Giant Whiteflies (Sternorrhyncha, Aleyrodidae) a Discussion of Their Taxonomic

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    IntroductionIn an account of the history of higher systematicsin the Aleyrodidae, Russell (2000) stated that fivewhitefly subfamily names have been used for extanttaxa. Of these, Uraleyrodinae Sampson & Drews(1941) was found to be synonymous with Aleyro-dinae Westwood (1840), based on a study of adultcharacters by Russell (1986). Takahashi (1932) haderected the subfamily Siphonaleyrodinae solely forhis new species Siphonaleyrodes formosanus, whichis clearly a member of the psylloid family Triozidae,and which was placed as a junior synonym of Triozacinnamomi (Boselli, 1930) by Mound & Halsey(1978), a view with which Russell (2000) concurred.The oldest-established subfamily, Aleyrodinae, isgenerally accepted and regarded as well defined byadult and nymphal [puparial] characters (Gill 1990).This leaves Udamoselinae Enderlein (1909) andAleurodicinae Quaintance & Baker (1913) whosecontroversial relationship is the subject of this paper.The genus Udamoselis, the species U. pigmentariaand the subfamily Udamoselinae were all proposedby Enderlein (1909), based upon his study of a

    single adult male specimen. Enderleins specimen hassubsequently never been traced, and is thought tohave been lost during the upheavals of the SecondWorld War. As well as being described from a singlespecimen, no satisfactory collecting locality is knownand Enderlein simply gave this as in all probabilitySouth America, indicating that the specimen musthave been given to him. Enderlein also includedAleurodicus Douglas (1892) in his new subfamily,without any discussion.Quaintance & Baker (1913) discussed whitefly wingvenation in detail, illustrating a range of actual andtheoretical patterns (Fig. 33). They proposed an-other new subfamily, Aleurodicinae, accommodat-ing Aleurodicus, Dialeurodicus Cockerell (1902),their own new genus Leonardius and ParaleyrodesQuaintance (1909), whilst continuing to acceptEnderleins subfamily Udamoselinae for Udamoselisalone. Their reason for supporting a separate sub-family for Udamoselis was the more complex wingvenation described and illustrated by Enderlein(Fig. 7), but the insects enormous size (Table 1) maywell have also been a factor in their decision. The

    Giant whiteflies (Sternorrhyncha, Aleyrodidae):a discussion of their taxonomic and evolutionarysignificance, with the description of a new speciesof UdamoselisEnderlein from Ecuador

    Jon H. Martin

    Three adult male whitefly specimens from Ecuador are described as Udamoselisestrellamarinaesp. n. This genus and its subfamily are reappraised on adultcharacters, including wing venation, paronychium structure, and distribution ofabdominal wax glands. In the absence of associated puparia nomenclatural caution

    is preferred, but the subfamilies Udamoselinae and Aleurodicinae are likely to besynonymous. Wing venation of other very large whiteflies is illustrated, and isdiscussed in comparison with fossil taxa. Speculation is made on the possible biologyof such giant whitefly species.Jon H. Martin, Department of Entomology, Natural History Museum, CromwellRoad, London SW7 5BD, UK. j. [email protected]

    Tijdschrift voor Entomologie 150: 1329, Figs. 133, Table 1. [ISSN 0040-7496]. 2007 Nederlandse Entomologische Vereniging. Published 1 June 2007.

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    relative complexity of the wing venation and greatbody size, in turn, have been regarded as possible evi-dence that Udamoselismight be a particularly primi-tive whitefly, and thus form a link between the otherpresent-day taxa and species described from the fossilrecord (such as by Schlee 1970, Shcherbakov 2000,and Hamilton 1990).U. pigmentariawas regarded as a nomen dubiumbyMound & Halsey (1978: 250), on the basis of in-adequate description of the adult, combined withthe absence of knowledge of the puparial stage uponwhich most whitefly taxonomy is now based. Ender-leins description of the solitary male did indeed omitmention of some characters that are now thoughtlikely to be important in the systematics of adults,and this supported the proposal that it be regardedas nomen dubium. Perhaps most importantly, End-

    erleins description and illustration of the fore andhind wings (see Fig. 7, here) showed a venation thatis considerably more complex than had been seen inany other known extant whiteflies: this raised a ques-tion as to whether all the firm lines in Enderleinsdrawings were truly veins and, hence, whether hisillustrations were accurate.Leaving aside the uncertainty over wing venation,the absence of any detail of such characters as ab-dominal wax glands or tarsal paronychium, the lossof the antennal flagellum in his sole specimen, and

    the lack of optical resolution available to Enderlein[he stated that no empodial paronychium was vis-ible with his magnifying glass], his description wasnonetheless remarkably detailed if sometimes ratherambiguous. With the considerable importance ofthis taxon Quaintance & Baker (1913) provided acomplete English translation of Enderleins descrip-tion of U. pigmentariaand this translation has beenextensively consulted in the course of the presentstudy. In the absence of study material, Mound andHalseys (1978) decision to regard U. pigmentariaasnomen dubiumwas pragmatic, allowing the continu-ing use of Aleurodicinae as the name for the numeri-cally smaller of only two extant whitefly subfamilies,accommodating about eight percent of describedwhiteflies.Schlee (1970) stated that The systematic position ofUdamoselis within the Aleyrodina cannot be elabo-rated until a new find is made, because of the insuffi-cient present knowledge based upon the single specimen,which has probably been destroyed. The assumed closekinship relation between Udamoselis and the Aleu-rodicidae [i.e. Aleurodicinae] is unproved.In contrast,

    Shcherbakov (2000) said: . despite an incompleteknowledge of the type genus, the name Udamoselinaeshould be used in the broad sense of Enderlein (1909)and Sampson (1943), i.e. including Aleurodicinae.

