED. Holotype puparium (male), East Malaysia, SARAWAK, Gunung Mulu mas-
sif, Melinau Gorge, on unidentified broadleaf tree, 11.iii.1989 (J.H.Martin #5451) (BMNH). Paratypes: 28
puparia (male), 46 puparia (female), 8 third-instar nymphs, 1 second-instar nymph, 1 first-instar nymph, 9
adult females, 2 adult males, same data as holotype (BMNH, USNM); duplicate dry puparial material on
leaves, same data (BMNH).
ETYMOLOGY. The specific epithet reflects the known domicile of this species, the island of Borneo.
COMMENTS. P. borneensis is unusual in having been discovered with the colony feeding on woody
twigs of its tree host. The individual puparia had their dorsal surfaces significantly raised from the venters, the
sides being covered (probably supported) by waxy secretions (Fig. 136, 137). Third-instar nymphs are some-
what different from those of other Palaealeurodicus species, in the extreme degree of reduction in legs and
antennae, the legs not being discernible and the antennae reduced to mere triangular points – these characters
have not been illustrated because there is little to see! Adult females of P. borneensis are highly unusual in
having five pairs of abdominal wax plates (Fig. 85).
Palaealeurodicus cinnamomi (Takahashi) comb. nov.

(Fig. 77 )
Aleurodicus cinnamomi Takahashi, 1951: 1. Lectotype here designated, West Malaysia (BMNH).
DISTRIBUTION. Austro-oriental Region – Singapore, West Malaysia.

MATERIAL EXAMINED. Lectotype puparium (female), West Malaysia, Cameron Highlands, on Machi-
lus sp. (Lauraceae), 01.x.1944 (R. Takahashi) (BMNH, Brit. Mus. 1955-799); paralectotypes, 27 puparia
(female), 5 puparia (male), 5 dissected pre-emergence adult females, 3 dissected pre-emergence adult males,
same data as lectotype (BMNH, USNM). Other material: 4 parasitised puparia (formerly syntypes), Singapore
[“Malaya, Shonan”], no host data, 21.xii.1942 (R. Takahashi) (BMNH, Brit. Mus. 1955-799); 15 puparia, Sin-
gapore, Bukit Timah Forest Reserve, on Nothaphoebe umbelliflora (Lauraceae), 13.v.2000 (Martin) (BMNH).
COMMENTS. Takahashi (1951) specified both Singapore and Malaya as original localities for his syn-
type series, but two slides in BMNH (presented by the Director of the then Selangor Museum) are both
labelled “Malaya”. However, one also has the collecting locality “Shonan”, which is the name which the Jap-
anese (who were occupiers of the region from 1942-1945) bestowed upon Singapore: the material was col-
lected in 1942. Singapore / Shonan was regarded as part of Malaya at that time. The Singapore slide is a cavity
slide containing four parasitised puparia only, with the parasitoid pupae having been removed from three of
the whitefly puparia and mounted separately under the same coverslip: this slide has not warranted remount-
ing.
The second syntypic slide in BMNH contained over 30 specimens (detailed above), which had received
little preparation prior to mounting. As part of this study, this material has been macerated, de-waxed, stained
and remounted onto seven new slides, one slide bearing the original labels. The determination “Aleurodicus
cinnamomi Takahashi” over-lay an earlier label “Aleurodicus machili Takahashi”, which has now been left
exposed for historical reasons – Takahashi had apparently initially thought it to be A. machili which he him-
self had described in 1931. The remainder of the data from this second original slide are “Machilus, Cameron
Highlands, 1.x.1944, R. Takahashi, Brit. Mus. 1955-799”. The lectotype is one of four female puparia on the
slide bearing the original labels, and is clearly indicated; one of the new paralectotype slide preparations has
been presented to USNM.

Palaealeurodicus holmesii (Maskell) comb. nov.
(Figs 78, 98, 99)
Aleurodes holmesii Maskell, 1896: 435. Lectotype puparium, Fiji [designated by Martin, 1999: 19, NZAC, examined].
Aleurodicus holmesii (Maskell) Cockerell, 1903: 664.
Aleurodicus malayensis Takahashi, 1951: 2. Lectotype puparium, West Malaysia [designated by Martin, 1999: 19,
BMNH, examined]. Synonymised by Martin, 1985: 312.
DISTRIBUTION. Oriental Region – India, Sri Lanka, Thailand; Austro-oriental Region – Java, New Guinea,
West Malaysia; Australia – Queensland, [New South Wales]; Pacific Region – Fiji.
MATERIAL EXAMINED (all BMNH). 19 paralectotype puparia of Aleurodes holmesii, Fiji, on Psidium
sp. (Myrtaceae) (ex-Maskell dry collection 1984); lectotype puparium and 7 paralectotype puparia of Aleuro-
dicus malayensis, Malaya, Kuala Lumpur, on undetermined tree, 10.iii.1943 (Takahashi); 1 puparium, “Cey-
lon”, Kandy, 02.i.1910 (Green); 4 puparia, “Siam”, Pah Meeung Mountains, 18.i.1928 (Cockerell); 5 mounted
puparia + duplicate dry material, Java, 1911 (Hargreaves); all nymphal stages, adult males, Australia, Queen-
sland, beach-top forest 2km north of Cape Tribulation, on ?Lauraceae, 13.xii.1996 (Martin); 10 puparia, 2
adult females, Papua New Guinea, Morobe Province coast, Buso, on undetermined host and Guioa sp. (Sapin-
daceae), ix-x.1979 (Martin).
COMMENTS. P. holmesii is the most widely distributed of the Palaealeurodicus species, with voucher
material in BMNH from the Malay Peninsula in the west to Fiji in the east. The species is regarded as being
variable in the length of its compound pore axial processes (which was the main character used to separate
malayensis from holmesii), and it differs from P. wallaceus mainly in the characteristics of the marginal teeth
(see key, p. 47).
Palaeleurodicus indicus (Regu & David) comb. nov.

