Lipaleyrodes emiliae sp. nov.

(Figs 1–53)

PUPARIUM (Figs 1A, 10). Pale to yellowish, average size 0.66 mm long and 0.47 mm wide, 1.4 times as long as wide. Elongate to ovoid, broadest at metathorax. Found in groups on the undersurface of leaves. Margin crenulate (Fig. 11). Pairs of anterior and posterior marginal setae evident, anterior 10 µm long; posterior 25 µm long. Thoracic and caudal tracheal area not evident. Lingula appearing reddish-brown pigmented (Fig. 48). Most pupae with a marginal ring of uneven, fine, white wax (Fig. 51). This waxy secretion associated with a kind of minute pore (Figs 13–14). At a distance, groups of immatures appear to have been dusted with a white powder, although closer examination indicates the presence of white, fluffy and contorted wax filaments (Fig. 52). Some specimens show accumulation of this waxy secretion on the thoracic area. Dorsum. Dorsum with 2 pairs of dorsal setae; cephalic setal length ranging widely from 21 to 82 µm, 102 µm apart; abdominal VIII setae much longer than vasiform orifice, 102 µm long, 82 µm apart. Longitudinal molting suture reaching margin, but transverse one failing to. Abdominal sutures ending on subdorsum. Caudal setae submarginal, 101 µm long, 52 µm apart; submarginal tiny setae 7 pairs, 2 pairs on cephalic area, 5 pairs posterioral to transverse moulting suture; porettes ill-defined; abdominal segment I 33 µm long; segments II–V slightly subequal, 28, 30, 30 and 29 µm long, respectively, segment VI 38 µm long; segment VIII longest at 42 µm; segment VII much reduced, medially shortened, 9 µm long. Vasiform orifice triangular to trapezoidal, posterior margin poorly defined, with lateral teeth on both sides; a little longer than wide at 70 x 63 µm (Fig. 16). Operculum semicircular, 1.41 times as wide as long at 36 x 51 µm; truncate at hind margin, occupying about half of vasiform orifice. Lingula exposed, 74 µm long, almost included within vasiform orifice except for the slightly tapering, spinulose apex that is exposed; a pair of subapical setae present, 29 µm long. Caudal furrow and ridges barely indented (Figs 1C, 15). Dorsal disc separated from submargin by a distinct fold. Submargin broad with 11 pairs of wax plate clusters, wax plates oval (Figs 1B, 12), clusters I and II with fewest (<5) wax plates; clusters IV–VI and XI with most wax plates, each with 1015; each of clusters III and VII–IX with 79. Ven t er (Fig. 17). A pair of ventral abdominal setae present, 4.6 µm long, 75 µm apart. Antenna not extending beyond base of prothoracic leg (Fig. 18). Outline of legs curved. Setae tiny at base of legs and rostrum.

THIRD INSTAR (Figs 2A, 19). Pale to yellowish, with an average size 1.43 times as long as wide at 0.46 x 0.32 mm. Elongate to ovoid, broadest at metathorax. Margin crenulate. Pairs of anterior and posterior marginal setae evident: anterior 5.6 µm long, posterior 20 µm long. Thoracic and caudal tracheal areas not evident. Lingula appears reddish-brown. Dorsum. Dorsum with 2 pairs of dorsal setae; cephalic setae varied in length at 8–28 µm, 79 µm apart; abdominal VIII setae 54 µm long, 66 µm apart. Abdominal sutures ending on subdorsum. Caudal setae submarginal, 84 µm long, 37 µm apart. Abdominal segment I 26 µm long; segments II–VI subequal, 22, 20, 20, 18, and 22 µm long, respectively; segment VIII longest at 34 µm; segment VII much reduced, medially shortened, 7.5 µm long. Vasiform orifice triangular to trapezoidal; wider than long, at 49 µm long x 52 µm wide. Operculum semicircular, 1.61 times as wide as long at 26 x 42 µm; truncate at hind margin, occupying about half of vasiform orifice. Lingula exposed, 59 µm long, almost enclosed in vasiform orifice, except a small part of posterior end exposed outside, awl-shaped, slightly tapering, spinulose; a pair of subapical setae present, 18 µm long. Caudal furrow and ridges barely indented (Figs 2B, 20). Dorsal disc separated from submargin by a distinct line. Ve n te r (Fig. 21). Antenna thin and not extending beyond base of prothoracic leg. Legs appearing circular-conical. Setae tiny on base of legs and rostrum.

