Aleyrodes spiraeoides Quaintance, 1900

Aleyrodes spiraeoides Quaintance, 1900 View in CoL

Fig. 10–12 View Figures 7–12 , 21–28 View Figures 21–24 View Figures 25–28 , 32 View Figures 29–32 , 37–41 View Figures 33–41 , 45 View Figures 42–45

Aleurodes spiraeoides Quaintance 1900: 36 (description of nymph, puparium, and adult male, plate 4, fig. 45–49, plate 8, fig. 74; syntypes from U.S.A., California, Los Angeles, on Fuchsia View in CoL , 23.x., A. Craw coll.; Los Angeles on Sonchus View in CoL , 21.x.1887, D.W. Coquillett coll.; Alameda on Convolvulus View in CoL [now Calystegia View in CoL ] occidentalis View in CoL , xi.1887, A. Koebele coll.; same but on Malvia rotundifolia [now Malva pusilla View in CoL ], 5.xi.1885; and California, no locality given, on Iris View in CoL , 20.x.1880, J.H. Comstock coll., at USNM).

Aleyrodes spiraeoides View in CoL : Bemis 1904: 530 (description of egg, adult male and female, additional host plant records; plate 35, fig. 56–60) — Quaintance and Baker 1914: 101, illustrations of puparium and adults, photograph of puparium on leaf, plate 38, fig. 1–13; plate 46, fig. 6).

Aleyrodes diasemus Bemis 1904: 516 View in CoL (description of nymph and puparium; syntypes from U.S.A., California, on Symphoricarpos racemosus View in CoL [now S. albus View in CoL ], Leland Stanford Junior University campus, along San Francisquito Creek, 18.ix.1901; on Ribes glutinosum View in CoL [now R. sanguineum var. glutinosum View in CoL ] near Menlo Park, ix.1901, Alameda, vi.1901, and King Mountain, viii.1901, at USNM but currently presumed lost) — Russell 1948: 78 (change of combination; discussion of type material). New synonymy.

Asterochiton diasemus : Quaintance and Baker 1914: 105 (change of combination).

Trialeurodes diasemus : Quaintance and Baker 1915: xi (change of combination).

Aleyrodes essigi Penny 1922: 23 View in CoL (description of larva, puparium, and adult; illustrations of puparium and adult, fig. 1; syntypes from U.S.A., California, Mission San Jose , on Ulmus sp. , ix.1916, E.O. Essig coll., at CAS). New synonymy.

Aleyrodes osmaroniae Sampson 1945: 58 View in CoL (description of puparium, fig. 1; holotype from U.S.A., California, Berkeley University campus, Strawberry Creek Canyon, on Osmaronia [now Oemleria View in CoL ] cerasiformia, 14.vi.1941, W.W. Sampson coll., at BME). New synonymy.