(Fig. 81)
Aleurodicus indicus Regu & David, 1992: 100. Holotype puparium, India (Tamil Nadu) [depository not stated, not exam-
ined].
DISTRIBUTION. Oriental Region – India; Austro-oriental Region – Brunei.

MATERIAL EXAMINED. 5 puparia, Brunei, Belait District, 20 km east of Tutong, on unidentified host,
23.iii.1989 (Martin) (BMNH).
COMMENTS. A field note, made when the sample from Brunei (see above) was collected, described the
puparial body as greenish and each puparium as secreting 10 long, white, waxy filaments; puparia were feed-
ing under main leaf veins (personal observations). Each puparium has 6 pairs of compound pores and it there-
fore seems possible that it is the single pair of very large pores (that do not possess axial processes, see below)
that also do not secrete waxy filaments in this species.
This curious species displays a combination of puparial characters that are typical for Palaealeurodicus
(lack of submedian dorsal disc setae, lack of dorsal simple pores, lack of leg claws, and most compound pores
bearing a robust dagger-like axial process). However, it also displays puparial characters that are unique
within Palaealeurodicus (one pair of abdominal compound pores large and ring-shaped and without an evi-
dent axial process, puparial margin not regularly dentate but somewhat irregular, submargin with 12 pairs of
setae). Thus, indicus possesses a mixture of characters that also suggest possible affinities with Aleuroctar-
thrus (q.v., p. 51, 52).

Palaealeurodicus machili (Takahashi) comb. nov.
(Fig. 80)
Aleurodicus machili Takahashi, 1931: 208. Syntypes, Taiwan [TARI, not examined].
Aleurodicus formosanus Takahashi, 1932: 7. Syntypes, Taiwan [TARI, not examined]. Synonymised by Mound & Hal-
sey, 1978: 234.
Aleurodicus philomenae David, 1987: 1247. Holotype puparium, India (Maharashtra) [depository not stated, not exam-
ined]. Synonymised by David & Selvakumaran, 1990: 139.
DISTRIBUTION. Oriental Region – Hong Kong, India, Taiwan.

MATERIAL EXAMINED (all BMNH). Puparia, third-instar nymphs and adult females from several sam-
ples, Hong Kong, on Litsea and Machilus species, 1990-2003 (Martin); 4 puparia, 3 adult males, 2 adult
females, Taiwan, 1986 & 2006 (Ko).
COMMENTS. This species is common on Machilus, Litsea and some other members of the Lauraceae in
Hong Kong and Taiwan, where the immature stages tend to develop against the lower midribs of leaves, and is
also recorded from India. This species will probably prove to be more widely distributed in the more temper-
ate parts of the Oriental Region.
The synonymy of formosanus with machili arose because Takahashi had described a third-instar nymph as
formosanus, mistakenly regarding it as a puparium of a distinct species (Mound & Halsey, 1978). The synon-
ymy of A. philomenae with A. machili was drawn to the attention of David & Selvakumaran by Louise Russell
(USNM), as they acknowledged in their short communication (1990).
Palaealeurodicus wallaceus (Martin) comb. nov.

(Fig. 82)
Aleurodicus wallaceus Martin, 1988: 59. Holotype puparium, Sulawesi [BMNH, examined].
DISTRIBUTION. Austro-oriental Region – Brunei, Sarawak, Sulawesi.

MATERIAL EXAMINED (all BMNH). Holotype puparium and 30 paratype puparia, Indonesia, Sulawesi
Utara, Danau Mooat, near Kotamobagu, on Persea americana (Lauraceae), 23.iii.1985 (Martin); puparia &
third-instar nymphs, Brunei, Belait District, Penanjong army base, on undetermined host, 03.iii.1989 (Mar-
tin); 1 puparium, East Malaysia, Sarawak, Gunung Mulu National Park, Long Pala base camp, on Annon-
aceae, 27.vi.1978 (Eastop).
COMMENTS. P. wallaceus was originally discovered feeding on an avocado tree in Sulawesi. Although
avocado is a neotropical plant introduced into Asia, this whitefly is immediately recognisable as a member of
the old-world group of species that were then accommodated in Aleurodicus. A single puparium was collected
in Sarawak as part of the Royal Geographical Society’s expedition to Gunung Mulu (1977-1978), and the
author has since also collected material in Brunei.
This species differs from P. holmesii principally in the characteristics of the marginal teeth, and may also
be distinguished from the other species of Palaealeurodicus by use of the key, pp. 46, 47.
ALEUROCTARTHRUS gen. nov.

