Giant Whiteflies (Sternorrhyncha, Aleyrodidae): a Discussion of Their Taxonomic and Evolutionary Significance, with the Descript

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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]

Introduction single adult male specimen. Enderlein’s specimen has In an account of the history of higher systematics subsequently never been traced, and is thought to in the Aleyrodidae, Russell (2000) stated that five have been lost during the upheavals of the Second whitefly subfamily names have been used for extant World War. As well as being described from a single taxa. Of these, Uraleyrodinae Sampson & Drews specimen, no satisfactory collecting locality is known (1941) was found to be synonymous with Aleyro- and Enderlein simply gave this as ‘in all probability dinae Westwood (1840), based on a study of adult South America’, indicating that the specimen must characters by Russell (1986). Takahashi (1932) had have been given to him. Enderlein also included erected the subfamily Siphonaleyrodinae solely for Aleurodicus Douglas (1892) in his new subfamily, his new species Siphonaleyrodes formosanus, which without any discussion. is clearly a member of the psylloid family Triozidae, Quaintance & Baker (1913) discussed whitefly wing and which was placed as a junior synonym of Trioza venation in detail, illustrating a range of actual and cinnamomi (Boselli, 1930) by Mound & Halsey theoretical patterns (Fig. 33). They proposed an- (1978), a view with which Russell (2000) concurred. other new subfamily, Aleurodicinae, accommodat- The oldest-established subfamily, Aleyrodinae, is ing Aleurodicus, Dialeurodicus Cockerell (1902), generally accepted and regarded as well defined by their own new genus Leonardius and Paraleyrodes adult and nymphal [puparial] characters (Gill 1990). Quaintance (1909), whilst continuing to accept This leaves Udamoselinae Enderlein (1909) and Enderlein’s subfamily Udamoselinae for Udamoselis Aleurodicinae Quaintance & Baker (1913) whose alone. Their reason for supporting a separate sub- controversial relationship is the subject of this paper. family for Udamoselis was the more complex wing The genus Udamoselis, the species U. pigmentaria venation described and illustrated by Enderlein and the subfamily Udamoselinae were all proposed (Fig. 7), but the insect’s enormous size (Table 1) may by Enderlein (1909), based upon his study of a well have also been a factor in their decision. The

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relative complexity of the wing venation and great Three new specimens recently collected in Ecuador body size, in turn, have been regarded as possible evi- correlate with Enderlein’s description sufficiently well dence that Udamoselis might be a particularly primi- to be regarded as belonging to Udamoselis, thus al- tive whitefly, and thus form a link between the other lowing this intriguing controversy to be reappraised. present-day taxa and species described from the fossil Many of the attributes described by Enderlein for record (such as by Schlee 1970, Shcherbakov 2000, U. pigmentaria are apparently accurate, although and Hamilton 1990). other parts of his description remain ambiguous U. pigmentaria was regarded as a nomen dubium by through the absence of the original specimen, com- Mound & Halsey (1978: 250), on the basis of in- bined with Enderlein’s failure to provide any illus- adequate description of the adult, combined with trations beyond the wings. Nevertheless the author the absence of knowledge of the puparial stage upon now considers it quite likely that U. pigmentaria it- which most whitefly taxonomy is now based. Ender- self will prove to be identifiable, in the event of new lein’s description of the solitary male did indeed omit material becoming available, and its identity should mention of some characters that are now thought no longer be regarded as nomen dubium. However, likely to be important in the systematics of adults, the subfamilial position of Udamoselis remains some- and this supported the proposal that it be regarded what uncertain, as will be discussed later in this as nomen dubium. Perhaps most importantly, End- paper. erlein’s description and illustration of the fore and Examination of the three males from Ecuador has hind wings (see Fig. 7, here) showed a venation that revealed their wings (Figs 5, 6) to display the identi- is considerably more complex than had been seen in cal venation illustrated by Enderlein (Fig. 7), but has any other known extant whiteflies: this raised a ques- confirmed that not all veins are as distinct as implied tion as to whether all the firm lines in Enderlein’s by Enderlein’s simplified line drawings. Comparison drawings were truly veins and, hence, whether his of the Ecuadorean material with the description of illustrations were accurate. U. pigmentaria leads to the conclusion that the two Leaving aside the uncertainty over wing venation, taxa are congeners but are distinct species. Despite the absence of any detail of such characters as ab- the small sample size, and frustrating lack of females dominal wax glands or tarsal paronychium, the loss and (especially) of puparia, it is felt that naming the of the antennal flagellum in his sole specimen, and Ecuadorean species is valid because of the wider in- the lack of optical resolution available to Enderlein terest in higher systematics that these specimens are [he stated that no empodial paronychium was vis- likely to generate. Udamoselis estrellamarinae is there- ible with his ‘magnifying glass’], his description was fore here described, and is named for its discoverer nonetheless remarkably detailed if sometimes rather (see below). ambiguous. With the considerable importance of this taxon Quaintance & Baker (1913) provided a complete English translation of Enderlein’s descrip- Materials, methods and terminology tion of U. pigmentaria and this translation has been extensively consulted in the course of the present Background study. In the absence of study material, Mound and In 2005 the author visited Ecuador, in company Halsey’s (1978) decision to regard U. pigmentaria as with Dra Estrella Hernández-Suarez and Sr Elicio nomen dubium was pragmatic, allowing the continu- Tapia. The purpose was to search for whitefly col- ing use of Aleurodicinae as the name for the numeri- onies that might yield natural enemies of the pest cally smaller of only two extant whitefly subfamilies, species, Lecanoideus floccissimus Martin, Hernán- accommodating about eight percent of described dez-Suárez & Carnero, 1997, in connection with whiteflies. achieving its natural control in the Canary Islands. Schlee (1970) stated that ‘The systematic position of Whilst sorting collected material for possible rearing Udamoselis within the Aleyrodina cannot be elabo- of parasitoids, Hernández-Suarez noticed three very rated until a new find is made, because of the insuffi- large, darkly-pigmented and relatively wax-free adult cient present knowledge based upon the single specimen, male whiteflies inside a bag containing a substantial which has probably been destroyed. The assumed close colony of Lecanoideus mirabilis (Cockerell, 1898) kinship relation between Udamoselis and the Aleu- on Annona leaves. An extensive search of other bags rodicidae [i.e. Aleurodicinae] is unproved’. In contrast, of material from the same garden tree failed to re- Shcherbakov (2000) said: ‘…. despite an incomplete veal any additional specimens. The three specimens knowledge of the type genus, the name Udamoselinae were brought back to the laboratory at the Natural should be used in the broad sense of Enderlein (1909) History Museum, London, for further study. and Sampson (1943), i.e. including Aleurodicinae.’

