Giant whiteflies (Sternorrhyncha, Aleyrodidae): a ...that are needed for the thorax / legs and...

IntroductionIn an account of the history of higher systematics in the Aleyrodidae, Russell (2000) stated that five whitefly subfamily names have been used for extant taxa. Of these, Uraleyrodinae Sampson & Drews (1941) was found to be synonymous with Aleyro-dinae Westwood (1840), based on a study of adult characters by Russell (1986). Takahashi (1932) had erected the subfamily Siphonaleyrodinae solely for his new species Siphonaleyrodes formosanus, which is clearly a member of the psylloid family Triozidae, and which was placed as a junior synonym of Trioza cinnamomi (Boselli, 1930) by Mound & Halsey (1978), a view with which Russell (2000) concurred. The oldest-established subfamily, Aleyrodinae, is generally accepted and regarded as well defined by adult and nymphal [puparial] characters (Gill 1990). This leaves Udamoselinae Enderlein (1909) and Aleurodicinae Quaintance & Baker (1913) whose controversial relationship is the subject of this paper.The genus Udamoselis, the species U. pigmentaria and the subfamily Udamoselinae were all proposed by Enderlein (1909), based upon his study of a

single adult male specimen. Enderlein’s specimen has subsequently never been traced, and is thought to have been lost during the upheavals of the Second World War. As well as being described from a single specimen, no satisfactory collecting locality is known and Enderlein simply gave this as ‘in all probability South America’, indicating that the specimen must have been given to him. Enderlein also included Aleurodicus Douglas (1892) in his new subfamily, without any discussion.Quaintance & Baker (1913) discussed whitefly wing venation in detail, illustrating a range of actual and theoretical patterns (Fig. 33). They proposed an-other new subfamily, Aleurodicinae, accommodat-ing Aleurodicus, Dialeurodicus Cockerell (1902), their own new genus Leonardius and Paraleyrodes Quaintance (1909), whilst continuing to accept Enderlein’s subfamily Udamoselinae for Udamoselis alone. Their reason for supporting a separate sub-family for Udamoselis was the more complex wing venation described and illustrated by Enderlein (Fig. 7), but the insect’s enormous size (Table 1) may well have also been a factor in their decision. The

Giant whiteflies (Sternorrhyncha, Aleyrodidae):a discussion of their taxonomic and evolutionary significance, with the description of a new species of Udamoselis Enderlein from Ecuador

Jon H. Martin

Three adult male whitefly specimens from Ecuador are described as Udamoselis estrellamarinae sp. n. This genus and its subfamily are reappraised on adult characters, including wing venation, paronychium structure, and distribution of abdominal wax glands. In the absence of associated puparia nomenclatural caution is preferred, but the subfamilies Udamoselinae and Aleurodicinae are likely to be synonymous. Wing venation of other very large whiteflies is illustrated, and is discussed in comparison with fossil taxa. Speculation is made on the possible biology of such giant whitefly species.Jon H. Martin, Department of Entomology, Natural History Museum, Cromwell Road, London SW7 5BD, UK. j. [email protected]

Tijdschrift voor Entomologie 150: 13–29, Figs. 1–33, Table 1. [ISSN 0040-7496]. http://www.nev.nl/tve© 2007 Nederlandse Entomologische Vereniging. Published 1 June 2007.

Tijdschrift voor Entomologie, volume 150, 200714

relative complexity of the wing venation and great body size, in turn, have been regarded as possible evi-dence that Udamoselis might be a particularly primi-tive whitefly, and thus form a link between the other present-day taxa and species described from the fossil record (such as by Schlee 1970, Shcherbakov 2000, and Hamilton 1990).U. pigmentaria was regarded as a nomen dubium by Mound & Halsey (1978: 250), on the basis of in-adequate description of the adult, combined with the absence of knowledge of the puparial stage upon which most whitefly taxonomy is now based. Ender-lein’s description of the solitary male did indeed omit mention of some characters that are now thought likely to be important in the systematics of adults, and this supported the proposal that it be regarded as nomen dubium. Perhaps most importantly, End-erlein’s description and illustration of the fore and hind wings (see Fig. 7, here) showed a venation that is considerably more complex than had been seen in any other known extant whiteflies: this raised a ques-tion as to whether all the firm lines in Enderlein’s drawings were truly veins and, hence, whether his illustrations were accurate.Leaving aside the uncertainty over wing venation, the absence of any detail of such characters as ab-dominal wax glands or tarsal paronychium, the loss of the antennal flagellum in his sole specimen, and the lack of optical resolution available to Enderlein [he stated that no empodial paronychium was vis-ible with his ‘magnifying glass’], his description was nonetheless remarkably detailed if sometimes rather ambiguous. With the considerable importance of this taxon Quaintance & Baker (1913) provided a complete English translation of Enderlein’s descrip-tion of U. pigmentaria and this translation has been extensively consulted in the course of the present study. In the absence of study material, Mound and Halsey’s (1978) decision to regard U. pigmentaria as nomen dubium was pragmatic, allowing the continu-ing use of Aleurodicinae as the name for the numeri-cally smaller of only two extant whitefly subfamilies, accommodating about eight percent of described whiteflies.Schlee (1970) stated that ‘The systematic position of Udamoselis within the Aleyrodina cannot be elabo-rated until a new find is made, because of the insuffi-cient present knowledge based upon the single specimen, which has probably been destroyed. The assumed close kinship relation between Udamoselis and the Aleu-rodicidae [i.e. Aleurodicinae] is unproved’. In contrast, Shcherbakov (2000) said: ‘…. despite an incomplete knowledge of the type genus, the name Udamoselinae should be used in the broad sense of Enderlein (1909) and Sampson (1943), i.e. including Aleurodicinae.’