Type species: Aleurodicus destructor Mackie, 1912: 142
DIAGNOSIS.
PUPARIA (Figs 90-93) – characters of most diagnostic importance are given in bold. As interpreted here,
Aleuroctarthrus is defined by the following combination of characters: margin almost smooth, or with very

shallow marginal crenulations (Fig. 90); 12 pairs of submarginal setae present (including the nominal
caudal pair); 4 pairs of tiny submedian cephalothoracic setae and pair of 8th abdominal setae, present;
anterior marginal setae absent; submargin and/or dorsal disc sparsely provided with tiny simple pores of
one type only (sometimes appearing loculate and sometimes appearing to have a minute central spine); ceph-
alic pair and 6 abdominal pairs of subdorsal compound pores present, each with an acute axial process (Fig.
92), but these processes often not evident (see discussion, below); all abdominal compound pores very
large and subequal in size, much larger than the cephalic pair; abdominal compound pore pairs located on
segments III-VIII (as in Aleurodicus, see Fig. A); two pairs of cicatrices present on thoracic area (scars of
third-instar compound pores); with only two large lingular setae (Fig. 91) (sometimes with two very small
additional setae visible under 400X magnification); lingular apex excluded beyond vasiform orifice. Ventrally,
each leg very large and with an apical claw (Fig. 90); antennae very long, reaching at least to middle legs;
underlying abdominal segment VII / VIII boundary (between operculum and fourth abdominal pair of
compound pores) is a pair of circular secretory glandular areas (Figs 90, 93), each comprising subcircu-
lar cells (often with secreted material somewhat obscuring the structure, and the diaphanous nature of the
ventral cuticle often leads to displacement). [2nd-instar with 3 pairs of compound pores, on cephalus, pro-
thorax and last abdominal segment; 3rd-instar with 3 pairs of cicatrices in same positions.]
ADULTS – with eight-segmented antennae; antennae of males not provided with numerous promi-
nent sensoria; abdominal wax plates numbering four in females and three in males, the posterior two
pairs in females with stout setae along their anterior edges. Wings clear, unpigmented.
ETYMOLOGY. The chosen name reflects the characteristic eight-segmented antennae.
COMMENTS. The suite of characters given in bold, above, serve to define this new genus and to distin-
guish it from Aleurodicus and Palaealeurodicus. The antennae of adults of both sexes comprising eight seg-
ments is a character thought to be unique amongst known whiteflies, and is a major factor in proposing a
separate genus. The presence of four abdominal wax plates in females, but only three in males, is a character
shared with Aleurodicus but distinguishes Aleuroctarthrus from Palaealeurodicus whose known males pos-
sess four pairs of abdominal wax plates. Of particular note in the puparium are the presence of leg claws, pres-
ence of 12 pairs of submarginal setae and presence of submedian cephalothoracic setae (characters absent in
Palaealeurodicus). The reduction of puparial lingular setae to only one stout pair distinguishes this genus
from both Aleurodicus and Palaealeurodicus, but this reduction in lingular setae is seen in some other Old
World aleurodicine genera (Pseudosynaleurodicus Gillespie, Stenaleyrodes Takahashi, Synaleurodicus
Solomon). The dearth of dorsal simple pores distinguishes Aleuroctarthrus from almost all Aleurodicus spe-
cies.
The decision to propose a new genus to accommodate only a single species was taken reluctantly. How-
ever, it has long been clear that Mackie’s species destructor possesses characters that set it apart from the New
World assemblage of Aleurodicus species, but which it does not share with any of the other palaeotropical
aleurodicine taxa.
One further species, Palaealeurodicus indicus (Regu & David), has proved difficult to place generically,
especially in the absence of adults and earlier-instar nymphs, because it shares some puparial characters with
A. destructor (12 pairs of submarginal setae, margin smooth-to-irregular, ring-shaped compound pores, see
figure 81), that are not shared by the other species of Palaealeurodicus. It is hoped that adults and 2nd & 3rd-
instar nymphs of indicus might eventually provide some clarification, but indicus (see p. 50) is here placed in
Palaealeurodicus based on other characters.

Aleuroctarthrus destructor (Mackie) comb. nov.
(Figs 90–93)
Aleurodicus destructor Mackie, 1912: 142. Neotype, Philippines [designated by Martin, 1999: 17, USNM, examined].
Aleurodes albofloccosa Froggatt, 1918: 436. Lectotype puparium, Australia [designated by Martin, 1999: 17, USNM,
examined]. Synonymised by Dumbleton, 1956: 160.
DISTRIBUTION. Oriental Region – Thailand; Austro-oriental Region – Brunei, West Malaysia, New Britain,
New Guinea, Philippines, Sarawak, Solomon Is, Sulawesi; Australia - NT, Qld, NSW, Vic, WA.
MATERIAL EXAMINED. Material from all countries listed above (BMNH).
COMMENTS. As the only included species, the generic diagnosis given above serves to provide all the
salient characters for A. destructor. The presence in puparia of six pairs of very large ring-shaped abdominal
compound pores, which most often appear to be devoid of axial processes, combined with only a single pair
of stout lingular setae, renders A. destructor immediately recognisable. The presence of eight-segmented
antennae in adults is apparently unique to A. destructor, but see discussion of Palaealeurodicus indicus, above
and on p. 50.
Despite its name, A. destructor seems only rarely to cause problems for agriculturalists, and it has not
been submitted to BMNH for identification for over 20 years. A single puparium, sent to BMNH from Liver-
pool Garden Festival in 1984, and stated to have been collected on Olea from Spain, is regarded as having
erroneous data.
Revised generic placements
Bakerius marmoratus (Hempel) comb. nov.
Aleurodicus marmoratus Hempel, 1922b: 1124 (Portuguese), 1161 (English). Syntypes (adults), Brazil [MZUSP, exam-
ined].
DISTRIBUTION. Neotropical Region – Brazil.