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Treatment of adult whiteflies for examination Wing venation terminology It is not easy to dissect any adult whitefly for mi- The wing venation of Udamoselis pigmentaria is croscopic examination on slides, in contrast with the important for the discussion of its genus and other situation with Psylloidea (jumping plant lice) which larger whiteflies. Enderlein (1909) employed a termi- are routinely dissected prior to slide-display. Never- nology (Fig. 7) that is not accepted today, but there theless, the interest in wing venation of larger white- appears to be little consensus on which alternative flies leads the author now to remove all wings from to use. Accordingly, for this communication I have such specimens, prior to maceration of the body, and followed the system used by Quaintance & Baker to mount the completely untreated wings under a (1913; see Fig. 33, here), Solomon (1935) and Gill separate coverslip. This avoids distortion and dete- (1990). For Udamoselis this is shown in figures 5 and rioration of any subtle wing pigmentation, and the 6, where ‘?’ indicates my uncertainty over whether separate coverslip allows for a very thin (hence flat) the posteriormost feature on each wing is truly a mount for the wings. vein. This venation terminology is also used here in With very large taxa a partial dissection of the body discussion of the wings of several other large whitefly also assists in better displaying certain characters for species (Figs 19-28). study, but with very small sample sizes this inevita- bly results in some other characters being distorted Depositories or obscured. The author usually dissects the head BMNH The Natural History Museum, London, (sometimes with the rostrum still attached) for exam- U.K. ination frontally, the abdomen for lateral mounting, USNM Entomological collections of the U.S. Na- male forceps for dorso-ventral display, and the aedea- tional Museum of Natural History, housed gus for lateral examination. The thorax, with legs still at U.S. Department of Agriculture, Belts- attached, may be mounted laterally (Fig. 1) for the ville, Maryland, U.S.A. practical reason that whitefly coxae are so strongly The holotype and paratype 2 of U. estrellamarinae ventrally-directed that normal dorso-ventral display are deposited in BMNH; paratype 1 of U. estrellama- is virtually impossible. Alternatively, the specimen rinae is deposited in USNM, and all three specimens may be more extensively dissected (Fig. 4), allow- have been slide-mounted as described above, with ing more constituent parts to be displayed parallel varying degrees of dissection. All the other specimens to the slide surface. With the much thicker mounts discussed and illustrated here are also housed in the that are needed for the thorax / legs and abdomen, in collection of BMNH. comparison with that required for the genitalia and head, the constituent parts of the specimen may be Udamoselis Enderlein distributed between two to four coverslips of 10 mm or 13 mm diameter (Fig. 4), but always with all the Udamoselis Enderlein, 1909: 230. Type species: Udamoselis parts of one adult on a single slide. The practice of pigmentaria Enderlein, by monotypy, but sole original dividing the constituents of one individual between specimen not traced. two slides, sometimes used by workers on Psylloidea, is to be avoided. Udamoselis estrellamarinae sp. n. When adult whiteflies are not dissected, the favoured Figs 1–6, 8–18 alignment is generally lateral, with wings dorsally (as in the resting position of Rhopalocera) and with Type material. Holotype: adult ?, Ecuador, Manabí legs and rostrum displayed ventrally as seen in Fig. Province coast, Briseña, 5 km north of San Vicente, 1. With much smaller adults, usually members of on Annona sp., probably A. muricata (Annonaceae), the Aleyrodinae, displaying them is more challeng- 10.ii.2005 (Hernández-Suarez coll., Martin ref. ing, because manipulation requires even more care #8103) (BMNH). Paratypes. adult ?: same data as and osmotic collapse becomes a greater risk. When holotype (BMNH; USNM). a large sample is at hand, the author usually resorts to placing many individuals under a single coverslip, Adult male (n=3) with minimal effort being expended on each indi- Measurements. For basic measurements of many vidual: the result is that usually all characters may body parameters, see Table 1. be seen, but only by examining several specimens. Coloration. Body colour very dark, with little ap- With very small adults, better results are sometimes parent waxy bloom covering the cuticle, resembling achieved by clearing and dehydrating un-macerated small Psylloidea to the naked eye. Pigmentation specimens and placing them directly onto a slide. of the body cuticle as shown in the photographs (Figs 1, 4), but is generally evenly dark brown; the