Three new specimens recently collected in Ecuador correlate with Enderlein’s description sufficiently well to be regarded as belonging to Udamoselis, thus al-lowing this intriguing controversy to be reappraised. Many of the attributes described by Enderlein for U. pigmentaria are apparently accurate, although other parts of his description remain ambiguous through the absence of the original specimen, com-bined with Enderlein’s failure to provide any illus-trations beyond the wings. Nevertheless the author now considers it quite likely that U. pigmentaria it-self will prove to be identifiable, in the event of new material becoming available, and its identity should no longer be regarded as nomen dubium. However, the subfamilial position of Udamoselis remains some-what uncertain, as will be discussed later in this paper.Examination of the three males from Ecuador has revealed their wings (Figs 5, 6) to display the identi-cal venation illustrated by Enderlein (Fig. 7), but has confirmed that not all veins are as distinct as implied by Enderlein’s simplified line drawings. Comparison of the Ecuadorean material with the description of U. pigmentaria leads to the conclusion that the two taxa are congeners but are distinct species. Despite the small sample size, and frustrating lack of females and (especially) of puparia, it is felt that naming the Ecuadorean species is valid because of the wider in-terest in higher systematics that these specimens are likely to generate. Udamoselis estrellamarinae is there-fore here described, and is named for its discoverer (see below).

Materials, methods and terminology

Background In 2005 the author visited Ecuador, in company with Dra Estrella Hernández-Suarez and Sr Elicio Tapia. The purpose was to search for whitefly col-onies that might yield natural enemies of the pest species, Lecanoideus floccissimus Martin, Hernán-dez-Suárez & Carnero, 1997, in connection with achieving its natural control in the Canary Islands. Whilst sorting collected material for possible rearing of parasitoids, Hernández-Suarez noticed three very large, darkly-pigmented and relatively wax-free adult male whiteflies inside a bag containing a substantial colony of Lecanoideus mirabilis (Cockerell, 1898) on Annona leaves. An extensive search of other bags of material from the same garden tree failed to re-veal any additional specimens. The three specimens were brought back to the laboratory at the Natural History Museum, London, for further study.

Martin: Giant whiteflies and new Udamoselis 15

Treatment of adult whiteflies for examination It is not easy to dissect any adult whitefly for mi-croscopic examination on slides, in contrast with the situation with Psylloidea (jumping plant lice) which are routinely dissected prior to slide-display. Never-theless, the interest in wing venation of larger white-flies leads the author now to remove all wings from such specimens, prior to maceration of the body, and to mount the completely untreated wings under a separate coverslip. This avoids distortion and dete-rioration of any subtle wing pigmentation, and the separate coverslip allows for a very thin (hence flat) mount for the wings.With very large taxa a partial dissection of the body also assists in better displaying certain characters for study, but with very small sample sizes this inevita-bly results in some other characters being distorted or obscured. The author usually dissects the head (sometimes with the rostrum still attached) for exam-ination frontally, the abdomen for lateral mounting, male forceps for dorso-ventral display, and the aedea-gus for lateral examination. The thorax, with legs still attached, may be mounted laterally (Fig. 1) for the practical reason that whitefly coxae are so strongly ventrally-directed that normal dorso-ventral display is virtually impossible. Alternatively, the specimen may be more extensively dissected (Fig. 4), allow-ing more constituent parts to be displayed parallel to the slide surface. With the much thicker mounts that are needed for the thorax / legs and abdomen, in comparison with that required for the genitalia and head, the constituent parts of the specimen may be distributed between two to four coverslips of 10 mm or 13 mm diameter (Fig. 4), but always with all the parts of one adult on a single slide. The practice of dividing the constituents of one individual between two slides, sometimes used by workers on Psylloidea, is to be avoided.When adult whiteflies are not dissected, the favoured alignment is generally lateral, with wings dorsally (as in the resting position of Rhopalocera) and with legs and rostrum displayed ventrally as seen in Fig. 1. With much smaller adults, usually members of the Aleyrodinae, displaying them is more challeng-ing, because manipulation requires even more care and osmotic collapse becomes a greater risk. When a large sample is at hand, the author usually resorts to placing many individuals under a single coverslip, with minimal effort being expended on each indi-vidual: the result is that usually all characters may be seen, but only by examining several specimens. With very small adults, better results are sometimes achieved by clearing and dehydrating un-macerated specimens and placing them directly onto a slide.