MATERIAL EXAMINED. Adult syntypes – see details and discussion below.
COMMENTS. In connection with an earlier study (Martin, 2004) the author was sent a syntypic slide of
A. marmoratus on loan, through the kindness of MZUSP. This slide comprised only adult specimens, but no
puparia could be located (Sonia Casari, MZUSP, personal communication). With our currently limited under-
standing of the taxonomy of adult whiteflies, puparia would be necessary for any definitive assessment of this
species. However, some conclusions can be drawn from the original description, as discussed below.
Hempel’s description of the “larva” of marmoratus details three pairs of large circular compound glands
[compound pores], one pair cephalically and two pairs abdominally, which strongly suggests a puparium of
Bakerius (Fig. 115) or Leonardius [third-instar nymphs of Aleurodicus species possess just two pairs of com-
pound pores]. The “pupa” [puparium] is described as having the median area distinctly darker than the periph-
ery, another typical character for Bakerius puparia. The description continues to detail six (not six pairs of)
white wax tubes [filaments] produced by the compound pores. Hempel also stated that compound pores were
not present in “very young” nymphs, whereas they are present in 2nd and 3rd nymphal instars of Aleurodicus
species. However, nymphal compound pores are not present in Bakerius species.
It is therefore concluded that marmoratus is best placed in Bakerius (comb. nov.), in the absence of any
descriptive suggestion of the additional presence of agglomerate pores, that would then indicate inclusion in
Leonardius instead.

Metaleurodicus essigi (Sampson & Drews) comb. nov.
(Figs 73–75)
Aleurodicus essigi Sampson & Drews, 1941: 147. Lectotype puparium, here designated, Mexico (UCD).
Redescription, from lectotype specimen

PUPARIUM. Habitus. Original description said “body covered by coating of mealy wax and four pairs of
shiny wax filaments which arise from the dorsum [from the large abdominal compound pores], and are many
times the length of the body and make the insect very conspicuous”. Margin. Outline oval, 0.8 mm long, 0.5
mm wide, widest at abdominal segment IV (n=1). Margin irregular, not toothed. Dorsum. Longitudinal
moulting suture reaching puparial margin; transverse moulting sutures damaged (Fig. 75). Abdominal seg-
ment VII not significantly reduced medially. Abdominal segmentation distinct, especially between segments
III/IV to VI/VII, where the division is visible almost to puparial margin (Fig. 75). Pro-/mesothoracic and
meso-/metathoracic divisions subtly visible submedially. Two pairs of thoracic cicatrices present (scars of
third-instar compound pores, figure 73). Vasiform orifice (Fig. 74) rounded-cordate; operculum trapezoidal,
its posterior margin very shallowly “m”-shaped, bearing a pair of stout setae; lingula head tongue-shaped,
densely covered by seta-like spinules, bearing the normal 4 stout setae; lingula not reaching posterior puparial
margin, but its setae overlapping margin. Chaetotaxy. A probable anterior marginal seta visible on one
side. A pair of posterior marginal setae (56μ long), and a pair of eighth abdominal setae (8μ long, anterior to
vasiform orifice) present; submargin with 12 pairs of setae (including the nominal caudal pair), ranging from
15μ (anteriorly) to 30μ (the posteriormost pair); submedian cephalothoracic setae entirely absent (Fig. 73).
Pores. Cephalic and posterior two pairs of compound pores 22μ in outer diameter, rather “bell-shaped” (Fig.
74). Abdominal compound pores on segments III–VI extremely large, 60–80μ in outer diameter, each with a
sword-like axial process. Immediately within puparial margin, on the abdomen only, lies a single row of wide-
rimmed pores about 9μ in diameter (Fig. 74), about 16 on one side of body and 18 on other (damage makes
accurate count impossible); on the cephalothoracic plates pores of this type are absent (Fig. 73). Dorsal disc
sparsely punctutated by loculate pores and unassociated porettes, with a very few (slightly larger) loculate
pores situated near puparial margin on cephalothoracic plates. Venter. Ventral abdominal setae not discernible
in lectotype. Legs typically robust and two-segmented, smooth, each with a pronounced apical claw. Antennal
apices extending to bases of fore legs, their own bases anterolateral to fore legs. Tracheal folds absent.
DISTRIBUTION. Neotropical Region – Mexico.
MATERIAL EXAMINED. Lectotype puparium here designated, MEXICO, on cliffs at Acapulco, Guer-
rero, on shrub, iii.1926 (G.F.Ferris, #347) (UCD 850).
COMMENTS. The absence of submedian cephalothoracic setal pairs, combined with the presence of loc-
ulate simple pores across the dorsum precludes inclusion in Aleurodicus, within the definition of Martin
(2004) and adopted here. Indeed, this species only has dorsal simple pores of two types (loculate and porette),
whereas the majority of Aleurodicus species have at least three types of simple pores, combined with a much
greater pore density in at least some areas of the dorsal surface. Whilst the four pairs of very large compound
pores are certainly atypical for Metaleurodicus, they are not typical for Aleurodicus, either. If anterior mar-
ginal setae are indeed present then that character also suggests exclusion from Aleurodicus. It is felt that essigi
is most suitably accommodated within Metaleurodicus (comb. nov.).
Only a single specimen is known, but the description did not state this. Thus, the sole known specimen
must be regarded as a syntype: it is therefore formally here designated the lectotype for future stability.

