Hemiptera: Sternorrhyncha: Aleyrodidae: Aleyrodinae - BioOne

THE PAN-PACIFIC ENTOMOLOGIST 89(2):84–101, (2013)

Two new species of whiteflies (Hemiptera: Sternorrhyncha: Aleyrodidae: Aleyrodinae) intercepted in quarantine on plants from Asia JOHN W. DOOLEY III AND ALLAN SMITH-PARDO United States Department of Agriculture, Animal and Plant Health Inspection Service, Plant Protection and Quarantine, 389 Oyster Point Blvd, Suite 2A, South San Francisco, California 94080 e-mail: [email protected] Corresponding author e-mail: [email protected] Abstract. Two new species of whiteflies are described and illustrated based on specimens intercepted at US Ports-of-Entry. Aleurolobus tomkinsae Dooley & Smith-Pardo sp. nov. was intercepted on the leaves of Bergera koenigii Linnaeus (Rutaceae) from India and Sri Lanka and on Magnolia L. (Magnoliaceae) from Thailand. Asialeyrodes lateropapilliformis Dooley & Smith-Pardo sp. nov. was intercepted on the leaves of Syzygium P. Browne ex Gaertn (Myrtaceae) from Indonesia. Keys to the species of the genus Asialeyrodes Corbett 1935 and to the whitefly species found on Bergera koenigii and Syzygium are also provided. Key Words. Aleurolobus, Asialeyrodes, Bergera, India, Indonesia, Magnolia, Sri Lanka, Thailand.

Resumen. Se describen e ilustran dos especies nuevas de Aleyrodidae, a partir de especı´menes interceptados en puertos de entrada a los EE.UU. Aleurolobus tomkinsae Dooley & SmithPardo sp. nov. fue interceptada en Bergera koenigii L. (Rutaceae) provenientes de la India y Sri Lanka y Magnolia Linnaeus (Magnoliaceae) de Tailandia. Asialeyrodes lateropapilliformis Dooley & Smith-Pardo sp. nov. fue interceptada en hojas de Syzygium P. Browne ex Gaertn (Myrtaceae) provenientes de Indonesia. Se proporcionan claves taxono´micas para la identificacio´n de las especies del genero Asialeyrodes Corbett 1935 asi como para la especies de mosca blanca encontradas normalmente en B. koenigii y Syzygium. Palabras claves. Aleurolobus, Asialeyrodes, Bergera, India, Indonesia, Magnolia, Sri Lanka, Tailandia.

INTRODUCTION The genus Aleurolobus Quaintance & Baker 1914 is comprised of 88 species distributed worldwide. Of these, 71 species are described from Asia and only two species are known to occur in the New World (Evans 2008). The type species of the genus, A. marlatti Quaintance 1903, was described from specimens collected on citrus in Japan. Aleurolobus tomkinsae Dooley & Smith-Pardo sp. nov. intercepted at US ports of entry since 2003 on leaves of Bergera koenigii L. (Rutaceae) in passenger baggage from India and Sri Lanka, were re-examined and identified from the PPQ collection by the author. In 2011, A. tomkinsae Dooley & Smith-Pardo sp. nov. was intercepted by USDA/APHIS/PPQ Safeguard Specialist Marilyn Tomkins at the San Francisco Plant Inspection Station on propagated plants of Magnolia L. (Magnoliaceae) from Thailand. The genus Asialeyrodes Corbett 1935 is comprised of 15 species occurring only in the eastern Palearctic and Indomalayan regions. The type species of the genus, A. lumpurensis Corbett 1935, was described from specimens collected on an undetermined plant in Malaysia. Asialeyrodes lateropapilliformis Dooley & SmithPardo sp. nov. was intercepted in quarantine on Syzygium P. Browne ex Gaertn (Myrtaceae) cuttings for propagation from Indonesia as a mail shipment.