Specimens examined. Slide mounted material: 460 puparia, 68 ♂♂, 55 ♀♀ [at CSCA unless indicated otherwise]: U.S.A., California, 14 puparia [identified as A. osmaroniae by W.W. Sampson] #27; Imperial County: 5 puparia, El Centro, on Aster exilis , 12.vii.1973, R.A. Flock coll.; 2 puparia, Brawley, 7.xi.1975, on yerba santa ( Eriodictyon californicum ), Flock and Pineda coll.; San Diego County: 9 puparia, Fallbrook, on Fraxinus sp. and milkweed ( Asclepias sp. ), 19.xi.1973, N. Metcalf coll.; 15 puparia, 2 ♂♂, 2 ♀♀, Rancho Santa Fe, 26.x.1973, P. Gomes coll.; 6 puparia, San Diego, on unknown shrub, 9.xii.1987, J. Kenyon coll.; 2 puparia, San Ysidro, on Myoporum sp. , 22.viii.1988, J. Kenyon coll.; 2 puparia, La Mesa, on Myoporum sp. , 10.xi.1996, D. Kellum coll.; 2 puparia, El Cajon, 16.v.1934, L.W. Fox coll.; Orange County: 9 puparia, 1 ♀, Santa Ana, on Jerusalem cherry ( Solanum pseudocapsicum ), 5.i.1932, T.E. McLead coll.; 1 ♂, 1 ♀, Westminster, on Myoporum sp. and iceplant ( Carpobrotus edulis ), 16.i.1973, Ellis and Lilly coll.; 16 puparia [some with long setae], Laguna Beach, on Myoporum laetum , 5.xi.2010, J. Bethke and N. Nisson coll.; Riverside County: 2 puparia [with long setae], Riverside, on Euphorbia supina [now Euphorbia maculata ], viii.1982, J. LaSalle coll.; 1 puparium [with long setae], Moreno, on potatoes ( Solanum tuberosum ), 25.ix.1979, E. Reeves coll.; San Bernadino County: 2 puparia, San Bernardino Mountains, on Burchellia sp. , W.W. Sampson coll.; 8 puparia, Ontario, 5.xii.1937, on Rosa sp. , G.A. Pohl coll.; Los Angeles County: 1 ♂, 1 ♀, Anaheim, on Citrus sp. , 1.xi.1932, M.A. Olsen; 1 ♀, San Marino, on Citrus sp. , 30.xii.1935, T. Gallion coll.; 9 puparia, Long Beach, on Iris sp. , 20.ii.1936, A.E. Bottel coll.; 5 puparia, El Monte, 23.vii.1938, on weed in ‘berry patch’, E.H. Schlenz coll.; 5 puparia, same data but 31.vii.1938; Santa Catalina Island, Toyon Canyon, on sow thistle ( Sonchus sp. ), 29.v.1981, R.J. Gill coll.; 10 puparia [with long setae], Los Angeles, on Fuchsia sp. , 18.xi.1986, Burke coll.; Ventura County: 2 puparia, Ojai Valley, on weed, 20.i.1932, E. Smith coll.; 16 puparia, 13 ♂♂, 2 ♀♀, Saticoy, on Penstemon sp. , 18.xi.1933, Barrett coll.; 4 puparia, Hueneme, on flowering apple ( Malus sp. ), 20.ix.1933; 15 puparia, Camarillo, on Iris sp. , 9.iii.1939, A.H. Calf coll.; Monterey County: 6 puparia, Salinas, on rose ( Rosa sp. ) and Malva parviflora , viii.1986, W.C. Wasik coll.; 2 ♂♂, 5 ♀♀, San Ardo, on potato ( Solanum tuberosum ), 31.viii.2001, Oliver coll.; Santa Barbara County: 4 puparia, Santa Barbara, 16.v.1972, on pansy ( Viola sp. ), C.C. Benedict coll.; 4 puparia, same data but on Iris sp. , 26.v.1972; 3 puparia, Carpinteria, on Gerbera sp. , 18.xi.2001, J. Davison coll.; 3 puparia, on Ceiba speciosa , 9.x.2003, M. Perry coll.; San Luis Obispo County: 2 puparia, 1 ♂, on Fuchsia sp. , 14.xii.1967, J. Williams coll.; 5 puparia, San Luis Obispo, on ‘skeleton weed’, 11.vii.1980, B. Lilley coll.; 6 puparia, 1 ♂, Nipomo (in nursery), on Iris sp. , 17.viii.2001, C. Taylor coll.; Tulare County: 2 puparia, Exeter, on ornamental, 4.x.1935, J.M. Awbrey coll.; Merced County: 29 puparia, Merced, on Veronica sp. , 15.xi.1928, D.P. Wheeler coll.; 1 puparium [with long setae], Los Baños, on tomatoes ( Solanum lycopersicum ), 9.vi.1992, V. Castellano coll; Santa Cruz County: 14 puparia, Watsonville, sow thistle ( Sonchus sp. ), 8.viii.2001, Bowman coll.; 1 ♂, same data but on kale ( Brassica oleracea ); Stanislaus County: 14 puparia [some with long setae], Knights Ferry, on Bidens sp. , 18.x.1978, Bingham coll.; 2 puparia [with long setae], on Verbena sp. , 4.v.1981, Zorn and Bingham coll.; Contra Costa County: 2 puparia, Antioch, on Iris sp. , 21.i.1931; 2 puparia, 1 ♂, on peppers ( Capsicum sp. ), 20.vii.1992, J. Caprile coll.; Alameda County: 13 puparia [some with long setae], 8 ♂♂ [syntypes of Aleyrodes essigi, CAS type #6189], Mission San Jose, on Ulmus sp. , ix.1916, E.O. Essig