Wing venation terminologyThe wing venation of Udamoselis pigmentaria is important for the discussion of its genus and other larger whiteflies. Enderlein (1909) employed a termi-nology (Fig. 7) that is not accepted today, but there appears to be little consensus on which alternative to use. Accordingly, for this communication I have followed the system used by Quaintance & Baker (1913; see Fig. 33, here), Solomon (1935) and Gill (1990). For Udamoselis this is shown in figures 5 and 6, where ‘?’ indicates my uncertainty over whether the posteriormost feature on each wing is truly a vein. This venation terminology is also used here in discussion of the wings of several other large whitefly species (Figs 19-28).

Depositories BMNH The Natural History Museum, London,

U.K.USNM Entomological collections of the U.S. Na-

tional Museum of Natural History, housed at U.S. Department of Agriculture, Belts-ville, Maryland, U.S.A.

The holotype and paratype 2 of U. estrellamarinae are deposited in BMNH; paratype 1 of U. estrellama-rinae is deposited in USNM, and all three specimens have been slide-mounted as described above, with varying degrees of dissection. All the other specimens discussed and illustrated here are also housed in the collection of BMNH.

Udamoselis EnderleinUdamoselis Enderlein, 1909: 230. Type species: Udamoselis

pigmentaria Enderlein, by monotypy, but sole original specimen not traced.

Udamoselis estrellamarinae sp. n.Figs 1–6, 8–18

Type material. Holotype: adult ?, Ecuador, Manabí Province coast, Briseña, 5 km north of San Vicente, on Annona sp., probably A. muricata (Annonaceae), 10.ii.2005 (Hernández-Suarez coll., Martin ref. #8103) (BMNH). Paratypes. adult ?: same data as holotype (BMNH; USNM).

Adult male (n=3)Measurements. For basic measurements of many body parameters, see Table 1.Coloration. Body colour very dark, with little ap-parent waxy bloom covering the cuticle, resembling small Psylloidea to the naked eye. Pigmentation of the body cuticle as shown in the photographs (Figs 1, 4), but is generally evenly dark brown; the


middle parts of the femora and the whole of the tibiae, tarsal segments and antennal flagellar seg-ments are paler. The wings are rather leathery and their dark pigmentation (Figs 2–3) can best be de-scribed as blotchy; each fore wing has two ovoid pale patches, and each hind wing a single trapezoidal pale zone; the degree of darkening of the wings varies be-tween the three specimens, with the holotype (Figs 2–3) the darkest.Legs. Femoral hairs normal on all legs, bristle-like; many hairs on distal parts of tibiae thickened but still apically acute; each middle tibia with a group of thickened hairs forming a distinct comb at three-quarters length, each hind tibia with a more subtle comb formed by a smaller number of thickened hairs, but each fore tibia without such a comb of hairs; each hind tibia with an apical arc of 9–11 pale dagger-like spines (Fig. 18) that may be saltatorial in function (as in Psylloidea), but these are absent from other legs. Tarsi two-segmented, basal segment much longer than apical segment; hairs on basal segment stouter and more numerous than those on apical segment; hind basal tarsal segment with two rows of stout ventral spines (Fig. 17), which (like the api-cal hind tibial spines) may be saltatorial in function; claws paired, each pair with a single stout ventrally- directed spine-like paronychium between them (Fig. 9). Tibial and tarsal surfaces with very fine