SECOND INSTAR (Figs 3A, 22). Pale to yellowish, with an average size 1.6 times as long as wide at 0.37 x 0.23 mm. Elongate to ovoid, broadest at metathorax. Margin crenulate. Pairs of anterior and posterior marginal setae evident, anterior 5.9 µm long; posterior 17 µm long. Thoracic and caudal tracheal area not evident. Lingula appears reddish-brown. Dorsum. Dorsum with 2 pairs of dorsal setae; cephalic setae 13 µm long, 53 µm apart; abdominal VIII setae much shorter than vasiform orifice, 19 µm long, 49 µm apart. Abdominal sutures ending on subdorsum. Caudal setae marginal, 59 µm long, 26 µm apart; abdominal segment I 18 µm long; segments II–V subequal, 13, 15, 14, and 12 µm long, respectively; segment VI 17 µm long; segment VIII longest at 27 µm; segment VII much reduced, medially shortened, 4.3 µm long. Vasiform orifice triangular to trapezoidal, posterior margin truncated, with lateral teeth on both sides; as long as wide at 38 x 39 µm. Operculum semicircular, 1.60 times as wide as long at 20 x 32 µm; truncate at hind margin, occupying about half of vasiform orifice. Lingula exposed, 42 µm long, almost enclosed within vasiform orifice; exposed posterior end awl-shaped, slightly tapering, spinulose; a pair of subapical setae present, 13 µm long. Caudal furrow and ridges barely indented (Fig. 23). Ve n te r (Fig. 24). Antenna not extending beyond base of prothoracic leg. Legs circular-conical. Setae tiny on base of legs and rostrum.

FIRST INSTAR (Figs 3B, 25). Pale to yellowish, with an average size 1.69 times as long as wide at 0.23 x 0.14 mm. Elongate to ovoid, broadest at mesothorax. Margin crenulate, surrounded by 15 pairs of variably sized setae (including posterior and caudal setae). Posterior marginal setae 17 µm long. Thoracic and caudal tracheal area not evident. Dorsum. Abdominal segment VIII setae much shorter than vasiform orifice, 2.3 µm long, 32 µm apart. Abdominal sutures ending on subdorsum, suture between segments II and III much thicker than others. Caudal setae marginal, 57 µm long, 20 µm apart; abdominal segments I and II usually not visible, segment III 8 µm long; segments IV–VI subequal, 11, 11, and 10 µm long, respectively; segment VIII longest at 21 µm; segment VII much reduced, medially shortened, 2.6 µm long. Vasiform orifice triangular to trapezoidal, about as long as wide at 27 x 26 µm; posterior margin truncated, with lateral teeth on both sides. Operculum dumbbell-shaped, 2.15 times as wide as long at 11 x 23 µm; truncate at hind margin, occupying less than half of vasiform orifice. Lingula exposed, 23 µm long, extending slightly beyond vasiform orifice, the posterior end semicircular, spinulose; a pair of subapical setae present, 6.9 µm long. Caudal furrow and ridges barely indented (Fig. 26). Ve n te r (Fig. 27). A pair of ventral abdominal setae present, 4.3 µm long, 25 µm apart. Antenna 62 µm long, 3-segmented, with apical seta. Legs segmented. Setae present at base of mid and hind legs.

EGG (Figs 9D, 28). Elongate to oval, surface smooth with brown metallic luster, 132 µm long, often found on undersurface of leaf and covered with waxy secretion. Base with a short pedicel, 26 µm long.