coll. [ CAS]; 16 puparia, Berkeley, 5.iii.1934, E.A. Drews and W.W. Sampson coll.; 4 puparia, same data but on cerasiformis , 20.xi.1934, W.W. Sampson coll.; 8 puparia [paratypes of Aleyrodes osmaroniae ], Strawberry Creek Canyon, U.C. Berkeley, on Osmaronia [now Oemleria ] cerasiformis, W.W. Sampson coll.; 1 puparia [holotype of Aleyrodes osmaroniae , circled, BME type #850] + 4 puparia [paratypes of Aleyrodes osmaroniae ], same data but [ BME]; 1 ♂, Hayward, on azalea ( Rhododendron sp. ), 22.x.1938, F.J. March coll.; San Francisco County: 18 puparia, Golden Gate Park, on Iris sp. , 24.i.1934 & 24.i.1939, W.W. Sampson and E.A. Drews coll.; 26 puparia, 1 ♂, 1 ♀, same data but on Myoporum laetum ; 1 ♂, 2 ♀♀, San Francisco, on primrose ( Primula sp. ), 5.ix.1936, R. Kausen coll.; 4 ♂♂, 8 ♀♀, Albany, on Iris sp. , x.1992, B. Campbell; San Joaquin County: 9 puparia [with long setae], Stockton, on tomato ( Solanum lycopersicum ), 4.vii.1984, T. Gantenbein coll.; Calaveras County: 1 puparium, 2 ♂♂, Big Trees, 13.viii.1992, D. Norfolk and K. Kenston coll.; Sacramento County: 2 ♂♂, Sacramento, on rose ( Rosa sp. ), 6.xii.1932, McFarlane coll.; 3 ♂♂, 2 ♀♀, Courtland, on Malva sp. , 19.iv.1933, Bachman coll.; 2 ♂♂, 2 ♀♀, Sacramento, on Mahonia sp. , 18.xi.1936, Wilkinson coll.; 1 ♂, 2 ♀♀, Folsom, on Citrus sp. , 31.viii.1937, W.P.A. coll.; 3 ♀♀, Mills District, on privet ( Ligustrum sp. ), 7.x.1937, W. Travioli coll.; 1 ♂, 1 ♀, Mayhews, 11.x.1937, W. Travioli coll.; 3 ♂♂, 1 ♀, Folsom, on privet ( Ligustrum sp. ), 8.ix.1937, H.H. Keifer coll.; 3 ♂♂, 4 ♀, same data but 8.ix.1939; 1 puparium, 1 ♂, on Aster sp. , 31.vii.1939, C. Schiller coll.; 3 puparia, Sacramento, on Penstemon sp. , 16.xii.1942, J.B. Steinweden coll.; 4 puparia, same data but on Iris sp. , vi.1977, R.J. Gill coll.; 2 ♂♂, 4 ♀♀, Sacramento, on rose ( Rosa sp. ), 24.i.1987, R.J. Gill coll.; 12 puparia, Galt, on Stevia sp. , 6.viii.1997, Wilson and Thompson coll.; Yolo County: 5 puparia, Woodland, on Cercis sp. , xi.1937, Wymore coll.; 1 puparium, Davis, on Arizona ash ( Fraxinus velutina ), 23.ix.1937, R.W. Downey coll.; 1 ♂, 3 ♀♀, Bryte, on Oregon grape ( Berberis aquifolium ), 26.ix.1937; 5 puparia [with long setae], on strawberry ( Fragaria sp. ), 3.x.1985, J. Wagoner coll.; Sonoma County: 1 puparium, Sonoma, on Hypericum sp. , 13.viii.1973, I. Violotti coll.; 5 puparia, Cloverdale, on honeysuckle ( Lonicera sp. ), 10.vi.1980, R.J. Gill coll; Napa County: 4 puparia, St. Helena , on Hypericum sp. , 6.iii.1934, Niles coll.; Yuba County: 2 ♂♂, Wheatland, on orange ( Citrus × sinensis ), 14.vii.1937, Schaffer coll.; Mendocino County: 2 puparia, 4 ♂♂, 7 ♀♀, Fort Bragg, on Rhododendron sp. , 17.v.1936, M.L. Jones coll; Humboldt County: 3 ♂♂, 2 ♀♀, Feildbrook, on Rhododendron sp. , 22.ii.2000, Haggard coll; Bute County: 8 puparia, Chico, on Fuchsia sp. , 26.x.1938, R. Swett coll.; 6 puparia, Wheatland, on Jerusalem cherry ( Solanum pseudocapsicum ), 12.iii.1936, Branson and Lehrer coll.; 3 puparia, Sutter County, Yuba City, on rose ( Rosa sp. ), 27.viii.1931, L.S. Jones coll.; Lassen County: 1 puparium [with long setae], Hillside Cemetery, on Hemizonia [now Deinandra ] lobbii, T.C. Fuller coll.; Shasta County: 1 puparium [with long setae], on Fragaria sp. , 5.ix.1961, J. Rodigou coll.; Arizona, Coconino County: 17 puparia, Flagstaff, on Euphorbia sp. , xi.2004, D. Byrne coll.; interception from Texas, 2 ♀♀, on salal ( Gaultheria shallon ), 5.ii.1996, W. Bianchi coll. Dry material: U.S.A., California, Riverside Co.: 1 puparium, Banning, on Tecomaria capensis , 26.viii.1942, Lower coll.; San Benito County: 4 puparia, Hollister, on honeysuckle ( Lonicera sp. ), 22.iv.1948, R.P. Allen coll.; 1 puparium, same data but on privet ( Ligustrum sp. ), 23.iv.1948, Gammou and Allen coll.; Madera County: 50 puparia, Chowchilla, on Iris sp. , 11.viii.1948, E. Danison coll.; Sacramento County: 1 puparium, Sacramento, on Penstemon sp. , 16.xii.1942, J.B. Steinweden coll.