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1 3

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.

middle parts of the femora and the whole of the spinules, becoming dense enough on tarsi to be re- tibiae, tarsal segments and antennal flagellar seg- garded as a fine pubescence. ments are paler. The wings are rather leathery and Abdomen. With a pronounced furrow running along their dark pigmentation (Figs 2–3) can best be de- each side of segments III–VII, above the mid-line; scribed as blotchy; each fore wing has two ovoid pale segment VIII about as long as deep, slightly ex- patches, and each hind wing a single trapezoidal pale panded at distal end, without a ventral spur at its zone; the degree of darkening of the wings varies be- distal extremity; segment IX about 3 times as long as tween the three specimens, with the holotype (Figs deep; sternites III–V each with a pair of wax plates 2–3) the darkest. (Fig. 14), the anterior two pairs significantly larger Legs. Femoral hairs normal on all legs, bristle-like; than the posterior pair; each anterior wax plate with many hairs on distal parts of tibiae thickened but a deep invagination from its anterior side, lined with still apically acute; each middle tibia with a group hairs and accommodating a patch of rounded reticu- of thickened hairs forming a distinct comb at three- lations; middle wax plates with a similar reticulate quarters length, each hind tibia with a more subtle patch situated in an indentation of the anterior edge comb formed by a smaller number of thickened of the plate; hind wax plates evenly ovoid, without hairs, but each fore tibia without such a comb of indentations and not fringed with hairs; surfaces hairs; each hind tibia with an apical arc of 9–11 pale of wax plates extremely finely imbricate-reticulate dagger-like spines (Fig. 18) that may be saltatorial in (400 magnification), with a few hairs on their sur- function (as in Psylloidea), but these are absent from faces as shown in figure 14; segments VIII–IX devoid other legs. Tarsi two-segmented, basal segment much of hairs but other segments with hairs surrounding longer than apical segment; hairs on basal segment wax plates (Fig. 14) and near postero-ventral margins stouter and more numerous than those on apical of sternites; tergites III–IV each with a subcircular segment; hind basal tarsal segment with two rows of patch of rounded reticulations on each side (Fig. 16); stout ventral spines (Fig. 17), which (like the api- whole of abdomen punctuated by tiny pale ovoid cal hind tibial spines) may be saltatorial in function; pores (Figs 14–16). Lingula and presumed oper- claws paired, each pair with a single stout ventrally- culum (Fig. 15) situated between one-third and directed spine-like paronychium between them half way from base to apex of segment IX; lingula (Fig. 9). Tibial and tarsal surfaces with very fine with fine spinule-pubescence but no evident hairs;

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5 C Sc

R R1

?A Cu

7 r1 r m 6

R cu 1 an R ax r

?Cu Rs an m cu

Figs 5–7. Wings of Udamoselis species. – 5, U. estrellamarinae sp. n., fore wing with venation annotated using terminology 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] presumed operculum with similar spinule-pubes- antennal flagellum (Fig. 11) without hairs but with cence but only with a single apical hair visible in finely spinulose transverse striations throughout its lateral aspect [it is to be expected that there is a pair length (Fig. 11, expanded detail); flagellar segments of such hairs]; claspers evenly curved in dorso-ven- with many extremely small, pale spots that are likely tral aspect (Fig. 10) but flat in lateral aspect (Fig. 1), to be sensoria, with a few slightly larger sensoria generally smooth and with a few fine hairs as shown on the apical segments; ultimate rostral segment in figure 10; aedeagus with a distinct elbow at half- pigmented dark brown in apical third, with many length (Fig. 12), about 0.325 mm from apex to el- hairs. bow, apically with two large and possibly four much Thorax. Thoracic dorsal plates smooth, pigmented smaller finger-like protrusions (Fig. 13). brown and sparsely provided with fine hairs, except Head. As shown in antero-ventral view in figure 8; for a fringe of longer hairs anteriorly on prothorax. small for the size of the insect (1 mm wide), much Wings. Rather narrow, dimensions given in Table 1, wider than long; compound eyes very large and in- fore wing margin flattened to slightly emarginate op- dented to accommodate the antennal insertions; posite apices of veins Rs and M (Fig. 5). Pterostigma lateral ocelli well-developed, as shown in figure 8, well defined by surface roughening (Fig. 5) rather but median ocellus not evident in any of the three than by differentiated coloration (Fig. 2). Venation mounted specimens; between the eyes is situated a of fore wing as shown in figure 5; Sc contiguous with conical protrusion from the frons; clypeus densely C until diverging as it approaches the pterostigma; hairy; basal two antennal segments dark brown, R1 short, becoming indistinct in the pterostigma; smooth and with fine hairs; the five-segmented Rs almost reaching wing margin; M very long, Downloaded from Brill.com09/30/2021 04:39:28AM via free access 18 Tijdschrift voor Entomologie, volume 150, 2007

8 9

10 11 12

b c

14 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.