ADULT MALE. Length from vertex to claspers, 0.79 mm. One ocellus present laterally on each side, located close to upper compound eyes. Upper and lower compound eyes joined by a single ommatidium as in B. tabaci (Figs 5, 31). Tip of labium with sensorial hairs (Figs 29, 30). Antenna (Fig. 4A). As in other Lipaleyrodes species, near compound eyes, 7-segmented; segment I of short cylindrical form, 14 µm long; segment II 39 µm long; segment III longest, sub-cylindrical, 87 µm long, armed with 2 primary sensoria and 1 sensorial cone, sensoria located apically and sub-apically on segment III, tip of cone extending to base of sub-apical primary sensoria (Fig. 32); segment IV subcylindrical, 17 µm long, segment V sub-cylindrical, 25 µm long, with a primary sensoria apically (Fig. 33); segment VI sub-cylindrical, 19 µm long, with a sensorial cone subapically, distal end extending to junction of segments VI and VII; segment VII subfusiform, 30 µm long, with a primary sensorium and sensorial cone in the distal half and a seta on the tip; cone long at over one-third of segment VII (Fig. 34). Wings. Forewing 604 µm long, 215 µm wide. Hindwing 527 µm long, 185 µm wide. Wings not mottled, limpid, covered with white waxy secretions. Wing venation reduced to costal-subcostal veins and radial vein. Legs. Mesotibia 184 µm long with ordinary rows of spines, brushes with 2 and 3 setae; proximal tarsus 59 µm long; distal tarsus 55 µm long (Figs 6 A, 35). Hindtibia 249 µm long; proximal tarsus 74 µm long; distal tarsus 64 µm long (Fig. 7 A); ending with 2 claws and paronychium, a little shorter than claw and covered with fine microtrichia at base (Fig. 37). Metatibial comb ordinarily with rows of 15 spines (Fig. 36) (in most individuals), numbers of spines not the same on all observed specimens (range, 1417). Genitalia. Aedeagus not bifurcate, 77 µm long, 11 µm wide, shorter than clasper. Clasper 23 µm at base, 88 µm long; 7 setae in mid-region (Figs 9B, 38 39).

ADULT FEMALE. Most of the characters are similar to those of males. Vertex to tip of ovipositor 0.89 mm in length. Antenna (Fig. 4B). Segments I–VII 15, 45, 99, 20, 28, 22, and 32 µm long, respectively. Wings. Forewing 748 µm long, 285 µm wide (Fig. 8A). Hindwing 649 µm long, 241 µm wide (Fig. 8B). Legs. Mesotibia 204 µm long, proximal tarsus 70 µm long, distal tarsus 67 µm (Fig. 6 B). Hindtibia 291 µm long, proximal tarsus 88 µm long, distal tarsus 74 µm (Fig. 7 B). Genitalia. Nine pairs of setae on gonapophysis, 2 pairs centrally, and 7 pairs laterally (Figs 40 41). Bulb-like orifice of cement gland trapezoidal, extremely curved, not mottled, full length 36 µm (Fig. 9A).

COMMENTS. In Taiwan, two different species are now included in the genus Lipaleyrodes: L. breyniae and L. emiliae . The puparia of both species have 11 pairs of wax plate clusters on the submarginal band, and all plates are oval in the two species. Each cluster consists of 4 or 5 wax plates in L. breyniae, but of 5 to 15 in L. emiliae . Lipaleyrodes breyniae occurs on euphorbiaceous and papilionaceous plants, whereas L. emiliae is restricted to Asteraceae, mostly Emilia sonchifolia . This new species may be distinguished from L. breyniae by the shorter the puparium and longer abdominal VIII setae, which are much longer than the vasiform orifice. In contrast, L. breyniae has a longer puparium and the abdominal VIII setae are shorter than the vasiform orifice. Moreover, the lingula apex of L. emiliae is particularly distinctive.

In Taiwan, we often found L. emiliae together with Bemisia tabaci (Gennadius) on the lower surface of the leaves of Emilia sonchifolia . These two species can be distinguished because the waxy secretion of L. emiliae is always dense, white, and fluffy, composed of filaments full of twists and turns (Fig. 52), whereas B. tabaci puparia secrete no noticeable white wax. Moreover, the lingula of live L. emiliae is dark brownish-black in all stages, particularly at the edges, whereas the lingula of B. tabaci is pale yellowish (Fig. 48). It should be noted that females are larger and longer than males in both species.

No adult character states have been recognized that can be used to define membership of the genus Lipaleyrodes, including L. euphorbiae, L. crossandrae, L. emiliae, and L. vernoniae . For example, the upper and lower compound eyes are joined by just one ommatidium; the numbers and position of the primary sensoria and sensorial cones on the antenna are the same in both male and female of these species; the wing venation is reduced to costal-subcostal veins and the radial vein. We also found it difficult initially to differentiate adults of L. emiliae and B. tabaci . because the color and appearance of L. emiliae are similar to those of B. tabaci, although L. emiliae specimens look a little larger than B. tabaci . The adult lingula of L. emiliae is club-shaped (Fig. 42), being inflated at the distal end. In B. tabaci, it looks like a cylinder with a uniform diameter throughout (Fig. 47). The other obvious difference between them concerns the wax plates in both male and female adults. In B. tabaci, the wax plates are separated into two parts by a little distance, which is about one-fifth of the width of the sternum in both the male and female. Wax plates in L. emiliae are much closer than those of B. tabaci (Figs 43–46). In live adults, there is a heart-shaped spot on the thoracic pleuron and a mouth-shaped spot on the prothoracic tergum of L. emiliae (Figs 49–50), but in B. tabaci, the thoracic pleuron and prothoracic tergum are evenly yellowish and not mottled.