Diagnostic characters. Puparium translucent yellow to tan ( Fig. 10–12 View Figures 7–12 ), elliptical but outline sometimes distorted (undulate or elongate) due to development amongst leaf hairs. Medio-dorsal area of cephalothorax and anterior abdomen never with pebble-like ornamentation and usually with only up to eight simple pores per segment on each side ( Fig. 21–24 View Figures 21–24 , 32 View Figures 29–32 ). Adult male with paramere narrowing apically, with a narrow subapical flange, as wide as 0.30–0.35 of its length and carina along medial margin of paramere contiguous to tip; aedeagus gradually curved at a 45-degree angle ( Fig. 45 View Figures 42–45 ).

Remarks. Puparium of A. proletella appears to be identical but that species prefers cruciferous hosts; reliable identification can be achieved by examination of male genitalia (see under A. proletella ). Puparia of A. pruinosus also seem to be undistinguishable morphologically from those of A. spiraeoides , usually differing by their relatively larger size and larger vasiform orifice (although ranges overlap; see Table 1) and color pattern, uniformly grayish brown usually with narrow margin yellow, although some specimens of A. spiraeoides present also an extensive tan coloration (e. g., Fig. 10 View Figures 7–12 ); we have been unable to distinguish between the adults of these two species. Puparia with long setae could be mistaken for the European A. lonicera Walker, 1852 which can also have long setae (known so far in the USA only from Florida) but differ from it by vasiform orifice yellow and abdominal segments lacking median tubercles, versus vasiform orifice often dark and usually with shallow median tubercles on abdominal segments II–V or II–VI in A. lonicera ( Martin et al. 2000; Stocks 2012). Their adults differ by the two diffuse dark spots on fore wing in A. spiraeoides , versus only one in A. lonicerae ( Martin et al. 2000; Stocks 2012).