    Three new specimens recently collected in Ecuadorcorrelate with Enderleins description sufficiently wellto be regarded as belonging to Udamoselis, thus al-lowing this intriguing controversy to be reappraised.Many of the attributes described by Enderlein forU. pigmentaria are apparently accurate, althoughother parts of his description remain ambiguousthrough the absence of the original specimen, com-bined with Enderleins failure to provide any illus-trations beyond the wings. Nevertheless the authornow considers it quite likely that U. pigmentariait-self will prove to be identifiable, in the event of newmaterial becoming available, and its identity shouldno longer be regarded as nomen dubium. However,the subfamilial position of Udamoselisremains some-what uncertain, as will be discussed later in thispaper.

    Examination of the three males from Ecuador hasrevealed their wings (Figs 5, 6) to display the identi-cal venation illustrated by Enderlein (Fig. 7), but hasconfirmed that not all veins are as distinct as impliedby Enderleins simplified line drawings. Comparisonof the Ecuadorean material with the description ofU. pigmentarialeads to the conclusion that the twotaxa are congeners but are distinct species. Despitethe small sample size, and frustrating lack of femalesand (especially) of puparia, it is felt that naming theEcuadorean species is valid because of the wider in-

    terest in higher systematics that these specimens arelikely to generate. Udamoselis estrellamarinaeis there-fore here described, and is named for its discoverer(see below).

    Materials, methods and terminology

    BackgroundIn 2005 the author visited Ecuador, in companywith Dra Estrella Hernndez-Suarez and Sr ElicioTapia. The purpose was to search for whitefly col-onies that might yield natural enemies of the pestspecies, Lecanoideus floccissimus Martin, Hernn-dez-Surez & Carnero, 1997, in connection withachieving its natural control in the Canary Islands.Whilst sorting collected material for possible rearingof parasitoids, Hernndez-Suarez noticed three verylarge, darkly-pigmented and relatively wax-free adultmale whiteflies inside a bag containing a substantialcolony of Lecanoideus mirabilis (Cockerell, 1898)onAnnonaleaves. An extensive search of other bagsof material from the same garden tree failed to re-

    veal any additional specimens. The three specimenswere brought back to the laboratory at the NaturalHistory Museum, London, for further study.

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    Treatment of adult whiteflies for examinationIt is not easy to dissect any adult whitefly for mi-croscopic examination on slides, in contrast with thesituation with Psylloidea (jumping plant lice) whichare routinely dissected prior to slide-display. Never-theless, the interest in wing venation of larger white-flies leads the author now to remove all wings fromsuch specimens, prior to maceration of the body, andto mount the completely untreated wings under aseparate coverslip. This avoids distortion and dete-rioration of any subtle wing pigmentation, and theseparate coverslip allows for a very thin (hence flat)mount for the wings.With very large taxa a partial dissection of the bodyalso assists in better displaying certain characters forstudy, but with very small sample sizes this inevita-bly results in some other characters being distorted

    or obscured. The author usually dissects the head(sometimes with the rostrum still attached) for exam-ination frontally, the abdomen for lateral mounting,male forceps for dorso-ventral display, and the aedea-gus for lateral examination. The thorax, with legs stillattached, may be mounted laterally (Fig. 1) for thepractical reason that whitefly coxae are so stronglyventrally-directed that normal dorso-ventral displayis virtually impossible. Alternatively, the specimenmay be more extensively dissected (Fig. 4), allow-ing more constituent parts to be displayed parallel

    to the slide surface. With the much thicker mountsthat are needed for the thorax / legs and abdomen, incomparison with that required for the genitalia andhead, the constituent parts of the specimen may bedistributed between two to four coverslips of 10 mmor 13 mm diameter (Fig. 4), but always with all theparts of one adult on a single slide. The practice ofdividing the constituents of one individual betweentwo slides, sometimes used by workers on Psylloidea,is to be avoided.When adult whiteflies are not dissected, the favouredalignment is generally lateral, with wings dorsally (asin the resting position of Rhopalocera) and withlegs and rostrum displayed ventrally as seen in Fig.1. With much smaller adults, usually members ofthe Aleyrodinae, displaying them is more challeng-ing, because manipulation requires even more careand osmotic collapse becomes a greater risk. Whena large sample is at hand, the author usually resortsto placing many individuals under a single coverslip,with minimal effort being expended on each indi-vidual: the result is that usually all characters maybe seen, but only by examining several specimens.

    With very small adults, better results are sometimesachieved by clearing and dehydrating un-maceratedspecimens and placing them directly onto a slide.

    Wing venation terminologyThe wing venation of Udamoselis pigmentaria isimportant for the discussion of its genus and otherlarger whiteflies. Enderlein (1909) employed a termi-nology (Fig. 7) that is not accepted today, but thereappears to be little consensus on which alternativeto use. Accordingly, for this communication I havefollowed the system used by Quaintance & Baker(1913; see Fig. 33, here), Solomon (1935) and Gill(1990). For Udamoselis this is shown in figures 5 and6, where ? indicates my uncertainty over whetherthe posteriormost feature on each wing is truly avein. This venation terminology is also used here indiscussion of the wings of several other large whiteflyspecies (Figs 19-28).


    BMNH The Natural History Museum, London,U.K.

    USNM Entomological collections of the U.S. Na-tional Museum of Natural History, housedat U.S. Department of Agriculture, Belts-ville, Maryland, U.S.A.

    The holotype and paratype 2 of U. estrellamarinaeare deposited in BMNH; paratype 1 of U. estrellama-rinaeis deposited in USNM, and all three specimenshave been slide-mounted as described above, withvarying degrees of dissection. All the other specimens

    discussed and illustrated here are also housed in thecollection of BMNH.


    UdamoselisEnderlein, 1909: 230. Type species: UdamoselispigmentariaEnderlein, by monotypy, but sole originalspecimen not traced.