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Plant inspection stations are located at designated US Ports of Entry throughout the United States and its territories. Safeguarding specialists that work at these facilities are responsible for 1) the inspection and clearance of all propagative commodities being imported into the United States for planting into the field and 2) the identification of pests and pathogens intercepted in quarantine at the port-of-entry stations. This paper emphasizes the need to closely inspect the leaves of plants imported for consumption or propagation that are intercepted in quarantine. Immature whiteflies are only found on the leaves and the fourth instar nymph, known as the puparium or pupa, in most cases is the only stage that can be identified to species level. MATERIALS AND METHODS The specimens used for the description of the new species were collected on plant material for consumption or propagation inspected in baggage and mail intercepted at the California, Georgia, and Illinois ports-of-entry. Eighteen puparia (4th instar) and 3 nymphs (2nd instar) of A. tomkinsae and 5 puparia and 1 nymph (2nd instar) of A. lateropapilliformis were prepared on slides. The method used for preparation of slide mounted specimens follows the techniques for scale insects and whiteflies given by Dooley (2011). Puparia were placed into test tubes containing 5% potassium hydroxide (KOH) and heated in a hot water bath for 15 min. The pale puparia of Asialeyrodes (Corbett 1935) were then washed in water to remove potassium hydroxide and soaked in Essig’s aphid fluid with the addition of 2 drops of double stain for 10–15 min. The dark puparia of Aleurolobus followed the same procedure but were not stained. Specimens were placed in 70% ethanol for 15 min and then transferred into 90% ethanol for 10 min. Specimens were transferred to clove oil for 15 min prior to slide mounting in Canada balsam. Images and measurements were taken with a SMZ 1500 wide field and a Nikon Eclipse 80i compound microscope using a Nikon DS-Fi1 Digital Image camera. Measurements in micrometers (mm) were taken from all type specimens recording the high, low and average values of the specimens with the average listed in brackets ([ ]). The holotypes and paratypes were identified at this Plant Inspection Station (Animal and Plant Health Service, Plant Protection and Quarantine at San Francisco) using available publications, digital imaging, and comparing of specimens. They were validated by Dr. Greg Evans at the Systematic Entomology Lab (Agricultural Research Service) at Beltsville, MD. The following publications were consulted: Corbett (1935), Dubey & Ko (2009), Ko et al. (1993) and Sundararaj & David (1991). Specimen Depositories. CASC—California Academy of Sciences Collection, San Francisco, California; CSCA—California State Collection of Arthropods, California Department of Food and Agriculture, Sacramento, California; PPQC—Plant Inspection Station reference collection, United States Department of Agriculture, Animal and Plant Health Service, Plant Protection and Quarantine, San Francisco, California; USNM—National Coccoidea Collection, United States National Museum of Natural History, Beltsville, Maryland. WHITEFLY MORPHOLOGY AND TERMINOLOGY The family Aleyrodidae (Hemiptera: Sternorrhyncha) consists of one fossil family, the Bernaeinae, and three extant subfamilies: Aleurodicinae, Aleyrodinae, and Udamosellinae (Martin 2007), the latter including only two South American species.