In his description of this species, Quaintance (1900) mentioned the absence of pores on dorsum of puparium. Although pores might not be visible in specimens that have been improperly cleaned or are unstained, they are present in all well-preserved specimens that we examined (e. g., Fig. 21–24 View Figures 21–24 , 32 View Figures 29–32 ).

New synonymies. Bemis (1904) described A. diasemus based on an unspecified number of nymphs and puparia collected on leaves of Symphoricarpos racemosus (now S. albus ) and Ribes glutinosum from three localities in Alameda and San Mateo Counties in California deposited at USNM (Type No. 7096). Quaintance and Baker (1914) transferred the species to Asterochiton Maskell and later ( Quaintance and Baker 1915) to Trialeurodes Cockerell. Russell (1948: 78) examined four slides at USNM labeled by Bemis as A. diasemus , without type labels and slightly mismatching the date or locality given for the types in the description, that she suspected represented the syntypes of this species, and concluded that the species belonged in the genus Aleyrodes rather than Trialeurodes . Unfortunately, no specimens labeled as A. diasemus were currently located at the USNM (I. Stocks and G. Evans pers. comm.), and its types are therefore presumed lost. Bemis’ (1904) description of the puparium mentioned the presence of 12 pairs of extremely long, stout spines. The position that he described for these ‘spines’ agrees with the position of the marginal, dorsal, and caudal setae characteristic of Aleyrodes , and we believe that these ‘spines’ correspond to enlarged setae. Bemis (1904) also mentioned a considerable variation in the amount and kind of wax secretions: ‘specimens may have both lateral and dorsal wax, or either alone, or none; when present the lateral fringe is of coalesced crystalline wax rods either free from or covered by flocculent wax; the dorsal secretion is in the form of a submarginal series of separate crystalline wax rods, rather long and curved downward.’ According to Russell (1948), the specimens of A. diasemus studied by Bemis were collected and mounted together with Trialeurodes vaporariorum (Westwood) (identified by Bemis as Aleyrodes glacialis Bemis ). Therefore, it is possible that the wax pattern described by Bemis (1904) under A. diasemus for some puparia corresponded to specimens of T. vaporariorum , explaining the unusual variability he noted. Some puparia of A. spiraeoides at the CSCA collection collected from Fragaria sp. , Myoporum , Solanum lycopersicum , and Verbena sp. display a variable number of very long (longer than vasiform orifice) setae (e. g., Fig. 24 View Figures 21–24 ) which agree in their positions with the ‘long spines’ described by Bemis (1904). The number of long setae in these populations ranges from none to up to 12 pairs, with much variability regarding which particular setae are enlarged, although the caudal pair is most commonly involved, and in some specimens only one of a pair of setae is extremely long and the other one is of the short length usual for this species. Also, we observed some specimens on Euphorbia sp. , Hemizonia lobbii , Fuchsia sp. , and Solanum tuberosum with long caudal setae (e. g., Fig. 23 View Figures 21–24 ). This variability observed among specimens from the same populations on the same leaves and even within the same specimens indicates that the length of the setae of the puparia is variable and does not represent interspecific differences, being most likely the result of their development on pubescent leaves. Mound (1963) showed that the number, size, and placement of dorsal setae in Bemisia tabaci (Gennadius) may be correlated with the amount, distribution, and shape of leaf hairs on the host surfaces, and we suspect that a similar explanation applies in this case. Since other than the wax pattern, which is here interpreted to be the result of the mixed series used for the description, and the long ‘spines’, which are a match for the long setae observed on some puparia of A. spiraeoides developing on pubescent leaves, the description of A. diasemus is congruent with the characteristics of A. spiraeoides , A. diasemus is here considered a junior subjective synonym of A. spiraeoides .