spinules, becoming dense enough on tarsi to be re-garded as a fine pubescence.Abdomen. With a pronounced furrow running along each side of segments III–VII, above the mid-line; segment VIII about as long as deep, slightly ex-panded at distal end, without a ventral spur at its distal extremity; segment IX about 3 times as long as deep; sternites III–V each with a pair of wax plates (Fig. 14), the anterior two pairs significantly larger than the posterior pair; each anterior wax plate with a deep invagination from its anterior side, lined with hairs and accommodating a patch of rounded reticu-lations; middle wax plates with a similar reticulate patch situated in an indentation of the anterior edge of the plate; hind wax plates evenly ovoid, without indentations and not fringed with hairs; surfaces of wax plates extremely finely imbricate-reticulate (400 magnification), with a few hairs on their sur-faces as shown in figure 14; segments VIII–IX devoid of hairs but other segments with hairs surrounding wax plates (Fig. 14) and near postero-ventral margins of sternites; tergites III–IV each with a subcircular patch of rounded reticulations on each side (Fig. 16); whole of abdomen punctuated by tiny pale ovoid pores (Figs 14–16). Lingula and presumed oper-culum (Fig. 15) situated between one-third and half way from base to apex of segment IX; lingula with fine spinule-pubescence but no evident hairs;

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

1

2

4

3


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

antennal flagellum (Fig. 11) without hairs but with finely spinulose transverse striations throughout its length (Fig. 11, expanded detail); flagellar segments with many extremely small, pale spots that are likely to be sensoria, with a few slightly larger sensoria on the apical segments; ultimate rostral segment pigmented dark brown in apical third, with many hairs.Thorax. Thoracic dorsal plates smooth, pigmented brown and sparsely provided with fine hairs, except for a fringe of longer hairs anteriorly on prothorax.Wings. Rather narrow, dimensions given in Table 1, fore wing margin flattened to slightly emarginate op-posite apices of veins Rs and M (Fig. 5). Pterostigma well defined by surface roughening (Fig. 5) rather than by differentiated coloration (Fig. 2). Venation of fore wing as shown in figure 5; Sc contiguous with C until diverging as it approaches the pterostigma; R1 short, becoming indistinct in the pterostigma; Rs almost reaching wing margin; M very long,

CSc

R1

Rs

M

Cu?A

R

R1

Rs

67

5

R

r1 rm

cuanax r

cu

an m?Cu

Figs 5–7. Wings of Udamoselis species. – 5, U. estrellamarinae sp. n., fore wing with venation annotated using ter-minology adopted here; 6, U. estrellamarinae sp.n., hind wing with venation annotated using terminology adopted here; 7, U. pigmentaria, original figure from Enderlein (1909), showing Enderlein’s venation terminology. [Scale bars = 1 mm]


9

b

13

a

8

10 11

14

12

c

15

Figs 8–15. 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 III–V, holotype; 15, lateral aspect of operculum (left) and lingula (right), paratype 1.


16 17

18

19

20

21

22

Figs 16–22. – 16–18, Udamoselis estrellamarinae. 16, oval patches of reticulations on abdominal tergites III and IV, paratype 2; 17, distal half of hind basitarsus, paratype 1; 18, apex of hind tibia, showing fringe of spines and thickened setae proximad, paratype 1. 19–20. Wings of male syntype of Parudamoselis kesselyaki Visnya. 19, fore wing; 20, hind wing. 21–22. 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]


23

24

25

26

27

28

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


diverging from R near wing base, almost straight but slightly angled posteriorly in its apical quarter; Cu (see discussion, below) faintly-indicated basally and apically, parallel to and closest to M at half-length, markedly paler than the heavily pigmented wing surface; posterior to Cu is a subtle and irregular fold that is clearly what Enderlein regarded as the Axillaris [anal vein, here], but it is unclear whether this really warrants such status. Hind wing with R / R1 & Rs clearly marked, with M diverging from R at wing base; in a similar situation to that seen in the fore wing, a subtle thickening / fold indicates what Enderlein regarded as the Analis [vein Cu here]. Coloration and patterning of wings as shown in Figs 2–3.Whichever fore wing venation system is followed the entity named Cu (Fig. 5) is clearly associated with the claval furrow (sutura clavi of Enderlein), always almost straight for much of its length and distinctly pale on pigmented wings (a ‘very distinct bright line’ as described by Enderlein for his fore wing Analis in figure 7). In Psylloidea, the same observations are true for vein Cu2 which doubles as the clavus (Hod-kinson & White 1979). Indeed, the present author questions whether this is really a ‘vein’ at all.Remarks. U. estrellamarinae differs from U. pig-mentaria in some obvious respects, assuming End-erlein’s description of the latter to be accurate. The wings of U. estrellamarinae are more elongate in shape, clearly seen by comparing figures 5-6 with figure 7, and by comparing the parameters given in Table 1. The margin of the fore wing of U. estrel-lamarinae is distinctly emarginate opposite the apex of vein Rs, and slightly less so opposite the apex of M, in contrast to the evenly curved distal margin in U. pigmentaria. Enderlein described the hind wing of U. pigmentaria as ‘densely sprinkled with dark brown spots’, in contrast to the almost uniformly dark hind wing, punctuated by tiny pale spots (Fig. 3), in U. estrellamarinae: otherwise, the descrip-tion of the wing pigmentation in U. pigmentaria is torturous and demonstrates well why photographs are the best way to illustrate such characters.Despite the wings of U. pigmentaria being somewhat longer, and much broader than in U. estrellamarinae the probability is that the body of U. pigmentaria is smaller than in U. estrellamarinae, with abdominal segment IX (see Table 1) being a particularly clear indicator despite the ambigu-ity over whether Enderlein’s stated body length for U. pigmentaria was inclusive or exclusive of the forceps.The conical protuberance on the frons was described as black in U. pigmentaria but is concolorous with the remainder of the head in U. estrellamarinae.