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16 17

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]

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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]

Downloaded from Brill.com09/30/2021 04:39:28AM via free access Martin: Giant whiteflies and new Udamoselis 21 diverging from R near wing base, almost straight Giant size and wing venation in the but slightly angled posteriorly in its apical quarter; Aleyrodidae Cu (see discussion, below) faintly-indicated basally From the description and discussion, above, Uda- and apically, parallel to and closest to M at half- moselis is clearly a genus with exceptionally large length, markedly paler than the heavily pigmented males. With no females available for study, it is by no wing surface; posterior to Cu is a subtle and irregular means certain that females will prove to be as large. fold that is clearly what Enderlein regarded as the There is a good reason for sounding such a caution- Axillaris [anal vein, here], but it is unclear whether ary note (see below). this really warrants such status. Hind wing with R / Figures 19 and 20 show the fore and hind wings of R1 & Rs clearly marked, with M diverging from R the sole syntype (an adult male) of Parudamoselis at wing base; in a similar situation to that seen in the kesselyaki Visnya (1941) in BMNH. Described as a fore wing, a subtle thickening / fold indicates what ‘gigantic’ whitefly by Visnya (from a large introduced Enderlein regarded as the Analis [vein Cu here]. population in a Hungarian glasshouse), the BMNH Coloration and patterning of wings as shown in male is actually even larger than the Udamoselis males Figs 2–3. (see Table 1), but the body cuticle and wing colora- Whichever fore wing venation system is followed the tion are more typical for a whitefly, not being darkly entity named Cu (Fig. 5) is clearly associated with pigmented. Although P. kesselyaki is now regarded as the claval furrow (sutura clavi of Enderlein), always a junior synonym of Ceraleurodicus varus (Bondar, almost straight for much of its length and distinctly 1928), based on comparison of syntype puparia, pale on pigmented wings (a ‘very distinct bright line’ (see below) of both species in the USNM collection as described by Enderlein for his fore wing Analis (Martin et al. 2000), the discussions here are based in figure 7). In Psylloidea, the same observations are upon the BMNH adult male syntype of P. kesselyaki true for vein Cu2 which doubles as the clavus (Hod- and Visnya’s observations on the Hungarian mate- kinson & White 1979). Indeed, the present author rial from the 1940s; no other males are known at questions whether this is really a ‘vein’ at all. the present time. The fore wing of the BMNH syn- Remarks. U. estrellamarinae differs from U. pig- type of P. kesselyaki (6.00 mm long, 2.95 mm maxi- mentaria in some obvious respects, assuming End- mum width) displays the same much-curtailed R1, erlein’s description of the latter to be accurate. The terminating in a pronounced pterostigma, which is wings of U. estrellamarinae are more elongate in also seen in Udamoselis. There is also a suggestion shape, clearly seen by comparing figures 5-6 with that thickening of the proximal boundary of the figure 7, and by comparing the parameters given in pterostigma could be the equivalent of the putative Table 1. The margin of the fore wing of U. estrel- vein Sc in Udamoselis. As in the males of both spe- lamarinae is distinctly emarginate opposite the apex cies of Udamoselis and other large species discussed of vein Rs, and slightly less so opposite the apex of here, the result is a more robust fore wing. Visnya M, in contrast to the evenly curved distal margin in also remarked upon distinct adult size-dimorphism U. pigmentaria. Enderlein described the hind wing in P. kesselyaki. Four females reared from puparia on of U. pigmentaria as ‘densely sprinkled with dark Protium copal in Belize, and identified as C. varus brown spots’, in contrast to the almost uniformly from their puparia, are considerably smaller than the dark hind wing, punctuated by tiny pale spots only (male) syntype of P. kesselyaki in BMNH, and (Fig. 3), in U. estrellamarinae: otherwise, the descrip- also smaller than Visnya’s quoted measurements for tion of the wing pigmentation in U. pigmentaria is females of P. kesselyaki. torturous and demonstrates well why photographs Visnya further noted that the male puparia of are the best way to illustrate such characters. P. kesselyaki were significantly larger than those of fe- Despite the wings of U. pigmentaria being males, as follows: ‘The length of most L4 is between 3.8 somewhat longer, and much broader than in and 4.3 mm. These are all females and the well devel- U. estrellamarinae the probability is that the body of oped eggs can be seen in matured ones before emerging. U. pigmentaria is smaller than in U. estrellamarinae, There are L4 of 5 mm and above, these seem to be males, with abdominal segment IX (see Table 1) being a so far however I could only find one preparation with a particularly clear indicator despite the ambigu- larva of 5.3 mm length and 2.5 mm wide, in which the ity over whether Enderlein’s stated body length for penis and claspers are visible.’ U. pigmentaria was inclusive or exclusive of the [Author’s note: the observations below concern some forceps. undescribed whitefly taxa. The characteristics de- The conical protuberance on the frons was described tailed here are not intended to be for nomenclatu- as black in U. pigmentaria but is concolorous with ral purposes, but only to broaden this discussion the remainder of the head in U. estrellamarinae. topic.]