The type series of A. essigi Penny, 1922 , described from Ulmus sp. in Alameda County, was borrowed from CAS in order to diagnose this species. It comprises four slides and three vials with host leaves and dry puparia. The dry material shows numerous puparia on the underside of pubescent leaves ( Fig. 12 View Figures 7–12 ). One of the slides includes five puparia and is in very bad preservation state, with air having leaked inside and puparia barely visible. A second slide includes eight puparia, which are poorly cleared, and whose characters are also obscured by the pharate adults in several of them. The puparia display a variable number of long setae in different positions and not consistent among the specimens (e. g., Fig. 25, 26 View Figures 25–28 ), and some have the same setae very short (e.g., Fig. 27 View Figures 25–28 ). Not a single specimen seems to present all four pairs illustrated as being very long by Penny (1922: fig. 1) and his drawing seems to have been a composite. The contour of the puparia varies from broadly elliptical to undulate or slightly elongate ( Fig. 25–27 View Figures 25–28 ). The remaining two slides include adults, one an emerging male, and the other one seven adult males. None of the specimens were properly cleaned or stained, but the male genitalia is visible in all of them. No differences between the syntypes of A. essigi deposited at CAS and specimens of A. spiraeoides with enlarged setae were found, nor between the adult male syntypes and adult males of A. spiraeoides , and therefore A. essigi is here considered to represent a junior subjective synonym of A. spiraeoides , the name simply representing specimens of A. spiraeoides developing on pubescent leaves as in the case of A. diasemus discussed above. Penny (1922) described the adult ‘female’, likely a typo since all specimens in the type series are males, of A. essigi as having ‘immaculate’ wings. The apparent lack of the two diffuse spots on the wings usually visible in A. spiraeoides can be explained by the fact that these spots are not always evident in slide mounted specimens.

Sampson (1945) described Aleyrodes osmaroniae from puparia collected with specimens that he identified as A. spiraeoides on Osmaronia [now Oemleria ] cerasiformia in Alameda County, and he diagnosed it from A. spiraeoides by the presence of a chitinized margin. The holotype and paratypes deposited at BME and paratypes deposited at CSCA were examined. In some specimens there appears to be a fine submarginal line along the puparium, and the marginal area so delimited appears darker, but no ‘chitinized’ areas were detected. The unusual appearance of the margin is interpreted here to represent the result of uneven staining of the cuticle, which is highly variable in the type series and within separate areas of the margin on some specimens (e. g., Fig. 28 View Figures 25–28 ). The same submarginal line (on some sections of the puparium or along its entire contour) was observed in other specimens of A. spiraeoides (e. g., Fig. 23 View Figures 21–24 , 25, 27 View Figures 25–28 ) and in other Aleyrodes species (e. g., Fig. 14, 16 View Figures 13–16 , 17, 20 View Figures 17–20 ), and it might be an artifact caused by one of the surfaces (dorsal or ventral) of the puparium being folded away from the margin during slide mounting. No differences were found among the type specimens of A. osmaroniae and puparia of A. spiraeoides , and therefore A. osmaroniae is here considered a junior subjective synonym of A. spiraeoides .

Hosts. Polyphagous, in California most common on Iris Tournefort ex. L. and Gladiolus L. ( Iridaceae ) but recorded from a wide range of plants (see Table 2). Penny (1922) recorded it from Ceanothus among other hosts, Creek Canyon, U.C. Berkeley, on Oemleria cerasiformis, W.W. Sampson coll. [BME type #850]. Figures are not to scale. but since all the specimens we examined from Ceanothus correspond to A. amnicola , we consider that record as doubtful and omit it from its host plant list. Landis et al. (1958) provided additional host plants based on specimens from the Pacific Northwest (Washington, Oregon, and Idaho States).

Distribution. Throughout the state, especially in urbanized areas. Recorded also from Arizona, Colorado, Florida, Idaho, Hawaii, Louisiana, Nevada, Oregon, Utah, Texas, and Washington States in the U.S.A., and from Canada, Mexico, and Venezuela ( Landis et al. 1958; Evans 2007).