Acknowledgements
This study has benefitted enormously from the kind efforts of several colleagues who have arranged loans of
study material, and checked records and data on behalf of the author. In particular, I would like to voice my
thanks to Debra Creel (USNM), who organised the loan of substantial amounts of material and who also
investigated the background to several type-material slides which bore incomplete data. Gillian Watson and
Ray Gill (CDFA) assisted with the loan of material, and also checked California records. Steve Heydon (R.M.
Bohart Museum, UCD) very kindly arranged for the author to borrow the sole known specimen of Aleurodi-
cus essigi Sampson & Drews. Penny Gullan (UCD) kindly offered opinions on several taxonomic matters, and
it is a pleasure to acknowledge her support. Field and logistical support by Elroy Charles, during field sam-
pling in Guyana in 2006, was of great value and is acknowledged with thanks. The author is indebted to the
following colleagues for their agreement to use their original photographic images here: Luis Valencia V (fig-
ures 124 and 125), and David Haviland (figure 131).
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FIGURES 1–4. Aleurodicus spp, puparia, photomicrographs. 1, A. araujoi Sampson & Drews. 2, A. coccolobae Quain-
tance & Baker. 3, A. dispersus Russell. 4, A. dugesii Cockerell.

FIGURES 5–8. Aleurodicus spp, puparia, photomicrographs. 5, A. inversus Martin. 6, A. magnificus Costa Lima. 7, A.
maritimus Hempel. 8, A. niveus Martin.

FIGURES 9–12 Aleurodicus spp, puparia, photomicrographs. 9, A. pauciporus Martin. 10, A. pulvinatus (Maskell). 11,
A. vinculus Martin. 12, A. rugioperculatus Martin.

FIGURES 13–16 Aleurodicus spp, puparia, photomicrographs. 13, A. coccolobae Quaintance & Baker, region of vasi-
form orifice. 14, A. rugioperculatus Martin, cephalothoracic sculpture. 15, A. rugioperculatus, median abdominal detail.
16, A. rugioperculatus, operculum and lingula.

FIGURES 17–20 Aleurodicus spp, puparia, photomicrographs. 17, 18 A. chirripoensis sp. nov., Costa Rica. 19, 20 A.
talamancensis Martin, Costa Rica.

FIGURES 21–26 Aleurodicus spp, puparia. 21, A. araujoi Sampson & Drews, posterior dorsal detail, with inset (a)
showing posterior marginal seta and deflexed margin. 22, A. dugesii Cockerell, posterior dorsal detail, with section (a)
showing posterior marginal seta and deflexed margin. 23, A. inversus Martin, anterior abdominal dorsal detail, with inset
(a) showing margin deflexed ventrally. 24, A. inversus, photomicrograph of posterior part of puparium. 25, A. magnificus
Costa Lima, posterior dorsal detail, with inset (a) showing margin deflexed ventrally (modified after Costa Lima, 1928).
26, A. maritimus Hempel, cephalothoracic dorsal detail.

FIGURES 27–33 Aleurodicus spp, puparia. 27, Aleurodicus pauciporus Martin, abdominal segments V–VIII. 28, A.
pauciporus, lateral margin and submarginal detail. 29, A. pulvinatus (Maskell), abdominal segment IV. 30, A. vinculus
Martin, submedian thoracic detail, showing “chains” of septate pores. 31, A. vinculus, cephalothoracic margin and sub-
marginal detail. 32, A. rugioperculatus Martin, section of cephalothorax, with longitudinal moulting suture on right. 33,
A. niveus Martin, posterior detail.

FIGURES 34–40 Aleurodicus spp, puparia. 34, A. talamancensis Martin, Costa Rica, posterior detail. 35, A. talaman-
censis, submarginal detail and apex of transverse moulting suture. 36, A. talamancensis, fully flattened margin at conflu-
ence with longitudinal moulting suture. 37, A. chirripoensis sp. nov., Costa Rica, posterior detail (to same scale as fig.
34). 38, A. chirripoensis, surface detail of lingula. 39, A. chirripoensis, abdominal segment IV with intersegmental divi-
sions and submedian depressions. 40, A. chirripoensis, cephalic compound pore and adjacent wide-rimmed simple pores,
with inset detail of submarginal pore band.