Material examined. Holotype puparium, TAIWAN, Renwu, on Emilia sonchifolia (Asteraceae), 6-VII-2003, C. H. Hsieh (TW1931) (NTU).

Paratypes: TAIWAN: All collected from Emilia sonchifolia except TW2173, and deposited at NTU except where otherwise noted. 2 pupal cases, 4 adult females, same data as for holotype; Taipei Co.: Shuanghsi, 5 pupal cases (on 1 microscope slide), 28-II-2004, Y. C. Hung (TW2172); Yungho, 137 pupal cases, 73 adult females, 10 adult males (on 36 microscope slides), 21-VII-2003, C. C. Ko (TW1957) (ANIC; BNHM; CDFA; NMNS; TARI; USNM); Yungho, 133 pupal cases, 32 third instars, 13 second instars, 5 first instars (on 36 microscope slides), 15-I-2004, C. C. Ko (TW2137); Sanhsia, 14 pupal cases (on 3 microscope slides), 1-I-2004, H. T. Yeh (TW2136); Sanhsia, 10 pupal cases (on 1 microscope slide), 23-X-2004, C. C. Ko (TW2388); Taipei City: Neihu, 2 pupal cases (on 1 microscope slide), 10-X-2004, C. C. Ko (TW2354); Sungshan, 8 pupal cases (on 1 microscope slide), 27-VII-2002, C. C. Ko (TW1747); Kungkuan, 43 adult females, 2 adult males (on 4 microscope slides), 14-VIII-2002, Y. C. Hung (TW1777); Taoyuan Co.: Sanguang, 10 pupal cases (on 1 microscope slide), 30-VII-2003, C. C. Ko (TW1989); Chutung, 4 pupal cases (on 2 microscope slides), 11-I-2004, H. T. Yeh (TW2146); Miaoli Co.: Nanchuang, 3 pupal cases (on 1 microscope slide), 1-VII-2003, C. H. Hsieh (TW1917); Sanwan, 7 pupal cases, 4 adult females (on 2 microscope slides), 1-VII-2003, C. H. Hsieh (TW1918); Nantou Co.: Nantou City, 7 pupal cases (on 1 microscope slide), 26-X-2003, H. T. Yeh (TW2105); Jihyuetan (Sun Moon Lake), 32 pupal cases, 1 female adult (on 6 microscope slides), 13-VIII-2004, H. T. Yeh (TW2308); Chiayi Co.: Chuchi, 3 pupal cases (on 1 microscope slide), 6-VII-2002, C. C. Ko (TW1701); Fenglu, 8 pupal cases (on 1 microscope slide), 10-IV-2004, C. C. Ko (TW2210); Chungpu, 4 pupal cases (on 1 microscope slide), 8-IX-2003, C. C. Ko (TW2026); Yichu, 14 pupal cases (on 1 microscope slide), 14-VII-2002, C. C. Ko (TW1720); Yichu, 1 pupal case, 8 adult females (on 3 microscope slides), 3-II-2003, C. C. Ko (TW1809); Tapu, 1 pupal case (on 1 microscope slide), 20-VII-2002, C. H. Hsieh (TW1732); Chiayi City, 1 pupal case (on 1 microscope slide), 7-VII-2002, C. C. Ko (TW1708); Tainan Co.: Paiho, 13 pupal cases (on 1 microscope slide), 28-IX-2003, C. C. Ko (TW2073); Paiho, 4 pupal cases (on 1 microscope slide), 28-VIII-2004, C. C. Ko (TW2314); Hsuehchia, 1 adult female (on 1 microscope slide), 5-VII-2003, C. H. Hsieh (TW1928); Liuchia, 7 pupal cases (on 1 microscope slide), 23-X-2004, C. H. Hsieh (TW2365); Hsinhua, 3 pupal cases (on 1 microscope slide), 23-X-2004, C. H. Hsieh (TW2360); Tainan City: Kuantzuling, 12 pupal cases (on 1 microscope slide), 21-VII-2002, C. C. Ko (TW1738); Kaohsiung Co.: Tashe, 3 pupal cases (on 2 microscope slides), 14-IX-2003, C. H. Hsieh (TW2053); Kaohsiung City: Shoushan, 6 pupal cases (on 1 microscope slide), 23-X-2004, H. T. Yeh (TW2385); Kaohsiung City, 5 pupal cases (on 1 microscope slide), 6-III-2004, Y. C. Hung (TW2181); Pingtung Co.: Neipu, 7 pupal cases (on 1 microscope slide), 3-VIII-2002, C. C. Ko (TW1752); Chaochou, 4 pupal cases (on 1 microscope slide), 3-VIII-2002, C. C. Ko (TW1749); Hsinpei, 2 pupal cases (on 1 microscope slide), 9-X-2004, C. H. Hsieh (TW2357); Ilan Co.: Suao, 11 pupal cases (on 3 microscope slides), 5-III-2004, C. C. Ko (TW2175); Nanao, 8 pupal cases (on 1 microscope slide), 20-II-2004, H. T. Yeh (TW2156); Hualien Co.: Hsinchen, 11 pupal cases (on 1 microscope slide), 20-II-2004, H. T. Yeh (TW2157); Liyutan, 16 pupal cases, 2 adult females (on 2 microscope slides), 2-III- 2003, C. H. Hsieh (TW1826); Taitung Co.: Changkung, 16 pupal cases, 9 adult females, 8 adult males (on 16 microscope slides), 20-II-2004, H. T. Yeh (TW2159); Heping, 9 pupal cases (on 1 microscope slide), 24-II-2004, H. T. Yeh (TW2162); Tunghou, 10 pupal cases (on 1 microscope slide), 23-II-2004, H. T. Yeh (TW2160); Painan, 4 pupal cases (on 1 microscope slide), 28-VII-2003, C. H. Hsieh (TW1972); Painan, 7 pupal cases (on 1 microscope slide), 20-II-2004, H. T. Yeh (TW2158); Taitung City, 7 pupal cases (on 1 microscope slide), 20-II-2004, H. T. Yeh (TW2161); Lutao (Green I.), 3 pupal cases (on 1 microscope slide), 26-VI-2004, C. H. Hsieh (TW2284); Lanhsu (Orchid I.), 14 pupal cases (on 1 microscope slide), 29-VII-2003, C. H. Hsieh (TW1976); Penghu Co.: Makung, 2 pupal cases (on 1 microscope slide), 19-X-2003, C. H. Hsieh (TW2099); Chinchen, 7 pupal cases (on 1 microscope slide), 8-IX-2004, C. H. Hsieh (TW2336); Shuanghsi, 1 pupal case (on 1 microscope slide), on Crassocephalum crepidioides (Asteraceae), 28-II- 2004, Y. C. Hung (TW2173).