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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.

In 2004 a congregation of extraordinary adult white- in the smallest) are considerably smaller than those flies was discovered under a single leaf in Nicaragua of Udamoselis males, presumably reflecting a lower [Uncaria tomentosa (Rubiaceae), Rio San Juan / Rio body mass. Bartola confluence, 22.vi.2004, Martin #8078]. The JMH 8078 is also notable for extreme adult dimor- generic position of this undescribed species is un- phism (Fig. 29). The adult females (body length certain, because almost nothing is known about the 2.10–2.60 mm, fore wing length 1.75–2.20 mm, relative importance of different adult characters in n=7) are all very much smaller than the males, and whitefly generic definitions. This situation has arisen are similar in size to many other Aleurodicinae. The because for 150 years whitefly taxonomy has been difference in wing size, between males and females based almost solely upon puparial [i.e. final nymphal of JMH 8078, is at once obvious from comparison stage] characters (Martin 2003). This species is here of figures 23–24 with 25–26 (all shown to the same referred to simply by its collection number, JMH scale). Size apart, while the female fore wing displays 8078, and it is notable for two reasons. Firstly, one of an entirely typical aleurodicine venation, that of the the five males (Fig. 29) may be the largest individual male has vein R1 converging with the costal margin, whitefly ever collected, with an overall body length there forming a small pterostigma, and then contin- of over 1 cm (see Table 1). Despite this exceptional uing towards the wing apex and almost paralleling length, the body is possibly less massive than the the curve of Rs. The increased complexity of the fore males of Udamoselis, with the extreme length being wing venation in the male results in a more robust due to abdominal segments VIII and IX (Fig. 29). structure. At 3.78 mm long in this largest specimen, its ab- Figures 21 and 22 show the fore and hind wings dominal segment IX alone is longer than almost all of a single large (dissected) male specimen (fore other whitefly species, including those in the Aleuro- wing 3.92 mm long, 2.19 mm maximum width) dicinae. However, the fore wings of these males (3.80 from Ecuador [Tena, 23.ii.1923, F.X. Williams coll.], mm long in the largest specimen, down to 3.20 mm which also remains unplaced generically. Here, R1 is Downloaded from Brill.com09/30/2021 04:39:28AM via free access Martin: Giant whiteflies and new Udamoselis 23

32 31

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.

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

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reminiscent of JMH 8078 but, instead of a pte- the fore wing with R forked to form R1 and Rs, rostigma, there is a curious fusion with the putative and are larger insects than most members of the Sc. Also, M is broken at its proximal end, with the Aleyrodinae. Udamoselis also displays these at- basal ‘stub’ displaced towards Cu (this curious fea- tributes. ture is identical in both fore wings). Once again, the • Most members of the Aleyrodinae do not have a result is a more robust structure, possibly to support forked R, but only a single main vein. a larger-than-usual body mass. • The only Neotropical members of the Aleuro- Although very large males are sometimes associated dicinae that are physically very small, and thus with much smaller females (JMH 8078 and P. kes- similar in size to typical members of the Aleyro- selyaki [= C. varus], see above), there are also very dinae, are the members of the genus Paraleyrodes, large female whiteflies in existence. One such female whose fore wings have an unbranched main vein, [Chiococca alba (Rubiaceae), Belize, Chiquibul forest as is typical in the Aleyrodinae. Reserve, 01.vi.2004, Martin #7967], also of uncer- • A few unusually large members of the Aleyrodi- tain generic position, has fore wings 3.65 mm long, nae have their fore wings with R forked, or have 2.00 mm maximum width and hind wings 3.10 mm other complications to their venation. Examples long, 1.40 mm maximum width. The abdomen has include European species of Aleurochiton Tull- four pairs of wax glands, the posteriormost two pairs gren, 1907 and the northern Australian native, with coarsely reticulate facets (which are reminiscent Gagudjuia allosyncarpiae Martin, 1999 and at of those in Aleurodicus dispersus). The body, is not least one species of Aleuroparadoxus Quaintance dark brown and the wings have only isolated spots of & Baker, 1914 [an undescribed species from pigment, so it is not thought likely that this individ- Guyana]. ual will prove to be a species of Udamoselis. Despite • All the males discussed here, including Udamose- the large size of the wings of this female, the venation lis and JMH 8078, have three pairs of abdominal is typically aleurodicine, with R widely-forked and wax plates – normally regarded as a diagnostic no pterostigma. aleurodicine feature (Gill, 1990). Finally, figures 27 and 28 show the wings of the male • All the males discussed here, including Uda- of Dialeurodicus caballeroi Martin, 2004, a much moselis and JMH 8078, have a spine- or seta-like smaller species than those discussed above (fore wing paronychium, again regarded as a diagnostic 1.83 mm long, 0.86 mm maximum width). This is a aleurodicine feature. wing type commonly seen in aleurodicine species of • Size dimorphism is also quite common in the medium size, with R1 simply but well developed in Aleyrodinae but it is then usually the adult the fore wing, similar to that seen in the hind wing, males, and male puparia, that are smaller. In gen- but contiguous with a large pterostigma in the fore era such as Aleurocanthus, the male and female wing. There is no indication of size dimorphism in puparia have not infrequently been described as D. caballeroi, with female fore wings even slightly separate species. longer and broader than in most males. [The pterostigma in both sexes is much less distinct in specimens whose wings have been through the macera- The fossil dimension tion process prior to slide-mounting.] Schlee (1970) presented a detailed discussion of Cre- In all the taxa discussed above, the hind wings have taceous and Tertiary amber-fossilised whiteflies. As the putative Cu either short and poorly indicated, or part of his comparisons of these with extant taxa, he apparently absent, and are thus typical of aleurodi- said the following: cines. The only oddity amongst these hind wings is ‘The recent Udamoselis is by no means the ‘most primi- seen in the male syntype of P. kesselyaki (Fig. 20), tive’ recent Aleyrodid; it does not represent the Aley- where R1 has a curious (?vein) stub arising at its half- rodina’s ground plan. Moreover it is one of the highly length, a feature seen in both of the hind wings. derivative forms, exhibiting numerous autapomorphies (one of which is the enormous size). The relatively complete and distinct venation is connected with the large Observations on wings and other body size.’ He further stated that ‘The wing venation characters, with respect to subfamily gives no evidence for the kinship relations within the placement in extant taxa Aleyrodina; families must not be defined by this fea- Added to the detailed comments on the very large ture.’ These views are supported by the quoted small- taxa, above, the following broader observations may to-medium size of the following fossil taxa: Heidea be made: cretacica Schlee, 1970, body length 1.02 mm, fore • Almost all members of the Aleurodicinae have wing length 0.83 mm; Bernaea neocomica Schlee,