FIGURES 41–44 Aleurodicus spp, puparia. 41, A. clusiae sp. nov., Guyana, cephalothoracic plate. 42, A. clusiae, poste-
rior right-hand abdominal detail, with wedge-shaped inset showing detail of submarginal pore band. 43, A. pseudugesii
sp. nov., Ecuador, margin and submargin, showing small double-rimmed septate pores and narrow band of wide-rimmed
pores. 44, A. pseudugesii, posteror dorsal detail.

FIGURES 45–47 Aleurodicus spp, puparia. 45, A. flavus Hempel, Brazil, posterior dorsal detail. 46, A. charlesi sp.
nov., Guyana, posterior dorsal detail. 47, A. nicaraguensis sp. nov., Nicaragua, right-hand cephalothoracic detail with
cephalic compound pore and thoracic cicatrice, mesal boundary of band of submarginal wide-rimmed pores shown as
dashed line, and pores themslves shown in inset.

FIGURES 48–53 Aleurodicus spp, puparia. 48, A. septatissimus sp. nov., Peru, anterior dorsal submarginal detail. 49, A.
septatissimus, right-hand dorsal abdominal detail. 50, A. spectabilis sp. nov., Brazil, dorsal thoracic detail, showing cica-
trices. 51, A. spectabilis, posterior dorsal detail. 52, A. antillensis Dozier, Puerto Rico, posterior dorsal detail showing
small group of double-rimmed septate simple pores posterior to vasiform orifice, and its single pair of small compound
pores. 53, A. ornatus Cockerell, Jamaica, posterior dorsal detail, showing the single pair of small compound pores.

FIGURES 54–57 Aleurodicus spp, puparia. 54, A. etiennei sp. nov., French Guiana, dorsum. 55, A. capiangae Bondar,
Panamá, dorsum. 56, A. trinidadensis Quaintance & Baker, Trinidad, margin and dorsal submargin, abdominal segments
III & IV. 57, A. darienpalmae sp. nov., Panamá, posterior dorsal detail, with lingular spinule clusters shown as expanded
detail.

FIGURES 58–62 Aleurodicus spp, puparia. 58, A. magnificus, Belize, cephalic compound pore with notched simple
pores only posterior to it. 59, A. neglectus Quaintance & Baker, Guyana, cephalic margin and submargin, with notched
simple pores surrounding compound pore. 60, A. neglectus, detail of fully flattened margin and submargin. 61, A. jamai-
censis Cockerell, Jamaica, margin and dorsal submargin. 62, A. guppyii Quaintance & Baker, Trinidad, cephalic margin
and dorsal submargin.

FIGURES 63–66 Aleurodicus spp, puparia. 63, A. juleikae Bondar, lectotype, Brazil, dashed line marking mesal bound-
ary of submarginal wide-rimmed pore band. 64, A. juleikae, lectotype, detail of margin and dorsal submargin, abdominal
segments III/IV. 65, A. cocois (Curtis) [lectotype of A. anonae Morgan], Guyana, dashed line marking mesal boundary of
submarginal wide-rimmed pore band. 66, A. cocois [lectotype of A. anonae], abdominal margin and dorsal submargin.

FIGURE 67 Aleurodicus pulvinatus (Maskell), lectotype puparium, Trinidad, showing pigmentation to right and distri-
bution of simple pores to left: with (a) rows of modified 8-shaped simple pores and (b) cluster of tiny bright pores adja-
cent to compound pore.

FIGURE 68 Aleurodicus pulvinatus (Maskell), simplified puparial outlines, showing variation of cuticular pigmentation
in samples from (a) Costa Rica, Hura crepitans, (b) Belize, Lauraceae, (c) Belize, Guettarda combsii, (d) Belize, unde-
termined, (e) Panamá, Isertia hypoleuca. The dashed line marks the boundary of the submarginal pore band.

FIGURE 69 Aleurodicus floccissimus (Martin et al.) comb. nov., puparium, Tenerife, with arrow (a) indicating tiny 8th
abdominal compound pore almost underlying lingula.

FIGURE 70 A. mirabilis (Cockerell), puparium, Ecuador, posterior detail with deflexed submarginal detail shown bot-
tom-left, dorsal submarginal detail inset bottom-right, deflexed margin as dashed line, lingula & mesal boundary of sub-
marginal band of wide-rimmed pores as dotted lines.

FIGURE 71 Aleurodicus dispersus Russell, puparium, after Russell (1965), with inset detail of (a) modified 8-shaped
pore, (b) double-rimmed notched pore, (c) wide-rimmed pore, (d) minute wide-rimmed pore and (e) wide-rimmed
notched pore.

FIGURE 72 Aleurodicus decemensis sp. nov., holotype puparium, Panamá.

FIGURES 73–76. 73, Metaleurodicus essigi (Sampson & Drews) comb. nov., lectotype puparium, Mexico, cephalotho-
racic plate. 74, M. essigi, lectotype, posterior dorsal detail. 75, M. essigi, lectotype, photomicrograph. 76, Palaealeurodi-
cus antidesmae (Corbett) comb. nov., holotype puparium, Sri Lanka (after Corbett, 1926).