    Udamoselis estrellamarinaesp. n.Figs 16, 818

    Type material.Holotype: adult ?, Ecuador, ManabProvince coast, Brisea, 5 km north of San Vicente,onAnnonasp., probablyA. muricata(Annonaceae),10.ii.2005 (Hernndez-Suarez coll., Martin ref.#8103) (BMNH). ?: same data asholotype (BMNH; USNM).

    Adult male (n=3)Measurements. For basic measurements of manybody parameters, see Table 1.Coloration. Body colour very dark, with little ap-

    parent waxy bloom covering the cuticle, resemblingsmall Psylloidea to the naked eye. Pigmentationof the body cuticle as shown in the photographs(Figs 1, 4), but is generally evenly dark brown; the

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    middle parts of the femora and the whole of the

    tibiae, tarsal segments and antennal flagellar seg-ments are paler. The wings are rather leathery andtheir dark pigmentation (Figs 23) can best be de-scribed as blotchy; each fore wing has two ovoid palepatches, and each hind wing a single trapezoidal palezone; the degree of darkening of the wings varies be-tween the three specimens, with the holotype (Figs23) the darkest.Legs. Femoral hairs normal on all legs, bristle-like;many hairs on distal parts of tibiae thickened butstill apically acute; each middle tibia with a groupof thickened hairs forming a distinct comb at three-quarters length, each hind tibia with a more subtlecomb formed by a smaller number of thickenedhairs, but each fore tibia without such a comb ofhairs; each hind tibia with an apical arc of 911 paledagger-like spines (Fig. 18) that may be saltatorial infunction (as in Psylloidea), but these are absent fromother legs. Tarsi two-segmented, basal segment muchlonger than apical segment; hairs on basal segmentstouter and more numerous than those on apicalsegment; hind basal tarsal segment with two rows ofstout ventral spines (Fig. 17), which (like the api-

    cal hind tibial spines) may be saltatorial in function;claws paired, each pair with a single stout ventrally-directed spine-like paronychium between them(Fig. 9). Tibial and tarsal surfaces with very fine

    spinules, becoming dense enough on tarsi to be re-

    garded as a fine pubescence.Abdomen.With a pronounced furrow running alongeach side of segments IIIVII, above the mid-line;segment VIII about as long as deep, slightly ex-panded at distal end, without a ventral spur at itsdistal extremity; segment IX about 3 times as long asdeep; sternites IIIV each with a pair of wax plates(Fig. 14), the anterior two pairs significantly largerthan the posterior pair; each anterior wax plate witha deep invagination from its anterior side, lined withhairs and accommodating a patch of rounded reticu-lations; middle wax plates with a similar reticulatepatch situated in an indentation of the anterior edgeof the plate; hind wax plates evenly ovoid, withoutindentations and not fringed with hairs; surfacesof wax plates extremely finely imbricate-reticulate(400magnification), with a few hairs on their sur-faces as shown in figure 14; segments VIIIIX devoidof hairs but other segments with hairs surroundingwax plates (Fig. 14) and near postero-ventral marginsof sternites; tergites IIIIV each with a subcircularpatch of rounded reticulations on each side (Fig. 16);whole of abdomen punctuated by tiny pale ovoid

    pores (Figs 1416). Lingula and presumed oper-culum (Fig. 15) situated between one-third andhalf way from base to apex of segment IX; lingulawith fine spinule-pubescence but no evident hairs;

    Figs 14. Photomicrographs of Udamoselis estrellamarinae. 1, laterally-mounted body, holotype; 2, fore wing,holotype; 3, hind wing, holotype; 4, dissected thorax / legs and abdomen, paratype 2.





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    presumed operculum with similar spinule-pubes-cence but only with a single apical hair visible inlateral aspect [it is to be expected that there is a pairof such hairs]; claspers evenly curved in dorso-ven-tral aspect (Fig. 10) but flat in lateral aspect (Fig. 1),generally smooth and with a few fine hairs as shownin figure 10; aedeagus with a distinct elbow at half-length (Fig. 12), about 0.325 mm from apex to el-bow, apically with two large and possibly four muchsmaller finger-like protrusions (Fig. 13).Head. As shown in antero-ventral view in figure 8;small for the size of the insect (1 mm wide), muchwider than long; compound eyes very large and in-dented to accommodate the antennal insertions;lateral ocelli well-developed, as shown in figure 8,but median ocellus not evident in any of the three

    mounted specimens; between the eyes is situated aconical protrusion from the frons; clypeus denselyhairy; basal two antennal segments dark brown,smooth and with fine hairs; the five-segmented

    antennal flagellum (Fig. 11) without hairs but withfinely spinulose transverse striations throughout itslength (Fig. 11, expanded detail); flagellar segmentswith many extremely small, pale spots that are likelyto be sensoria, with a few slightly larger sensoriaon the apical segments; ultimate rostral segmentpigmented dark brown in apical third, with manyhairs.Thorax. Thoracic dorsal plates smooth, pigmentedbrown and sparsely provided with fine hairs, exceptfor a fringe of longer hairs anteriorly on prothorax.

    Wings. Rather narrow, dimensions given in Table 1,fore wing margin flattened to slightly emarginate op-posite apices of veins Rsand M (Fig. 5). Pterostigmawell defined by surface roughening (Fig. 5) ratherthan by differentiated coloration (Fig. 2). Venation

    of fore wing as shown in figure 5; Sc contiguous withC until diverging as it approaches the pterostigma;R1 short, becoming indistinct in the pterostigma;Rs almost reaching wing margin; M very long,












    r1 r m


    anax r


    an m?Cu

    Figs 57. Wings of Udamoselisspecies. 5, U. estrellamarinaesp. n., fore wing with venation annotated using ter-minology adopted here; 6, U. estrellamarinaesp.n., hind wing with venation annotated using terminology adoptedhere; 7, U. pigmentaria, original figure from Enderlein (1909), showing Enderleins venation terminology. [Scalebars = 1 mm]

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    10 11





    Figs 815. Udamoselis estrellamarinae. 8, head, anteroventral view, paratype 2; 9, claws and empodial spine;10, clasper, paratype 1; 11, antenna, with detail of apical 2.5 segments, paratype 1; 12, aedeagus, paratype 2;13, aedeagal apex of (a) paratype 2, (b) paratype 1, (c) holotype; 14, wax plates on abdominal sternites IIIV,holotype; 15, lateral aspect of operculum (left) and lingula (right), paratype 1.