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1970, body length 1.30 mm, fore wing length 1.08 discovered despite detailed searches. mm; Juleyrodes gilli Shcherbakov, 2000, fore wing A characteristic of the puparia of larger spe- length 2.1 mm (no associated body); Burmoselis cies currently accommodated in Ceraleurodicus, evelynae Shcherbakov, 2000, body length 0.95 mm, Dialeurodicus and Octaleurodicus Hempel, 1922 fore wing length 1.10 mm. The wing venation of (Figs 30–32) is that they feature nine pairs of radial H. cretacica was not discernible, but the venations of rays [‘peripheral intersegmental ridges’ of Shcherba- the other three species vary from similar to present- kov 2000]. At least some of these rays terminate in day Aleurodicinae to slightly more complex than very fine ‘combs’ of modified marginal teeth, and the in Udamoselis. Another fossil species, Megaleurodes secreted glassy peripheral filaments are thus narrower megocellata Hamilton, 1990 [Cretaceous], measured at these points, appearing more opaque and render- 11.5 mm in length but is clear to the present au- ing the rays clearly visible within the glassy skirt that thor that Shcherbakov (2000) has already correctly surrounds the puparium (Figs 30–32). Some rays in regarded this particular insect as being a member of some species also have apparent tracheal folds un- the Fulgoroidea (Auchenorrhyncha), having three- derlying them ventrally (Martin 2004). Shcherbakov segmented tarsi and legs that would be characteristi- considered that this tracheal feature indicates a more cally angular in cross-section. complete complement of spiracles than is usual in These limited data do not, of course, preclude the whitefly puparia. Such puparia also tend to have their possibility that very large whiteflies have occurred compound pores reduced in size and number (those in the fossil record. Nevertheless these observations entirely without compound pores being assigned combine to suggest that there has been an evolu- to Dialeurodicus), and do not produce the copious tionary trend towards the extremely simplified wing ‘woolly’ tangles of secretions that are so commonly venation seen in over 90 percent of extant whitefly associated with other members of the Aleurodicinae. species, and that this trend may have been reversed Three puparia of the type discussed above are shown for larger species, with a more complex venation re- here: figure 30 depicts Ceraleurodicus keris, with an tained or re-evolving in the cases of Udamoselis, most asymmetric puparium and with a very long and com- members of the Aleurodicinae and some larger mem- plex filament arising from each compound pore, but bers of the Aleyrodinae. otherwise highly cryptic; figure 31 shows a species of Octaleurodicus, with short and glassy ‘fingers’ of wax secreted by eight small, submedian compound pores; Observations on biology figure 32 features a species of Ceraleurodicus with ex- All specimens of the very large taxa, and specimens tremely reduced (submedian) compound pores, and of several other larger species, are represented in no discernible dorsal secretions at all. the BMNH collection only by small numbers of One can only speculate on the size and appearance adults, these sometimes having been discovered in of puparia of Udamoselis, or those of other giant apparent mating congregations with no associated males such as those of JMH 8078, but Visnya’s ob- puparia. A personal observation, that the author be- servation that the male puparia of P. kesselyaki are lieves to be correlated, is that puparia of several large significantly larger than those of females is likely to aleurodicine species currently placed in Ceraleu- be relevant. The puparia of both C. varus (recent- rodicus Hempel, 1922 are found widely scattered ly-collected material from Central America) and over their hosts, and not in the kind of aggregations C. keris have been recorded as reaching 4 mm in that are typical for most other species in the Aleu- length, but these may all be female. The widely scat- rodicinae. It is considered possible that the adults tered puparia of at least some species currently placed of such species congregate for mating, but that the in Ceraleurodicus are highly cryptic in life, hinder- females then distribute single eggs very widely. If this ing field searching. This was especially marked with is the strategy also adopted by Udamoselis then it is C. varus in Belize (Martin 2004). likely that the three individuals of U. estrellamarinae A few old puparia of C. keris were discovered on the were a nucleus for such a mating group, and it is also same host plant as the aggregation of adults of JMH likely that puparia will be difficult to find. 8078, and it remains a possibility that these two en- Visnya’s comments on male puparia of P. kes- tities may be conspecific [no adults of C. keris have selyaki concur with the present author’s field ob- been reared from puparia]. servations of this whitefly group (see description of Ceraleurodicus keris and account of C. varus by Martin 2004) – that males are even more difficult Provisional conclusions to find than are females, and no males or probable Based upon all the adult male characters for male puparia of either C. varus or C. keris have been Udamoselis, now available for study on actual speci-