FIGURES 77–79 Palaealeurodicus spp. 77, P. cinnamomi (Takahashi) comb. nov., lectotype puparium, West Malaysia.
78, P. holmesii (Maskell), puparium, Queensland. 79, P. borneensis sp. nov., Sarawak, adult female, completely divided
compound eye.

FIGURES 80–82 Palaealeurodicus spp, puparia. 80, P. machili (Takahashi) comb. nov., Hong Kong. 81, P. indicus
(Regu & David) comb. nov., Brunei. 82, P. wallaceus (Martin) comb. nov., paratype, Sulawesi.

FIGURES 83–86 Palaealeurodicus borneensis sp. nov., Sarawak. 83, holotype puparium, dorsum. 84, third-instar
nymph, dorsum, with one pair of ventrally-displaced setae marked as dashed lines. 85, Adult female, abdomen in lateral
view to show wax plates. 86, puparium (male), ventral detail of legs and antenna.

FIGURES 87–89 Palaealeurodicus borneensis sp. nov., Sarawak, adult female. 87, forewing. 88, hind wing. 89, anten-
nal segment VII.

FIGURES 90–93 Aleuroctarthrus destructor (Mackie) comb. nov., puparium. 90, whole puparium, Guadalcanal. 91,
vasiform orifice, showing presence of extremely reduced second pair of lingular setae, Guadalcanal. 92, abdominal dor-
sal submarginal detail, showing faintly visible axial processes in compound pores, New Guinea. 93, posteroventral detail,
showing ventral abdominal setae, posterior spiracles and pair of cellular secretory glands, Queensland.

FIGURE 94 Azuraleurodicus pentarthrus Martin, puparium, with expanded detail of (a) marginal glands and submar-
ginal double-rimmed 8-shaped septate pores, (b) angular septate pore, and (c) submarginal double-rimmed concentric
pores.

FIGURES 95–99. 95, Eudialeurodicus sp., puparium, Ecuador. 96, Nipaleyrodes elongata Takahashi, puparium,
Sulawesi. 97, Metaleurodicus cardini (Back), puparium, Dominican Republic. 98, Palaealeurodicus holmesii (Maskell)
comb. nov., forewing (male), Queensland. 99, P. holmesii, antennal segment III (male), Queensland.

FIGURE 100 Paraleyrodes pseudonaranjae Martin, paratype puparium, Hong Kong.

FIGURE 101 Nealeurodicus petiolaris Martin, paratype puparium, with ventral detail at left and expanded detail of (a)
thoracic margin, (b) lateral margin and dorsal submargin, (c) abdominal compound pore and (d) dorsal submedian detail
of abdominal segments VII and VIII, showing clusters of wide-rimmed pores and quadriloculate pores.

FIGURES 102–104. 102, Synaleurodicus serratus Martin, puparium, Western Australia, with detail of (a) vasiform ori-
fice with seta-less lingula, (b) subdorsal agglomerate pore, (c) anterior margin and dorsal submargin, (d) anterior margin
and ventral submargin, (e) submedian abdominal glandular depressions, and (f) expanded detail of legs, antenna and spi-
racles. 103, S. hakeae Solomon, Western Australia, subdorsal agglomerate pore cluster with ducts (teneral specimen).
104, Paraleyrodes pseudonaranjae Martin, paratype adult forewing, Hong Kong.

FIGURES 105–109 Dialeurodicus spp, puparia. 105, D. bondariae Martin, vasiform orifice and surrounds. 106, D.
bondariae, margin and dorsal submarginal detail of second-anteriormost “ray”. 107, D. caballeroi Martin, margin and
dorsal detail of abdominal “ray” with porate tubercle at proximal end. 108, D. caballeroi, vasiform orifice and surrounds.
109, D. caballeroi, photomicrograph.

FIGURES 110–113. 110, Kaieteurodicus cereus Martin, puparium, Guyana. 111, Stenaleyrodes vinsoni Takahashi,
puparium, New Caledonia. 112, Pseudosynaleurodicus mayoi Gillespie, puparium, Queensland. 113, Ceraleurodicus
varus (Bondar), puparium, Belize, showing radial “rays”, asymmetric provision of compound pores, and chaetotaxy rep-
resented by dots.

FIGURES 114–116. 114, Octaleurodicus nitidus Hempel, puparium, Ecuador. 115, Bakerius ?phrygilanthi Bondar,
puparium, Panamá. 116, Leonardius kellyae Martin, paratype puparium, abdominal segments IV–VI, with agglomerate
porous areas on each segment, and that on segment IV surrounding the compound pore.

FIGURES 117–120 photomicrographs, puparia. 117, Aleuronudus acapulcensis (Sampson & Drews), Panamá. 118,
Aleuronudus manni (Baker), Belize. 119, Austroaleurodicus pigeanus (Baker & Moles), Chile. 120, Azuraleurodicus
pentarthrus Martin, Belize.

FIGURES 121–123 habitus photographs of Aleurodicus spp, puparia. 121, A. niveus Martin, type colony, Las Ceuvas,
Belize. 122, A. dispersus Russell, Santa Cruz de Tenerife, Canary Islands. 123, A. coccolobae Quaintance & Baker, Las
Ceuvas, Belize.