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Morphologically, the Aleurodicinae and Udamosellinae are very similar, and the latter may prove to be a junior synonym of the former (Martin 2007). The Aleurodicinae and the Aleyrodinae can be easily separated in both the adult and immature stages (instars). Historically, the 4th stage (puparium) has been used to identify species because the puparium has many more diagnostic structures than the adult and prepupal immature stages. The puparium, with few exceptions, can easily be identified to subfamily by the presence of compound pores in most species of Aleurodicinae, which are absent in all Aleyrodinae and a few Aleurodicinae species, the number of pairs of subapical setae present on the lingula (Aleurodicinae usually with two pairs and the Aleyrodinae with one pair), and the presence of an apical terminal claw (Aleurodicinae) versus an adhesive pad (Aleyrodinae) on the tarsi. Adult whiteflies can be identified to subfamily easily by the wing venation, antennal segments, and ventral abdominal wax plates. Adult males and females of all the Aleurodicinae, except for the genus Paraleyrodes (Aleurodicinae), have the R1 vein of the fore wing forked. The genus Paraleyrodes and the Aleyrodinae have the R1 vein of the fore wing simple (not forked). Abbreviations used follow those of Gill (2012) and Martin (1999) for pores, segments, and setae. Paired geminate pores and associated porettes: ASDP 5 anterior subdorsal pores and porettes, ASMeP 5 anterior submedial pores and porettes, ASMP 5 anterior submarginal pores and porettes, PSDP 5 posterior subdorsal pores and porettes, PSMeP 5 submedial pores and porettes, PSMP 5 posterior submarginal pores and porettes. Segments: A1–A8 5 abdominal segments 1 through 8, C1 5 cephalon), T1 5 prothorax (thoracic segment 1), T2 5 mesothorax (thoracic segment 2), and T3 5 metathorax (thoracic segment 3). Setae: AMS 5 anterior marginal seta, ASDS 5 anterior subdorsal setae, ASMeS 5 anterior submedial seta, ASMS 5 anterior submarginal seta, CS 5 caudal setae, PSDS 5 posterior subdorsal seta, PSMeS(A1) 5 posterior dorsal submedial A-1 seta, PSMeS(A8) 5 dorsal submedian A-8 seta, PMS 5 posterior marginal seta, PSMS 5 posterior submarginal seta, PVS(A8) 5 ventral submedian A-8 seta. RESULTS AND DISCUSSION Keys to the puparia of whiteflies occurring on Bergera, Syzygium and the species of Asialeyrodes follow. Note that the original name of the host, Murraya koenigii (L.) Spreng (Rutaceae) is a synonym of Bergera koenigii. Key to the puparia of whiteflies found on Bergera koenigii 1.

Apex of all tarsi terminating in a claw; lingula long and spatulate, extending far beyond the posterior margin of the vasiform orifice, and with 2 pairs of subapical setae; 4 pairs of abdominal compound pores present, cuticle always pale . . . . . . . . . . . . . . . . . . . Aleurodicinae: Aleurodicus dispersus Russell 1965 Apex of all tarsi terminating in an adhesive pad; lingula not long and spatulate, usually obscured by operculum, if visible not extending far beyond the posterior margin of the vasiform orifice; 1 pair of subapical setae; compound pores absent; cuticle pale to black. Aleyrodinae . . . . . . . 2 2(1). Puparium black to brown in color; with or without dorsal spines; margin dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Puparium pale; dorsal spines absent; margin dentate to crenulate . . . . . . . 7

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Puparium black with large, thick dorsal spines forming a submarginal ring; subdorsal fold or suture absent; eyespots absent; thoracic and caudal clefts not differentiated. Aleurocanthus Quaintance & Baker 1914 . . . . . . . . . . . . . . 4 Puparium dark brown without spines but robust setae may be present; subdorsal fold or suture present; pair of crescent-shaped cephalothoracic eyespots present; thoracic and caudal clefts each with 3 teeth in the species that key here. Two species of Aleurolobus Quaintance & Baker 1914 . . . . . . . 6 4(3). Marginal teeth relatively slender, 7 to 11 teeth per 100 mm; submarginal spines subequal in length . . . . . Aleurocanthus spiniferus (Quaintance 1903) Marginal teeth wider with 35 to 5 teeth per 100 mm; submarginal spines not subequal in length but various from short to long . . . . . . . . . . . . . . 5 5(4). Ten pairs of submarginal spines; submedial spines on A1 subequal in length to those on A3; fourth medial pair of spines on the cephalothorax extend well beyond the margin . . . . . . Aleurocanthus husaini Corbett 1939 Eleven pairs of submarginal spines; submedial spines on A1 less than 75x the length to those on A3; fourth medial pair of spines on the cephalothorax terminates before the margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleurocanthus woglumi Ashby 1915 6(3). ASDS and PSDS minute not reaching puparial margin (less than 12 mm long); minute setae present adjacent to the proximal margin of the dorsal fold . . . . . . . . . . . . . . . . . .Aleurolobus marlatti (Quaintance 1903) ASDS robust extending to or beyond the puparial margin (range from 14–102 mm long), abdomen with PSDS bordering subdorsal fold on A2 and on submargin on A4 (ranging in length from 48–94 mm) both extending to or beyond puparial margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleurolobus tomkinsae Dooley & Smith-Pardo sp. nov. 7(2). Subdorsal fold present; submarginal row of papillae absent; marginal teeth with or without basal glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Subdorsal fold absent; submarginal row of papillae absent or present; margin without basal glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 8(7). Subdorsal fold disjuncted (not joined together in a smooth, uniform radially-concentric pattern) and joined below the vasiform orifice; marginal teeth with basal glands giving the appearance of a double row of teeth; caudal ridge absent; lingula obscured by operculum; posterior margin of vasiform orifice with or without a branching comb-like structure. . . . . . . . . . . . . . . . . . . . Aleurothrixus floccosus (Maskell 1895) Subdorsal fold smooth, radially concentric or as a lateral pair and not joined below the vasiform orifice; marginal teeth without basal glands; caudal ridge present; lingula visible or obscured by operculum; branching comb-like structure absent . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9(8). Subdorsal pair of lateral folds extending from the ASDS to the vasiform orifice; puparium more than 2x longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bemisia giffardi (Kotinsky 1907) Subdorsal pair of lateral folds replaced by an incomplete, radially concentric fold that is interrupted by the caudal ridge; puparium less than 15x as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10(9). A8 trilobed; operculum covers most of vasiform orifice obscuring the lingula . . . . . . . . . . . . . Aleurolobus confusus David & Subramaniam 1976