Downloaded from Brill.com09/30/2021 04:39:28AM via free access Martin: Giant whiteflies and new Udamoselis 27 mens, there is nothing that suggests to the author • They are likely to resemble those of aleurodicine that Udamoselis should be regarded as belonging to species with relatively cryptic puparia that have a different subfamily from Aleurodicus and other radial rays (Figs 30-32), and possibly also under- members of the Aleurodicinae. This concurs with the lying tracheal folds; conclusions of Solomon (1935), Sampson (1943), • Males of the size of U. pigmentaria and U. estrel- Schlee (1970) and Shcherbakov (2000), as discussed lamarinae will need puparia of at least 5-6 mm by Martin & Streito (2003). The same may be said long, as observed by Visnya (1941) for one prov- of the other large taxa with relatively complex wing en male puparium of similar-sized Parudamoselis venation, discussed here. In large part, this conclu- kesselyaki; sion is based upon the proposition that more com- • Discovery of Udamoselis puparia will finally con- plex wing venation is a response to larger body mass firm that Udamoselis and Aleurodicus are mem- (as promulgated by Schlee 1970), and is thus not bers of one subfamily. a feature of importance for subfamilial placement. This proposition is made because other major at- Should the above prove to be true, Udamoselinae tributes are entirely typical for Aleurodicinae. will then become the valid name for the numerically It therefore follows that there is no compelling rea- smaller of only two extant whitefly subfamilies (un- son for regarding Udamoselis as providing evidence less a case for exception is made to the International of a ‘missing link’ between fossil whitefly taxa and Commission for Zoological Nomenclature). 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, Acknowledgements rather than plesiomorphic, in some of its features. Field work in Ecuador was made possible by the However, the continuing (and frustrating) lack of enthusiastic logistical support provided by Profes- adult females and (especially) immature stages, reli- sor Giovanni Onore of the Pontificio Universidad ably associated with males of Udamoselis, prevents a Católica del Ecuador, Quito (PUCE), who was in- final conclusion from being drawn here on its subfa- strumental in obtaining necessary permissons, and milial position. Should future collecting yield females who advised on collecting locations. The field work and puparia, which further support the view that was carried out under Investigation Authorisation Udamoselis, Aleurodicus and the other taxa discussed #008 – IC – FAU.DNBAP/MA, and export certifi- above do indeed all belong to the same subfamily, cate #0687999. The energetic field assistance and then Udamoselinae is the older name that would incredibly sharp eyes of Elicio Tapia (PUCE) were then take precedence over Aleurodicinae. Solomon also extremely valuable and directly resulted in the (1935) did express the opinion that Aleurodicinae collection of the sample that yielded the specimens and Udamoselinae were one and the same, but then of Udamoselis that are the main subject of this com- incorrectly used the more recent name, Aleurodici- munication. The author would also like to thank the nae, as valid. Instituto Canario de Investigaciones Agrarias, La Shcherbakov (2000) has expressed the opinion that, Laguna, Spain for supporting his participation in the should puparia of Udamoselis display the feature of Ecuador field project. rays and additional tracheal openings at the puparial Field work in Nicaragua, which also yielded material margin (see above, and Figs 30-32) then all such taxa of importance to these discussions, was facilitated by might reasonably be placed in a separate tribe, Uda- the 2004 Universidad de Leon entomological expe- moselini, and those without them in a second tribe, dition, organised by Jean-Michel Maes. Aleurodicini. 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 A final prediction (1913). Scanning of line drawings was carried out by Although still a speculative opinion, the author’s Pat Hart (BMNH). interpretation of the biological evidence discussed My thanks are extended to Penny Gullan for her above does lead him to expect the following at- valuable comments on an earlier draft of this paper. 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);