Giant size and wing venation in the AleyrodidaeFrom the description and discussion, above, Uda-moselis is clearly a genus with exceptionally large males. With no females available for study, it is by no means certain that females will prove to be as large. There is a good reason for sounding such a caution-ary note (see below).Figures 19 and 20 show the fore and hind wings of the sole syntype (an adult male) of Parudamoselis kesselyaki Visnya (1941) in BMNH. Described as a ‘gigantic’ whitefly by Visnya (from a large introduced population in a Hungarian glasshouse), the BMNH male is actually even larger than the Udamoselis males (see Table 1), but the body cuticle and wing colora-tion are more typical for a whitefly, not being darkly pigmented. Although P. kesselyaki is now regarded as a junior synonym of Ceraleurodicus varus (Bondar, 1928), based on comparison of syntype puparia, (see below) of both species in the USNM collection (Martin et al. 2000), the discussions here are based upon the BMNH adult male syntype of P. kesselyaki and Visnya’s observations on the Hungarian mate-rial from the 1940s; no other males are known at the present time. The fore wing of the BMNH syn-type of P. kesselyaki (6.00 mm long, 2.95 mm maxi-mum width) displays the same much-curtailed R1, terminating in a pronounced pterostigma, which is also seen in Udamoselis. There is also a suggestion that thickening of the proximal boundary of the pterostigma could be the equivalent of the putative vein Sc in Udamoselis. As in the males of both spe-cies of Udamoselis and other large species discussed here, the result is a more robust fore wing. Visnya also remarked upon distinct adult size-dimorphism in P. kesselyaki. Four females reared from puparia on Protium copal in Belize, and identified as C. varus from their puparia, are considerably smaller than the only (male) syntype of P. kesselyaki in BMNH, and also smaller than Visnya’s quoted measurements for females of P. kesselyaki.Visnya further noted that the male puparia of P. kesselyaki were significantly larger than those of fe-males, as follows: ‘The length of most L4 is between 3.8 and 4.3 mm. These are all females and the well devel-oped eggs can be seen in matured ones before emerging. There are L4 of 5 mm and above, these seem to be males, so far however I could only find one preparation with a larva of 5.3 mm length and 2.5 mm wide, in which the penis and claspers are visible.’[Author’s note: the observations below concern some undescribed whitefly taxa. The characteristics de-tailed here are not intended to be for nomenclatu-ral purposes, but only to broaden this discussion topic.]


In 2004 a congregation of extraordinary adult white-flies was discovered under a single leaf in Nicaragua [Uncaria tomentosa (Rubiaceae), Rio San Juan / Rio Bartola confluence, 22.vi.2004, Martin #8078]. The generic position of this undescribed species is un-certain, because almost nothing is known about the relative importance of different adult characters in whitefly generic definitions. This situation has arisen because for 150 years whitefly taxonomy has been based almost solely upon puparial [i.e. final nymphal stage] characters (Martin 2003). This species is here referred to simply by its collection number, JMH 8078, and it is notable for two reasons. Firstly, one of the five males (Fig. 29) may be the largest individual whitefly ever collected, with an overall body length of over 1 cm (see Table 1). Despite this exceptional length, the body is possibly less massive than the males of Udamoselis, with the extreme length being due to abdominal segments VIII and IX (Fig. 29). At 3.78 mm long in this largest specimen, its ab-dominal segment IX alone is longer than almost all other whitefly species, including those in the Aleuro-dicinae. However, the fore wings of these males (3.80 mm long in the largest specimen, down to 3.20 mm

in the smallest) are considerably smaller than those of Udamoselis males, presumably reflecting a lower body mass.JMH 8078 is also notable for extreme adult dimor-phism (Fig. 29). The adult females (body length 2.10–2.60 mm, fore wing length 1.75–2.20 mm, n=7) are all very much smaller than the males, and are similar in size to many other Aleurodicinae. The difference in wing size, between males and females of JMH 8078, is at once obvious from comparison of figures 23–24 with 25–26 (all shown to the same scale). Size apart, while the female fore wing displays an entirely typical aleurodicine venation, that of the male has vein R1 converging with the costal margin, there forming a small pterostigma, and then contin-uing towards the wing apex and almost paralleling the curve of Rs. The increased complexity of the fore wing venation in the male results in a more robust structure.Figures 21 and 22 show the fore and hind wings of a single large (dissected) male specimen (fore wing 3.92 mm long, 2.19 mm maximum width) from Ecuador [Tena, 23.ii.1923, F.X. Williams coll.], which also remains unplaced generically. Here, R1 is