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    16 17






    Figs 1622. 1618, Udamoselis estrellamarinae. 16, oval patches of reticulations on abdominal tergites III andIV, paratype 2; 17, distal half of hind basitarsus, paratype 1; 18, apex of hind tibia, showing fringe of spines andthickened setae proximad, paratype 1. 1920. Wings of male syntype of Parudamoselis kesselyakiVisnya. 19, forewing; 20, hind wing. 2122. Wings of unidentified male whitefly from Tena, Ecuador. 21, fore wing; 22, hind wing.[Scale bars for wings only = 1 mm, one per wing pair]

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    Figs 2328. Wings. 2326, Wings of unidentified whitefly (JHM 8078) from Bartola, Nicaragua. 23, forewing, male; 24, hind wing, male; 25, fore wing, female; 26, hind wing, female; 2728. Wings of Dialeurodicuscaballeroi, male. 27, fore wing; 28 hind wing. [Scale bars = 1 mm, one per wing pair]

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    diverging from R near wing base, almost straightbut slightly angled posteriorly in its apical quarter;Cu (see discussion, below) faintly-indicated basallyand apically, parallel to and closest to M at half-length, markedly paler than the heavily pigmentedwing surface; posterior to Cu is a subtle and irregularfold that is clearly what Enderlein regarded as theAxillaris [anal vein, here], but it is unclear whetherthis really warrants such status. Hind wing with R /R1& Rs clearly marked, with M diverging from Rat wing base; in a similar situation to that seen in thefore wing, a subtle thickening / fold indicates whatEnderlein regarded as the Analis [vein Cu here].Coloration and patterning of wings as shown inFigs 23.Whichever fore wing venation system is followed theentity named Cu (Fig. 5) is clearly associated with

    the claval furrow (sutura clavi of Enderlein), alwaysalmost straight for much of its length and distinctlypale on pigmented wings (a very distinct bright lineas described by Enderlein for his fore wing Analisin figure 7). In Psylloidea, the same observations aretrue for vein Cu2which doubles as the clavus (Hod-kinson & White 1979). Indeed, the present authorquestions whether this is really a vein at all.Remarks. U. estrellamarinae differs from U. pig-mentaria in some obvious respects, assuming End-erleins description of the latter to be accurate. The

    wings of U. estrellamarinae are more elongate inshape, clearly seen by comparing figures 5-6 withfigure 7, and by comparing the parameters given inTable 1. The margin of the fore wing of U. estrel-lamarinaeis distinctly emarginate opposite the apexof vein Rs, and slightly less so opposite the apex ofM, in contrast to the evenlycurved distal margin inU. pigmentaria. Enderlein described the hind wingof U. pigmentaria as densely sprinkled with darkbrown spots, in contrast to the almost uniformlydark hind wing, punctuated by tiny pale spots(Fig. 3), in U. estrellamarinae: otherwise, the descrip-tion of the wing pigmentation in U. pigmentaria istorturous and demonstrates well why photographsare the best way to illustrate such characters.Despite the wings of U. pigmentaria beingsomewhat longer, and much broader than inU. estrellamarinaethe probability is that the body ofU. pigmentariais smaller than in U. estrellamarinae,with abdominal segment IX (see Table 1) being aparticularly clear indicator despite the ambigu-ity over whether Enderleins stated body length forU. pigmentaria was inclusive or exclusive of the

    forceps.The conical protuberance on the frons was describedas black in U. pigmentariabut is concolorous withthe remainder of the head in U. estrellamarinae.

    Giant size and wing venation in theAleyrodidaeFrom the description and discussion, above, Uda-moselis is clearly a genus with exceptionally largemales. With no females available for study, it is by nomeans certain that females will prove to be as large.There is a good reason for sounding such a caution-ary note (see below).Figures 19 and 20 show the fore and hind wings ofthe sole syntype (an adult male) of ParudamoseliskesselyakiVisnya (1941) in BMNH. Described as agigantic whitefly by Visnya (from a large introducedpopulation in a Hungarian glasshouse), the BMNHmale is actually even larger than the Udamoselismales(see Table 1), but the body cuticle and wing colora-tion are more typical for a whitefly, not being darklypigmented. Although P. kesselyakiis now regarded as

    a junior synonym of Ceraleurodicus varus (Bondar,1928), based on comparison of syntype puparia,(see below) of both species in the USNM collection(Martin et al. 2000), the discussions here are basedupon the BMNH adult male syntype of P. kesselyakiand Visnyas observations on the Hungarian mate-rial from the 1940s; no other males are known atthe present time. The fore wing of the BMNH syn-type of P. kesselyaki(6.00 mm long, 2.95 mm maxi-mum width) displays the same much-curtailed R1,terminating in a pronounced pterostigma,which is

    also seen in Udamoselis. There is also a suggestionthat thickening of the proximal boundary of thepterostigma could be the equivalent of the putativevein Sc in Udamoselis. As in the males of both spe-cies of Udamoselisand other large species discussedhere, the result is a more robust fore wing. Visnyaalso remarked upon distinct adult size-dimorphismin P. kesselyaki. Four females reared from puparia onProtium copal in Belize, and identified as C. varusfrom their puparia, are considerably smaller than theonly (male) syntype of P. kesselyakiin BMNH, andalso smaller than Visnyas quoted measurements forfemales of P. kesselyaki.Visnya further noted that the male puparia ofP. kesselyakiwere significantly larger than those of fe-males, as follows: The length of most L4is between 3.8and 4.3 mm. These are all females and the well devel-oped eggs can be seen in matured ones before emerging.There are L4of 5 mm and above, these seem to be males,so far however I could only find one preparation with alarva of 5.3 mm length and 2.5 mm wide, in which thepenis and claspers are visible.[Authors note: the observations below concern some

    undescribed whitefly taxa. The characteristics de-tailed here are not intended to be for nomenclatu-ral purposes, but only to broaden this discussiontopic.]