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A8 not trilobed, operculum covering half of vasiform orifice exposing at least the tip of the lingula . . . . . Africaleurodes citri (Takahashi 1932) Submarginal row of papillae present; lingula tri-lobed; legs with spines . . . . . . . . . . . . . . . . . . . . . . . . . . Trialeurodes ricini (Misra 1924) Submarginal row of papillae absent; lingula not lobed, or with only a basal pair of lobes, or obscured; legs lacking spines . . . . . . . . . . . . . . 12 Submargin with 14 to 16 pairs of setae that extend well beyond lateral margin; vasiform orifice triangular; tracheal cleft without finger-like comb of teeth; base of lingula with a pair of lateral lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parabemisia myricae (Kuwana 1927) Submarginal setae absent; tracheal cleft forming finger-like comb of 4 to 5 teeth; lingula obscured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleuroplatus translucidus Quaintance & Baker 1917 Key to the puparia of whiteflies found on Syzygium

1.

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2(1). 3(2).

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4(3).

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5(2).

6(5).

Apex of all tarsi terminating in a claw; lingula spatulate and exserted from the vasiform orifice; 2 pairs of subapical setae present on the lingula; 4 pairs of abdominal compound pores, cuticle always pale . . . . . . . . . . . . . . Aleurodicinae: Aleurodicus dispersus Russell 1965 Apex of all tarsi terminating in an adhesive pad; lingula usually obscured by operculum, if visible always inserted within the vasiform orifice and not spatulate shaped; 1 pair of subapical setae present on the lingula or obscured; compound pores absent (large, simple disc pores present in Dialeuropora); cuticle pale to black. Aleyrodinae . . . . . 2 Puparium with large, thick dorsal spines; margin dentate. Aleurocanthus Quaintance & Baker 1914. . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Puparium without large, thick dorsal spines, but setae may be present . . . . 5 Puparium pale; marginal teeth with acute apices; submargin with 12–13 pairs of spines with acute apices extending well beyond the lateral margin . . . . . . . . . . . . . . . . . . . Aleurocanthus pendleburyi Corbett 1935 Puparium dark brown to black; marginal teeth not acute; submargin with short to elongated, acute to bifurcated spines some of which may extend beyond the lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Submarginal spines with bifurcate apices and not reaching the lateral margin; 31–36 pairs of dorsal spines present in the male and 42–48 pairs in the female; subdorsal spines absent, 1 submedian pair of spines present on A1 . . . . . . . . . . . . . . Aleurocanthus eugeniae Takahashi 1933 Submarginal spines with acute apices and extending to or beyond the lateral margin; 11 pairs of dorsal spines; numerous pairs of subdorsal and submedian spines present including those on A1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleurocanthus woglumi Ashby 1915 Five large pairs of submarginal disc pores present each with its diameter subequal to width of the vasiform orifice; subdorsal fold always absent. Dialeuropora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Large disc pores absent; subdorsal fold absent or present . . . . . . . . . . . 7 Large disc pores forming a smooth, radially-concentric arc on the subdorsum; tracheal fold not evident and caudal fold faint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dialeuropora mangiferae (Corbett 1935)

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9(8). 10(8).