Fig. 29. Unidentified whitefly (JMH 8078) from Bartola, Nicaragua – largest male specimen (left) and a female specimen (right) from same collection, in alcohol prior to slide-preparation, showing extreme dimorphism.


Figs 30–32. Likely models for Udamoselis puparia – habitus photographs of puparia of species of Aleurodicinae with sparse wax, radial rays and additional ventral tracheal folds, that do not occur in aggregations. – 30, Ceraleurodicus keris Martin, on Lunania parviflora, Nicaragua; 31, Octaleurodicus sp., on ?Melastomataceae, Ecuador; 32, Ceraleu-rodicus sp., on Clusia grandiflora, Guyana.

30

3132


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


reminiscent of JMH 8078 but, instead of a pterostigma, there is a curious fusion with the putative Sc. Also, M is broken at its proximal end, with the basal ‘stub’ displaced towards Cu (this curious fea-ture is identical in both fore wings). Once again, the result is a more robust structure, possibly to support a larger-than-usual body mass.Although very large males are sometimes associated with much smaller females (JMH 8078 and P. kes-selyaki [= C. varus], see above), there are also very large female whiteflies in existence. One such female [Chiococca alba (Rubiaceae), Belize, Chiquibul forest Reserve, 01.vi.2004, Martin #7967], also of uncer-tain generic position, has fore wings 3.65 mm long, 2.00 mm maximum width and hind wings 3.10 mm long, 1.40 mm maximum width. The abdomen has four pairs of wax glands, the posteriormost two pairs with coarsely reticulate facets (which are reminiscent of those in Aleurodicus dispersus). The body, is not dark brown and the wings have only isolated spots of pigment, so it is not thought likely that this individ-ual will prove to be a species of Udamoselis. Despite the large size of the wings of this female, the venation is typically aleurodicine, with R widely-forked and no pterostigma.Finally, figures 27 and 28 show the wings of the male of Dialeurodicus caballeroi Martin, 2004, a much smaller species than those discussed above (fore wing 1.83 mm long, 0.86 mm maximum width). This is a wing type commonly seen in aleurodicine species of medium size, with R1 simply but well developed in the fore wing, similar to that seen in the hind wing, but contiguous with a large pterostigma in the fore wing. There is no indication of size dimorphism in D. caballeroi, with female fore wings even slightly longer and broader than in most males. [The pter-ostigma in both sexes is much less distinct in speci-mens whose wings have been through the macera-tion process prior to slide-mounting.]In all the taxa discussed above, the hind wings have the putative Cu either short and poorly indicated, or apparently absent, and are thus typical of aleurodi-cines. The only oddity amongst these hind wings is seen in the male syntype of P. kesselyaki (Fig. 20), where R1 has a curious (?vein) stub arising at its half-length, a feature seen in both of the hind wings.

Observations on wings and other characters, with respect to subfamily placement in extant taxaAdded to the detailed comments on the very large taxa, above, the following broader observations may be made:• Almost all members of the Aleurodicinae have

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

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

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

• A few unusually large members of the Aleyrodi-nae have their fore wings with R forked, or have other complications to their venation. Examples include European species of Aleurochiton Tull-gren, 1907 and the northern Australian native, Gagudjuia allosyncarpiae Martin, 1999 and at least one species of Aleuroparadoxus Quaintance & Baker, 1914 [an undescribed species from Guyana].

• All the males discussed here, including Udamose-lis and JMH 8078, have three pairs of abdominal wax plates – normally regarded as a diagnostic aleurodicine feature (Gill, 1990).

• All the males discussed here, including Uda-moselis and JMH 8078, have a spine- or seta-like paronychium, again regarded as a diagnostic aleurodicine feature.

• Size dimorphism is also quite common in the Aleyrodinae but it is then usually the adult males, and male puparia, that are smaller. In gen-era such as Aleurocanthus, the male and female puparia have not infrequently been described as separate species.