FIGURES 124–129 habitus photographs. 124, Aleurodicus ?juleikae Bondar developing, unusually, on fruits of mango,
Santiago de Surco, Peru (photograph © Luis Valencia V, with permission). 125, Aleurodicus ?juleikae, puparia and egg-
laying spiral, on Ficus, Santiago de Surco, Peru (photograph © Luis Valencia V, with permission). 126, Aleuroctarthrus
destructor (Mackie) comb. nov., tangle of coarse filaments secreted by puparia, Morobe Province coast, New Guinea.
127, Aleurodicus floccissimus (Martin et al.) comb. nov., secretions accumulated when puparia develop in sheltered con-
ditions, Yasuní National Park, Ecuador. 128, A. mirabilis (Cockerell) comb. nov., puparia developing along main leaf
veins and completely obscured by their secretions, Georgetown, Guyana. 129, Aleurodicus sp., adults, Esmeraldas, Ecua-
dor.

FIGURES 130–131 habitus photographs of Aleurodicus spp. 130, A. mirabilis (Cockerell), oviposition trails on Annona
leaf, La Independencia, Ecuador. 131, A. dugesii Cockerell, extensive waxy secretions lend a surreal appearance to vege-
tation, Bakersfield, California (photograph © David Haviland, with permission).

FIGURES 132–133 habitus photographs of Aleurodicus spp, puparia. 132, A. talamancensis Martin, Limón Province
banana plantations, Costa Rica. 133, A. chirripoensis Martin, part of type colony, Cerro Chirripó, Costa Rica.

FIGURES 134–135 habitus photographs of puparia. 134, Ceraleurodicus keris Martin, Bartola, Nicaragua, showing
extreme asymmetry of outline and compound pore provision. 135, Aleurodicus clusiae sp. nov., post-emergence pupal
cases, Kaieteur Falls, Guyana.

FIGURES 136–137 habitus photographs of Palaealeurodicus borneensis sp. nov., puparia, showing lateral waxy palli-
sade supporting delicate cuticle, and dorsal wax carapace in situ, Sungai Melinau, Sarawak.

FIGURES 138–142 Aleurodicus spp, adults. 138, A. guppyii Quaintance & Baker, forewing (after Quaintance & Baker,
1913). 139, A. pulvinatus (Maskell), forewing, Nevis. 140, A. floccissimus (Martin et al.), forewing, male, Tenerife. 141,
A. floccissimus, female abdomen, dorso-ventral, Tenerife. 142, A. floccissimus, female abdomen, lateral, showing three
of the four lateral wax plates.

FIGURES 143–144 habitus photographs of Aleurodicus mirabilis (Cockerell), on Annona sp., East Coast Demerara,
Guyana. 143, developing colony with characteristic secretion of individual puparia clearly visible. 144, mature colony,
completely obscured by puparial secretions that have been folded back to reveal feeding adults.

A Revision of Aleurodicus Douglas With Two New Genera Proposed For Paleotropical Natives An Identification Guide To World Genera

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A Revision of Aleurodicus Douglas With Two New Genera Proposed For Paleotropical Natives An Identification Guide To World Genera

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ZOOTAXA

A revision of Aleurodicus Douglas (Sternorrhyncha, Aleyrodidae),

FIRST PUBLISHED IN 2008 BY

© 2008 Magnolia Press

ISSN 1175-5326 (Print edition)

2 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

A revision of Aleurodicus Douglas (Sternorrhyncha, Aleyrodidae), with two new

Accepted by C. Schaefer: 7 May 2008; published: 30 Jul. 2008 3

Abbreviations for specimen depositories

BMNH – Natural History Museum, London SW7 5BD, U.K.

Materials & methods

4 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Morphological terminology, with special reference to dorsal puparial pores

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 5

6 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 7

8 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Designation of primary type specimens

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 9

Check list of taxa discussed in this paper

10 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Key to the world genera of Aleurodicinae – puparia

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 11

12 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 13

14 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Key to Aleurodicus species – puparia

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 15

16 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 17

18 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 19

Redescription, from neotype puparium

20 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

DISTRIBUTION. Neotropical Region – Belize, Costa Rica, Mexico.

Aleurodicus capiangae Bondar

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 21

Aleurodicus charlesi sp. nov.

22 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Aleurodicus chirripoensis sp. nov.

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 23

Aleurodicus clusiae sp. nov.

24 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Aleurodicus coccolobae Quaintance & Baker

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 25

Aleurodicus cocois Curtis

Aleurodicus darienpalmae sp. nov.

26 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Aleurodicus decemensis sp. nov.

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 27

Aleurodicus dispersus Russell

28 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Aleurodicus dugesii Cockerell

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 29

Aleurodicus etiennei sp. nov.

30 · Zootaxa 1835 © 2008 Magnolia Press REVISION OF ALEURODICUS

Aleurodicus flavus Hempel

Aleurodicus flavus Hempel, 1922a: 4. Syntypes, Brazil [MZUSP, not examined].

DISTRIBUTION. Neotropical Region – Brazil.

MARTIN Zootaxa 1835 © 2008 Magnolia Press · 31

Aleurodicus guppyii Quaintance & Baker

DISTRIBUTION. Neotropical Region – Trinidad.

Aleurodicus inversus Martin