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    In 2004 a congregation of extraordinary adult white-flies was discovered under a single leaf in Nicaragua[Uncaria tomentosa (Rubiaceae), Rio San Juan / RioBartola confluence,, Martin #8078]. Thegeneric position of this undescribed species is un-certain, because almost nothing is known about therelative importance of different adult characters inwhitefly generic definitions. This situation has arisenbecause for 150 years whitefly taxonomy has beenbased almost solely upon puparial [i.e. final nymphalstage] characters (Martin 2003). This species is herereferred to simply by its collection number, JMH8078, and it is notable for two reasons. Firstly, one ofthe five males (Fig. 29) may be the largest individualwhitefly ever collected, with an overall body lengthof over 1 cm (see Table 1). Despite this exceptionallength, the body is possibly less massive than themales of Udamoselis, with the extreme length beingdue to abdominal segments VIII and IX (Fig. 29).At 3.78 mm long in this largest specimen, its ab-

    dominal segment IX alone is longer than almost allother whitefly species, including those in the Aleuro-dicinae. However, the fore wings of these males (3.80mm long in the largest specimen, down to 3.20 mm

    in the smallest) are considerably smaller than thoseof Udamoselismales, presumably reflecting a lowerbody mass.JMH 8078 is also notable for extreme adult dimor-phism (Fig. 29). The adult females (body length2.102.60 mm, fore wing length 1.752.20 mm,n=7) are all very much smaller than the males, andare similar in size to many other Aleurodicinae. Thedifference in wing size, between males and femalesof JMH 8078, is at once obvious from comparisonof figures 2324 with 2526 (all shown to the samescale). Size apart, while the female fore wing displaysan entirely typical aleurodicine venation, that of themale has vein R1converging with the costal margin,there forming a small pterostigma, and then contin-uing towards the wing apex and almost parallelingthe curve of Rs. The increased complexity of the forewing venation in the male results in a more robuststructure.Figures 21 and 22 show the fore and hind wings

    of a single large (dissected) male specimen (forewing 3.92 mm long, 2.19 mm maximum width)from Ecuador [Tena, 23.ii.1923, F.X. Williams coll.],which also remains unplaced generically. Here, R1is

    Fig. 29. Unidentified whitefly (JMH 8078) from Bartola, Nicaragua largest male specimen (left) and a female

    specimen (right) from same collection, in alcohol prior to slide-preparation, showing extreme dimorphism.

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    Figs 3032. Likely models for Udamoselispuparia habitus photographs of puparia of species of Aleurodicinae withsparse wax, radial rays and additional ventral tracheal folds, that do not occur in aggregations. 30, CeraleurodicuskerisMartin, on Lunania parviflora, Nicaragua; 31, Octaleurodicussp., on ?Melastomataceae, Ecuador; 32, Ceraleu-rodicussp., on Clusia grandiflora, Guyana.


    31 32

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    Fig. 33. Reproduction of Plate 1 from Quaintance & Baker (1913), showing wing venations of present-day white-flies, theoretical origin of whitefly wing venation, wing venation of present-day Triozasp.(Psylloidea) and theoreti-cal origin of triozid wing venation.

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    reminiscent of JMH 8078 but, instead of a pte-rostigma, there is a curious fusion with the putativeSc. Also, M is broken at its proximal end, with thebasal stub displaced towards Cu (this curious fea-ture is identical in both fore wings). Once again, theresult is a more robust structure, possibly to supporta larger-than-usual body mass.Although very large males are sometimes associatedwith much smaller females (JMH 8078 and P. kes-selyaki [= C. varus], see above), there are also verylarge female whiteflies in existence. One such female[Chiococca alba(Rubiaceae), Belize, Chiquibul forestReserve,, Martin #7967], also of uncer-tain generic position, has fore wings 3.65 mm long,2.00 mm maximum width and hind wings 3.10 mmlong, 1.40 mm maximum width. The abdomen hasfour pairs of wax glands, the posteriormost two pairs

    with coarsely reticulate facets (which are reminiscentof those in Aleurodicus dispersus). The body, is notdark brown and the wings have only isolated spots ofpigment, so it is not thought likely that this individ-ual will prove to be a species of Udamoselis. Despitethe large size of the wings of this female, the venationis typically aleurodicine, with Rwidely-forked andno pterostigma.Finally, figures 27 and 28 show the wings of the maleof Dialeurodicus caballeroi Martin, 2004, a muchsmaller species than those discussed above (fore wing

    1.83 mm long, 0.86 mm maximum width). This is awing type commonly seen in aleurodicine species ofmedium size, with R1simply but well developed inthe fore wing, similar to that seen in the hind wing,but contiguous with a large pterostigma in the forewing. There is no indication of size dimorphism inD. caballeroi, with female fore wings even slightlylonger and broader than in most males. [The pter-ostigma in both sexes is much less distinct in speci-mens whose wings have been through the macera-tion process prior to slide-mounting.]In all the taxa discussed above, the hind wings havethe putative Cu either short and poorly indicated, orapparently absent, and are thus typical of aleurodi-cines. The only oddity amongst these hind wings isseen in the male syntype of P. kesselyaki (Fig. 20),where R1has a curious (?vein) stub arising at its half-length, a feature seen in both of the hind wings.