The fossil dimensionSchlee (1970) presented a detailed discussion of Cre-taceous and Tertiary amber-fossilised whiteflies. As part of his comparisons of these with extant taxa, he said the following:‘The recent Udamoselis is by no means the ‘most primi-tive’ recent Aleyrodid; it does not represent the Aley-rodina’s ground plan. Moreover it is one of the highly derivative forms, exhibiting numerous autapomorphies (one of which is the enormous size). The relatively com-plete and distinct venation is connected with the large body size.’ He further stated that ‘The wing venation gives no evidence for the kinship relations within the Aleyrodina; families must not be defined by this fea-ture.’ These views are supported by the quoted small-to-medium size of the following fossil taxa: Heidea cretacica Schlee, 1970, body length 1.02 mm, fore wing length 0.83 mm; Bernaea neocomica Schlee,


1970, body length 1.30 mm, fore wing length 1.08 mm; Juleyrodes gilli Shcherbakov, 2000, fore wing length 2.1 mm (no associated body); Burmoselis evelynae Shcherbakov, 2000, body length 0.95 mm, fore wing length 1.10 mm. The wing venation of H. cretacica was not discernible, but the venations of the other three species vary from similar to present-day Aleurodicinae to slightly more complex than in Udamoselis. Another fossil species, Megaleurodes megocellata Hamilton, 1990 [Cretaceous], measured 11.5 mm in length but is clear to the present au-thor that Shcherbakov (2000) has already correctly regarded this particular insect as being a member of the Fulgoroidea (Auchenorrhyncha), having three-segmented tarsi and legs that would be characteristi-cally angular in cross-section.These limited data do not, of course, preclude the possibility that very large whiteflies have occurred in the fossil record. Nevertheless these observations combine to suggest that there has been an evolu-tionary trend towards the extremely simplified wing venation seen in over 90 percent of extant whitefly species, and that this trend may have been reversed for larger species, with a more complex venation re-tained or re-evolving in the cases of Udamoselis, most members of the Aleurodicinae and some larger mem-bers of the Aleyrodinae.

Observations on biologyAll specimens of the very large taxa, and specimens of several other larger species, are represented in the BMNH collection only by small numbers of adults, these sometimes having been discovered in apparent mating congregations with no associated puparia. A personal observation, that the author be-lieves to be correlated, is that puparia of several large aleurodicine species currently placed in Ceraleu- rodicus Hempel, 1922 are found widely scattered over their hosts, and not in the kind of aggregations that are typical for most other species in the Aleu-rodicinae. It is considered possible that the adults of such species congregate for mating, but that the females then distribute single eggs very widely. If this is the strategy also adopted by Udamoselis then it is likely that the three individuals of U. estrellamarinae were a nucleus for such a mating group, and it is also likely that puparia will be difficult to find.Visnya’s comments on male puparia of P. kes-selyaki concur with the present author’s field ob-servations of this whitefly group (see description of Ceraleurodicus keris and account of C. varus by Martin 2004) – that males are even more difficult to find than are females, and no males or probable male puparia of either C. varus or C. keris have been

discovered despite detailed searches.A characteristic of the puparia of larger spe-cies currently accommodated in Ceraleurodicus, Dialeurodicus and Octaleurodicus Hempel, 1922 (Figs 30–32) is that they feature nine pairs of radial rays [‘peripheral intersegmental ridges’ of Shcherba-kov 2000]. At least some of these rays terminate in very fine ‘combs’ of modified marginal teeth, and the secreted glassy peripheral filaments are thus narrower at these points, appearing more opaque and render-ing the rays clearly visible within the glassy skirt that surrounds the puparium (Figs 30–32). Some rays in some species also have apparent tracheal folds un-derlying them ventrally (Martin 2004). Shcherbakov considered that this tracheal feature indicates a more complete complement of spiracles than is usual in whitefly puparia. Such puparia also tend to have their compound pores reduced in size and number (those entirely without compound pores being assigned to Dialeurodicus), and do not produce the copious ‘woolly’ tangles of secretions that are so commonly associated with other members of the Aleurodicinae. Three puparia of the type discussed above are shown here: figure 30 depicts Ceraleurodicus keris, with an asymmetric puparium and with a very long and com-plex filament arising from each compound pore, but otherwise highly cryptic; figure 31 shows a species of Octaleurodicus, with short and glassy ‘fingers’ of wax secreted by eight small, submedian compound pores; figure 32 features a species of Ceraleurodicus with ex-tremely reduced (submedian) compound pores, and no discernible dorsal secretions at all.One can only speculate on the size and appearance of puparia of Udamoselis, or those of other giant males such as those of JMH 8078, but Visnya’s ob-servation that the male puparia of P. kesselyaki are significantly larger than those of females is likely to be relevant. The puparia of both C. varus (recent-ly-collected material from Central America) and C. keris have been recorded as reaching 4 mm in length, but these may all be female. The widely scat-tered puparia of at least some species currently placed in Ceraleurodicus are highly cryptic in life, hinder-ing field searching. This was especially marked with C. varus in Belize (Martin 2004).A few old puparia of C. keris were discovered on the same host plant as the aggregation of adults of JMH 8078, and it remains a possibility that these two en-tities may be conspecific [no adults of C. keris have been reared from puparia].