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

sp. n. HT ? sp. n. PT ? 1 sp. n. PT ? 2 U. pig- Parud. kessel JHM 8078 indet. ? indet. / (BMNH) (USNM) (BMNH) menta- yaki ? largest Tena, JHM 7967 ria ? **** ? Ecuador Belize

body L 8.50 - - 7.00* 6.30*** 10.5 - - fore wing L 5.00 5.20 5.00 5.50 5.00 (6.00) 3.80 3.92 3.65 fore wing W 2.10 2.25 2.20 2.91** 2.70 (2.95) 1.70 2.19 2.00 hind wing L 3.50 3.6 3.55 3.75 4.00 (4.55) 2.70 3.15 3.10 hind wing W 1.25 1.26 1.28 1.69** 1.80 (2.00) 1.03 1.62 1.40 abd. L 5.90 6.75 5.80 4.75* 4.50 (6.30) 8.95 5.85 2.40 [/] abd. I-VII 2.10 2.40 2.10 (3.00) 2.40 1.70 - abd. VIII 0.90 1.00 0.80 (0.90) 1.65 0.70 - abd. IX 2.15 2.26 1.95 1.75 1.20 (1.55) 3.78 2.25 - fwpd 0.57 0.63 0.57 (0.55) 0.49 0.45 0.76 mwpd 0.60 0.62 0.58 0.48 0.50 - hwpd 0.36 0.44 0.37 (0.41) 0.26 ? 0.71 lingula 0.085 0.11 0.10 0.10 ? 0.10 forceps 1.35 1.45 1.32 1.25 0.80 (1.30) 1.60 1.20 n/a fore f+t 1.35, 1.35 1.47, 1.45 1.35, - 0.93 (1.12) 0.85, 0.87 0.72 0.90 fore tibia 1.35, 1.37 1.49, 1.48 1.36, 1.36 1.09 (1.15) 0.82, 0.84 0.73 0.90 fore pts 0.38, 0.39 0.42, 0.42 0.37, 0.38 0.37 (0.42) 0.22, 0.23 0.28 0.32 fore dts 0.27, 0.29 0.30, 0.30 0.24, 0.28 0.26 (0.27) 0.25, 0.24 0.23 0.23 fore claw 0.09, 0.09 0.095, 0.09 - , 0.10 (0.08) 0.09, 0.09 0.08 0.08 mid. f+t 1.43, 1.43 1.55, 1.55 1.38, 1.38 1.12 (1.30) 0.95, 0.96 0.83 1.05 mid. tibia 1.45, 1.43 1.52, 1.52 1.42, 1.37 1.33 (1.40) 0.91, 0.91 0.95 1.00 mid. pts 0.38, 0.36 0.41, 0.40 0.39, 0.36 0.45 (0.50) 0.18, 0.20 0.29 0.33 mid. dts 0.26, 0.28 0.29, 0.27 0.26, 0.26 0.27 (0.28) 0.22, 0.23 0.23 0.22 mid. claw 0.09, - 0.09, 0.09 0.09, - 0.09, - 0.08 0.07 hind f+t 1.55, 1.50 1.60, 1.67 1.55, 1.54 1.15 (1.30) 0.97, 0.98 1.00 1.15 hind tibia 2.17, 2.17 2.18, 2.20 2.11, 2.09 1.97 (2.10) 1.29, 1.30 1.29 1.36 hind pts 0.45, 0.44 0.50, 0.51 0.46, 0.46 0.82 (0.88) 0.46, 0.48 0.64 0.51 hind dts 0.27, 0.27 0.28, 0.28 0.26, - 0.32 (0.32) 0.29, 0.27 0.27 0.25 hind claw 0.09, 0.09 0.085, 0.09 0.09, - 0.09, - 0.08 0.07 head W - - 1.00 - - 0.76 fch 0.20 - - n/a ?n/a [small] ant. I 0.07 - 0.09, - 0.08 (0.08) 0.09, - 0.05 0.05 ant. II 0.15 0.15, 0.16 0.15, - 0.16 (0.18) 0.14, 0.14 0.15 0.15 ant. III 0.63, 0.65 0.71, 0.76 0.62, - 0.88 0.54, 0.50 0.58 0.49 ant. IV 0.31, 0.26 0.28, 0.30 0.26, - 0.35 0.28, 0.28 0.27 0.31 ant. V 0.12, 0.16 0.15, 0.165 0.14, - 0.28 0.18, 0.19 0.28 0.12 ant. VI 0.055 0.06, 0.055 0.06, - 0.17 0.07, 0.08 0.10 0.095 ant. VII 0.04 0.03, 0.035 0.03, - 0.13 0.07, 0.065 0.075 0.06 urs 0.43 0.41 0.40 (0.34 ) 0.33 0.34 0.34 vas no no no ? yes no no n/a

* Enderlein did not state whether this included the forceps ** Calculated from 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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