    Observations on wings and othercharacters, with respect to subfamilyplacement in extant taxa

    Added to the detailed comments on the very largetaxa, above, the following broader observations maybe made: Almost all members of the Aleurodicinae have

    the fore wing with R forked to form R1 and Rs,and are larger insects than most members of theAleyrodinae. Udamoselis also displays these at-tributes.

    Most members of the Aleyrodinae do not have aforked R, but only a single main vein.

    The only Neotropical members of the Aleuro-dicinae that are physically very small, and thussimilar in size to typical members of the Aleyro-dinae, are the members of the genus Paraleyrodes,whose fore wings have an unbranched main vein,as is typical in the Aleyrodinae.

    A few unusually large members of the Aleyrodi-nae have their fore wings with Rforked, or haveother complications to their venation. Examplesinclude European species of Aleurochiton Tull-gren, 1907 and the northern Australian native,

    Gagudjuia allosyncarpiae Martin, 1999 and atleast one species of AleuroparadoxusQuaintance& Baker, 1914 [an undescribed species fromGuyana].

    All the males discussed here, including Udamose-lisand JMH 8078, have three pairs of abdominalwax plates normally regarded as a diagnosticaleurodicine feature (Gill, 1990).

    All the males discussed here, including Uda-moselisand JMH 8078, have a spine- or seta-likeparonychium, again regarded as a diagnostic

    aleurodicine feature. Size dimorphism is also quite common in theAleyrodinae but it is then usually the adultmales, and male puparia, that are smaller. In gen-era such as Aleurocanthus, the male and femalepuparia have not infrequently been described asseparate species.

    The fossil dimensionSchlee (1970) presented a detailed discussion of Cre-taceous and Tertiary amber-fossilised whiteflies. Aspart of his comparisons of these with extant taxa, hesaid the following:The recent Udamoselisis by no means the most primi-tive recent Aleyrodid; it does not represent the Aley-rodinas ground plan. Moreover it is one of the highlyderivative forms, exhibiting numerous autapomorphies(one of which is the enormous size). The relatively com-plete and distinct venation is connected with the largebody size.He further stated that The wing venationgives no evidence for the kinship relations within theAleyrodina; families must not be defined by this fea-

    ture. These views are supported by the quoted small-to-medium size of the following fossil taxa: Heideacretacica Schlee, 1970, body length 1.02 mm, forewing length 0.83 mm; Bernaea neocomica Schlee,

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    Tijdschrift voor Entomologie, volume 150, 200726

    1970, body length 1.30 mm, fore wing length 1.08mm; Juleyrodes gilli Shcherbakov, 2000, fore winglength 2.1 mm (no associated body); BurmoselisevelynaeShcherbakov, 2000, body length 0.95 mm,fore wing length 1.10 mm. The wing venation ofH. cretacicawas not discernible, but the venations ofthe other three species vary from similar to present-day Aleurodicinae to slightly more complex thanin Udamoselis. Another fossil species, MegaleurodesmegocellataHamilton, 1990 [Cretaceous], measured11.5 mm in length but is clear to the present au-thor that Shcherbakov (2000) has already correctlyregarded this particular insect as being a member ofthe Fulgoroidea (Auchenorrhyncha), having three-segmented tarsi and legs that would be characteristi-cally angular in cross-section.These limited data do not, of course, preclude the

    possibility that very large whiteflies have occurredin the fossil record. Nevertheless these observationscombine to suggest that there has been an evolu-tionary trend towards the extremely simplified wingvenation seen in over 90 percent of extant whiteflyspecies, and that this trend may have been reversedfor larger species, with a more complex venation re-tained or re-evolving in the cases of Udamoselis, mostmembers of the Aleurodicinae and some larger mem-bers of the Aleyrodinae.

    Observations on biologyAll specimens of the very large taxa, and specimensof several other larger species, are represented inthe BMNH collection only by small numbers ofadults, these sometimes having been discovered inapparent mating congregations with no associatedpuparia. A personal observation, that the author be-lieves to be correlated, is that puparia of several largealeurodicine species currently placed in Ceraleu-rodicus Hempel, 1922 are found widely scatteredover their hosts, and not in the kind of aggregationsthat are typical for most other species in the Aleu-rodicinae. It is considered possible that the adultsof such species congregate for mating, but that thefemales then distribute single eggs very widely. If thisis the strategy also adopted by Udamoselisthen it islikely that the three individuals of U. estrellamarinaewere a nucleus for such a mating group, and it is alsolikely that puparia will be difficult to find.Visnyas comments on male puparia of P. kes-selyaki concur with the present authors field ob-servations of this whitefly group (see description of

    Ceraleurodicus keris and account of C. varus byMartin 2004) that males are even more difficultto find than are females, and no males or probablemale puparia of either C. varusor C. kerishave been

    discovered despite detailed searches.A characteristic of the puparia of larger spe-cies currently accommodated in Ceraleurodicus,Dialeurodicus and Octaleurodicus Hempel, 1922(Figs 3032) is that they feature nine pairs of radialrays [peripheral intersegmental ridges of Shcherba-kov 2000]. At least some of these rays terminate invery fine combs of modified marginal teeth, and thesecreted glassy peripheral filaments are thus narrowerat these points, appearing more opaque and render-ing the rays clearly visible within the glassy skirt thatsurrounds the puparium (Figs 3032). Some rays insome species also have apparent tracheal folds un-derlying them ventrally (Martin 2004). Shcherbakovconsidered that this tracheal feature indicates a morecomplete complement of spiracles than is usual inwhitefly puparia. Such puparia also tend to have their

