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 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. Downloaded from Brill.com09/30/2021 04:39:28AM via free access 14 Tijdschrift voor Entomologie, volume 150, 2007 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.’ Downloaded from Brill.com09/30/2021 04:39:28AM via free access Martin: Giant whiteflies and new Udamoselis 15 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