11(10). 12(11).

13(12).

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14(13).

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Large disc pores not forming a smooth radially-concentric arc, but rather the second pair of pores offset and adjacent to the first pair of legs; tracheal fold partially evident and caudal fold well defined. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dialeuropora papillata Cohic 1966 Concentric subdorsal fold present separating submargin from subdorsum; posterior portion of vasiform orifice notched or not . . . . . . . . 8 Concentric subdorsal fold absent; vasiform orifice not notched on the posterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Pupal case elongate-oval with narrow lateral margin, submargin 2x or more in width as the margin; posterior margin of vasiform orifice not notched . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes Corbett 1935 Pupal case elliptical to pyriform with narrow margin and submargin; posterior margin of vasiform orifice notched . . . . . . . . . . . . . . . . . . . . 9 Submargin pale and dorsal disk black; submarginal row of papillae absent . . . . . . . . . . . . . . . . . . . Aleuroclava neolitseae (Takahashi 1934) Pupal case entirely pale; submarginal row of papillae present . . . . . . . . . . . . . . . . . . . Aleuroclava takahashii (David & Subramaniam 1976) Submargin with a row of barrel-shaped glands; tracheal and caudal clefts comb-like with 10–12 narrow, elongate teeth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orchamoplatus louiserussellae Martin 1999 Submargin without barrel-shaped glands; tracheal clefts if modified not forming a comb of elongated teeth . . . . . . . . . . . . . . . . . . . . . . . 11 Caudal and tracheal clefts not differentiated; caudal and tracheal furrows absent . . . . . . . . . . . . . . . . . Aleuroinanis myrtacei Martin 1999 Caudal and tracheal clefts forming distinct pores or notches; caudal and tracheal furrows present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Dorsum with a rhachis and submarginal and subdorsal rows of capitate or mushroom-shape pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhachisphora queenslandica Martin 1999 Dorsum without a rhachis and differentiated pores . . . . . . . . . . . . . . 13 Cranial suture absent; tracheal and caudal clefts with deeply invaginated pores indented away from the margin; tracheal and caudal furrows stippled ventrally . . . . Indoaleyrodes pseudoculatus Martin 1985 Cranial suture absent or present; tracheal and caudal clefts with pores that are not deeply invaginated; caudal furrow not stippled, may be reticulated or punctuated by a band of polygonal markings . . . . . . . . 14 Cranial suture absent; subdorsal pair of PSMeS(A1) present; caudal clefts with pores that are lined internally with 2–3 spine-like teeth; caudal furrow punctuated by a band of polygonal markings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rabdostigma minahassai Martin 1988 Cranial suture present; subdorsal pair of PSMeS(A1) absent; caudal clefts with pores that are not lined internally; dorsal thoracic tracheal furrows not differentiated; caudal furrow reticulated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dialeurodes indicus David & Subramaniam 1976

Genus Aleurolobus Quaintance & Baker 1914 Diagnosis. Puparium habitus. Puparium oval with cuticle usually brown to black, sometimes pale. Dorsum. Submargin separated from dorsal disk by a subdorsal fold