    compound pores reduced in size and number (thoseentirely without compound pores being assignedto Dialeurodicus), and do not produce the copiouswoolly tangles of secretions that are so commonlyassociated with other members of the Aleurodicinae.Three puparia of the type discussed above are shownhere: figure 30 depicts Ceraleurodicus keris, with anasymmetric puparium and with a very long and com-plex filament arising from each compound pore, butotherwise highly cryptic; figure 31 shows a species ofOctaleurodicus, with short and glassy fingers of wax

    secreted by eight small, submedian compound pores;figure 32 features a species of Ceraleurodicuswith ex-tremely reduced (submedian) compound pores, andno discernible dorsal secretions at all.One can only speculate on the size and appearanceof puparia of Udamoselis, or those of other giantmales such as those of JMH 8078, but Visnyas ob-servation that the male puparia of P. kesselyaki aresignificantly larger than those of females is likely tobe relevant. The puparia of both C. varus (recent-ly-collected material from Central America) andC. keris have been recorded as reaching 4 mm inlength, but these may all be female. The widely scat-tered puparia of at least some species currently placedin Ceraleurodicus are highly cryptic in life, hinder-ing field searching. This was especially marked withC. varusin Belize (Martin 2004).A few old puparia of C. keriswere discovered on thesame host plant as the aggregation of adults of JMH8078, and it remains a possibility that these two en-tities may be conspecific [no adults of C. kerishavebeen reared from puparia].

    Provisional conclusionsBased upon all the adult male characters forUdamoselis, now available for study on actual speci-

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    Martin: Giant whiteflies and new Udamoselis 27

    mens, there is nothing that suggests to the authorthat Udamoselisshould be regarded as belonging toa different subfamily from Aleurodicus and othermembers of the Aleurodicinae. This concurs with theconclusions of Solomon (1935), Sampson (1943),Schlee (1970) and Shcherbakov (2000), as discussedby Martin & Streito (2003). The same may be saidof the other large taxa with relatively complex wingvenation, discussed here. In large part, this conclu-sion is based upon the proposition that more com-plex wing venation is a response to larger body mass(as promulgated by Schlee 1970), and is thus nota feature of importance for subfamilial placement.This proposition is made because other major at-tributes are entirely typical for Aleurodicinae.It therefore follows that there is no compelling rea-son for regarding Udamoselis as providing evidence

    of a missing link between fossil whitefly taxa andthose alive today. It is of particular note that at leastsome fossil taxa were not giants (see above), despitetheir more complex wing venations, leading Schlee toconclude that Udamoselisis actually highly derived,rather than plesiomorphic, in some of its features.However, the continuing (and frustrating) lack ofadult females and (especially) immature stages, reli-ably associated with males of Udamoselis, prevents afinal conclusion from being drawn here on its subfa-milial position. Should future collecting yield females

    and puparia, which further support the view thatUdamoselis,Aleurodicusand the other taxa discussedabove do indeed all belong to the same subfamily,then Udamoselinae is the older name that wouldthen take precedence over Aleurodicinae. Solomon(1935) did express the opinion that Aleurodicinaeand Udamoselinae were one and the same, but thenincorrectly used the more recent name, Aleurodici-nae, as valid.Shcherbakov (2000) has expressed the opinion that,should puparia of Udamoselisdisplay the feature ofrays and additional tracheal openings at the puparialmargin (see above, and Figs 30-32) then all such taxamight reasonably be placed in a separate tribe, Uda-moselini, and those without them in a second tribe,Aleurodicini.

    A final predictionAlthough still a speculative opinion, the authorsinterpretation of the biological evidence discussedabove does lead him to expect the following at-tributes to be displayed by Udamoselispuparia:

    They are likely to be rather cryptic (i.e. withoutcopious woolly secretions);

    They are likely to be scattered widely over theirhost(s);

    They are likely to resemble those of aleurodicinespecies with relatively cryptic puparia that haveradial rays (Figs 30-32), and possibly also under-lying tracheal folds;

    Males of the size of U. pigmentariaand U. estrel-lamarinaewill need puparia of at least 5-6 mmlong, as observed by Visnya (1941) for one prov-en male puparium of similar-sized Parudamoseliskesselyaki;

    Discovery of Udamoselispuparia will finally con-firm that Udamoselis and Aleurodicus are mem-bers of one subfamily.

    Should the above prove to be true, Udamoselinaewill then become the valid name for the numericallysmaller of only two extant whitefly subfamilies (un-less a case for exception is made to the International

    Commission for Zoological Nomenclature).

    AcknowledgementsField work in Ecuador was made possible by theenthusiastic logistical support provided by Profes-sor Giovanni Onore of the Pontificio UniversidadCatlica del Ecuador, Quito (PUCE), who was in-strumental in obtaining necessary permissons, andwho advised on collecting locations. The field work

    was carried out under Investigation Authorisation#008 IC FAU.DNBAP/MA, and export certifi-cate #0687999. The energetic field assistance andincredibly sharp eyes of Elicio Tapia (PUCE) werealso extremely valuable and directly resulted in thecollection of the sample that yielded the specimensof Udamoselisthat are the main subject of this com-munication. The author would also like to thank theInstituto Canario de Investigaciones Agrarias, LaLaguna, Spain for supporting his participation in theEcuador field project.Field work in Nicaragua, which also yielded materialof importance to these discussions, was facilitated bythe 2004 Universidad de Leon entomological expe-dition, organised by Jean-Michel Maes.All line drawings and photography are the work ofthe author, with the exception of figure 33, whichis a reproduction of plate 1 of Quaintance & Baker(1913). Scanning of line drawings was carried out byPat Hart (BMNH).My thanks are extended to Penny Gullan for hervaluable comments on an earlier draft of this paper.