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


mens, there is nothing that suggests to the author that Udamoselis should be regarded as belonging to a different subfamily from Aleurodicus and other members of the Aleurodicinae. This concurs with the conclusions of Solomon (1935), Sampson (1943), Schlee (1970) and Shcherbakov (2000), as discussed by Martin & Streito (2003). The same may be said of the other large taxa with relatively complex wing venation, discussed here. In large part, this conclu-sion is based upon the proposition that more com-plex wing venation is a response to larger body mass (as promulgated by Schlee 1970), and is thus not a feature of importance for subfamilial placement. This proposition is made because other major at-tributes are entirely typical for Aleurodicinae.It therefore follows that there is no compelling rea-son for regarding Udamoselis as providing evidence of a ‘missing link’ between fossil whitefly taxa and those alive today. It is of particular note that at least some fossil taxa were not giants (see above), despite their more complex wing venations, leading Schlee to conclude that Udamoselis is actually highly derived, rather than plesiomorphic, in some of its features.However, the continuing (and frustrating) lack of adult females and (especially) immature stages, reli-ably associated with males of Udamoselis, prevents a final conclusion from being drawn here on its subfa-milial position. Should future collecting yield females and puparia, which further support the view that Udamoselis, Aleurodicus and the other taxa discussed above do indeed all belong to the same subfamily, then Udamoselinae is the older name that would then take precedence over Aleurodicinae. Solomon (1935) did express the opinion that Aleurodicinae and Udamoselinae were one and the same, but then incorrectly used the more recent name, Aleurodici-nae, as valid.Shcherbakov (2000) has expressed the opinion that, should puparia of Udamoselis display the feature of rays and additional tracheal openings at the puparial margin (see above, and Figs 30-32) then all such taxa might reasonably be placed in a separate tribe, Uda-moselini, and those without them in a second tribe, Aleurodicini.

A final predictionAlthough still a speculative opinion, the author’s interpretation of the biological evidence discussed above does lead him to expect the following at-tributes to be displayed by Udamoselis puparia:• They are likely to be rather cryptic (i.e. without

copious woolly secretions);• They are likely to be scattered widely over their

host(s);

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

• Males of the size of U. pigmentaria and U. estrel-lamarinae will need puparia of at least 5-6 mm long, as observed by Visnya (1941) for one prov-en male puparium of similar-sized Parudamoselis kesselyaki;

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

Should the above prove to be true, Udamoselinae will then become the valid name for the numerically smaller of only two extant whitefly subfamilies (un-less a case for exception is made to the International Commission for Zoological Nomenclature).

AcknowledgementsField work in Ecuador was made possible by the enthusiastic logistical support provided by Profes-sor Giovanni Onore of the Pontificio Universidad Católica del Ecuador, Quito (PUCE), who was in-strumental in obtaining necessary permissons, and who advised on collecting locations. The field work was carried out under Investigation Authorisation #008 – IC – FAU.DNBAP/MA, and export certifi-cate #0687999. The energetic field assistance and incredibly sharp eyes of Elicio Tapia (PUCE) were also extremely valuable and directly resulted in the collection of the sample that yielded the specimens of Udamoselis that are the main subject of this com-munication. The author would also like to thank the Instituto Canario de Investigaciones Agrarias, La Laguna, Spain for supporting his participation in the Ecuador field project.Field work in Nicaragua, which also yielded material of importance to these discussions, was facilitated by the 2004 Universidad de Leon entomological expe-dition, organised by Jean-Michel Maes.All line drawings and photography are the work of the author, with the exception of figure 33, which is a reproduction of plate 1 of Quaintance & Baker (1913). Scanning of line drawings was carried out by Pat Hart (BMNH).My thanks are extended to Penny Gullan for her valuable comments on an earlier draft of this paper.


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

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

(BMNH) (USNM) (BMNH) menta- yaki ? largest Tena, JHM 7967 ria ? **** ? Ecuador Belize

body L 8.50 - - 7.00* 6.30*** 10.5 - -fore wing L 5.00 5.20 5.00 5.50 5.00 (6.00) 3.80 3.92 3.65fore 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 1.40abd. 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 Enderlein’s 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 author’s measurements of BMNH paratype in (bracketed italics)


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Received: 11 October 2006Accepted: 1 December 2006

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