HONG KONG: Kowloon, 6 pupal cases, 1 third instars, 2-XII-1995, C. S. K Lau (IIE23405); New Territories, 21 puparia, 2 third instars, 22-XI-1999, J. H. Martin (Martin 7255) (plus dry material on leaves); New Territories, 23 puparia, 26-XI-2005, J. H. Martin (Martin 8196) (plus dry material on leaves); Hong Kong Island, 28 puparia, 1 adult female, 5-XII-2005, J. H. Martin (Martin 8258) (plus dry material on leaves) (all in BMNH).

Etymology. The specific name emiliae is derived from the type host plant, Emilia sonchifolia .

Host plants. Asteraceae: Crassocephalum crepidioides and Emilia sonchifolia .

Distribution. Taiwan (Fig. 55), Hong Kong.

Natural ememies. Aphelinidae: Encarsia duorungu Hayat and E. sophia Timberlake. The two species are also parasitoids of B. tabaci on E. sonchifolia .

Biology. This species is apparently restricted to members of the Asteraceae, and is of no known economic importance. Emilia sonchifolia (Asteraceae) (Fig. 54) is an annual weed widely distributed in the South Pacific Islands from Indonesia to eastern Polynesia, and in Japan, Taiwan, and China, and is common in coastal regions, sandy places, wastelands, roadsides, and paddy ridges, from sea level to about 850 m. The new species of whitefly is therefore likely to have a wider geographic distribution than recorded here. The puparia were found in groups often heavily infestating the undersurfaces of a leaves amid dense white filaments of wax. No ant attendance was observed. It was also found together with Bemisia tabaci (Gennadius), intermingling with this species on the lower leaf surfaces of the plants.