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    Table 1. Measurements (in mm) of main body parameters of males of Udamoselis estrellamarinae sp. n.,U. pigmentariaEnderlein, and some other very large whiteflies.Key:L = length; W = width; abd. = abdominal; abd. L = abdominal length, including forceps in ?; ant. = antennalsegment(s); body L = body length, including forceps in?; dts = distal tarsal segment; fch = frontal cone height inrelief; fwpd = maximum dimension of fore wax plate; f+t = femur + trochanter; HT = holotype; hwpd = maximumdimension of hind wax plate; mwpd = maximum dimension of middle wax plate; PT = paratype; pts = proximal

    tarsal segment; vas = ventral abdominal spur on abdominal segment VIII.

    sp. n. HT ? sp. n. PT? 1 sp. n. PT? 2 U. pig- Parud. kessel JHM 8078 indet.? indet./

    (BMNH) (USNM) (BMNH) menta- yaki? largest Tena, JHM 7967

    ria? **** ? Ecuador Belize

    body L 8.50 - - 7.00* 6.30*** 10.5 - -fore wing L 5.00 5.20 5.00 5.50 5.00 (6.00) 3.80 3.92 3.65fore wing W 2.10 2.25 2.20 2.91** 2.70 (2.95) 1.70 2.19 2.00hind wing L 3.50 3.6 3.55 3.75 4.00 (4.55) 2.70 3.15 3.10hind wing W 1.25 1.26 1.28 1.69** 1.80 (2.00) 1.03 1.62 1.40

    abd. L 5.90 6.75 5.80 4.75* 4.50 (6.30) 8.95 5.85 2.40 [/

    ]abd. I-VII 2.10 2.40 2.10 (3.00) 2.40 1.70 -abd. VIII 0.90 1.00 0.80 (0.90) 1.65 0.70 -abd. IX 2.15 2.26 1.95 1.75 1.20 (1.55) 3.78 2.25 -fwpd 0.57 0.63 0.57 (0.55) 0.49 0.45 0.76mwpd 0.60 0.62 0.58 0.48 0.50 -hwpd 0.36 0.44 0.37 (0.41) 0.26 ? 0.71lingula 0.085 0.11 0.10 0.10 ? 0.10forceps 1.35 1.45 1.32 1.25 0.80 (1.30) 1.60 1.20 n/afore f+t 1.35, 1.35 1.47, 1.45 1.35, - 0.93 (1.12) 0.85, 0.87 0.72 0.90fore tibia 1.35, 1.37 1.49, 1.48 1.36, 1.36 1.09 (1.15) 0.82, 0.84 0.73 0.90fore pts 0.38, 0.39 0.42, 0.42 0.37, 0.38 0.37 (0.42) 0.22, 0.23 0.28 0.32fore dts 0.27, 0.29 0.30, 0.30 0.24, 0.28 0.26 (0.27) 0.25, 0.24 0.23 0.23fore claw 0.09, 0.09 0.095, 0.09 - , 0.10 (0.08) 0.09, 0.09 0.08 0.08mid. f+t 1.43, 1.43 1.55, 1.55 1.38, 1.38 1.12 (1.30) 0.95, 0.96 0.83 1.05mid. tibia 1.45, 1.43 1.52, 1.52 1.42, 1.37 1.33 (1.40) 0.91, 0.91 0.95 1.00mid. pts 0.38, 0.36 0.41, 0.40 0.39, 0.36 0.45 (0.50) 0.18, 0.20 0.29 0.33mid. dts 0.26, 0.28 0.29, 0.27 0.26, 0.26 0.27 (0.28) 0.22, 0.23 0.23 0.22mid. claw 0.09, - 0.09, 0.09 0.09, - 0.09, - 0.08 0.07hind f+t 1.55, 1.50 1.60, 1.67 1.55, 1.54 1.15 (1.30) 0.97, 0.98 1.00 1.15hind tibia 2.17, 2.17 2.18, 2.20 2.11, 2.09 1.97 (2.10) 1.29, 1.30 1.29 1.36hind pts 0.45, 0.44 0.50, 0.51 0.46, 0.46 0.82 (0.88) 0.46, 0.48 0.64 0.51hind dts 0.27, 0.27 0.28, 0.28 0.26, - 0.32 (0.32) 0.29, 0.27 0.27 0.25hind claw 0.09, 0.09 0.085, 0.09 0.09, - 0.09, - 0.08 0.07head W - - 1.00 - - 0.76fch 0.20 - - n/a ?n/a [small]ant. I 0.07 - 0.09, - 0.08 (0.08) 0.09, - 0.05 0.05ant. II 0.15 0.15, 0.16 0.15, - 0.16 (0.18) 0.14, 0.14 0.15 0.15ant. III 0.63, 0.65 0.71, 0.76 0.62, - 0.88 0.54, 0.50 0.58 0.49ant. IV 0.31, 0.26 0.28, 0.30 0.26, - 0.35 0.28, 0.28 0.27 0.31ant. V 0.12, 0.16 0.15, 0.165 0.14, - 0.28 0.18, 0.19 0.28 0.12ant. VI 0.055 0.06, 0.055 0.06, - 0.17 0.07, 0.08 0.10 0.095ant. VII 0.04 0.03, 0.035 0.03, - 0.13 0.07, 0.065 0.075 0.06urs 0.43 0.41 0.40 (0.34) 0.33 0.34 0.34vas no no no ? yes no no n/a* Enderlein did not state whether this included the forceps

    ** Calculated from Enderleins wing-length measurements, assuming outlines are accurate*** Body length measured in alcohol or in balsam, not stated if forceps included; sole male in BMNH in dissected

    condition**** Data from Visnya, 1941, with authors measurements of BMNH paratype in (bracketed italics)

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    Martin: Giant whiteflies and new Udamoselis 29

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    Received: 11 October 2006Accepted: 1 December 2006