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that is radially concentric and interrupted by the caudal furrow; transverse molting suture terminating at the subdorsal fold; lateral margin dentate or crenulated tracheal and caudal clefts differentiated from those of the lateral margin with a comb of 3 teeth in the species found on B. koenigii; cephalothorax and abdomen with or without tubercles; papillae absent. Vasiform orifice. Elongate-cordate to roundedtriangular with orifice covering most of operculum but exposing part of the lingula; orifice surrounded by a tri-lobed raised structure Chaetotaxy. ASDS, ASMeS, PSMeS(A1), PSMeS(A8), and CS present. Venter. Tracheal furrow with or without minute spinules; legs with adhesive pads; length of antennae variable, extending to the base of first pair of legs or to the middle of the second pair of legs. Comments. Aleurolobus, an Old World genus, is comprised of 88 described species. Of these, only Aleurolobus solitarius Quaintance & Baker 1917 and A. marlatti are known to occur in the New World. The following seven species have been intercepted at US ports of entry: Aleurolobus wunni (Ryberg 1938), A. confusus David & Subramaniam 1976, A. marlatti, A. olivinus (Silvestri 1911), A. setigerus Quaintance & Baker 1917, and A. subrotundus Silvestri 1927. Six of the 88 species occur in Thailand: A. barodensis (Maskell 1985), A. marlatti, A. rhododendri Takahashi 1934, A. setigerus, A. tomkinsae and A. tuberculatus Regu & David 1993. The genus Aleurolobus was first described as a dark brown pupa with an incomplete, radially-concentric subdorsal fold, eyespots, tracheal and caudal cleft with 3 teeth present (Quaintance & Baker 1914). Since then, other species have been described ranging from pale or dark brown, with long to minute marginal, submarginal, and subdorsal setae and with or without eyespots or dentate differentiated tracheal clefts. This genus is one of the most frequently intercepted genera of whiteflies at the US ports of entry but only 5 different species have been recorded. Of these, A. marlatti is the species most frequently intercepted in quarantine, often found on Bergera leaves from India. Recently however, A. tomkinsae has become more prevalent on Bergera leaves originating from Asia, representing more than 75% of the interceptions of whiteflies on this host. Of all the described species, A. oplismeni Takahashi 1931, A. tomkinsae, and A. setigerus have elongate setae that extend to, or beyond the lateral margin. However these species differ by the arrangement of subdorsal setae and number of reduced teeth in the tracheal and caudal clefts. Aleurolobus tomkinsae Dooley & Smith-Pardo sp. nov. Figs. 1 and 2 Description. Puparium habitus (Figs. 1 and 2). Puparia brown, oval to elliptical in shape with no wax visible, found on underside of leaf, solitarily or in a groups of 2–3 puparia. Slide mounted puparium. Measurements were taken in microns (mm) from 17 specimens of the type series with the range of measurements followed by the average measurement in brackets. Pupal case 626–1082 [877] long by 365–777 [616] wide, holotype 1005 long by 731 wide. Dorsum. Lateral margin dentate, teeth short with blunt apices; tracheal and caudal clefts differentiated from lateral margin by 3 teeth in each cleft; submargin with transverse striae from the lateral margin to the dorsal disk. Longitudinal molting suture extending beyond the dorsal fold to the anterior margin of the cephalothorax; transverse suture extending to the dorsal fold. A pair of large, crescent-shaped eyespots present subdorsally on the cephalon. Subdorsal fold separating the dorsal disk from the submargin, radially concentric, incomplete,

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Figure 1. Illustration of habitus and diagnostic characters of Aleurolobus tomkinsae. a) Habitus. b) Cephalic setae. c) Thoracic cleft. d) Vasiform orifice, operculum, caudal fold and setae.

interrupted below the vasiform orifice by the caudal furrow. Vasiform orifice. Length 62–105 [86] by 47–81 [68] wide, holotype 95 long by 72 wide. Vasiform orifice elongate cordate with operculum not completely covering the orifice, lingula elongate, apically wider, not exposed; large tri-lobate sclerotized area surrounds the orifice. The distance from the caudal margin to the posterior border of the vasiform orifice 46–90 [74], holotype 89 from the caudal margin to the posterior margin of the orifice which is shorter than the length of the vasiform orifice. Pores. Paired geminate pore with associated porette present from the submargin to submedian and cephalon to the caudal submargin below the vasiform orifice, with sclerotized margin and the porettes separated by no more than the 1.5 times the diameter of the larger pore. Number of pores as follows: ASMP 31–39 [34], PSMP 28–43 [36], ASDP 12–24 [20], ASMeP 8–10 [8], PSMeP 18–20 [18], PSDP 14–28 [20]. Submarginal pores and porettes are on the apical border near the margin and arranged in a single row although 2 or 3 anterior pores may be in a double row. Chaetotaxy. Cephalothorax (C1 to T3) with 5 pairs of anterior marginal, subdorsal (inner border of the subdorsal fold), and submedian setae with the following lengths: AMS 17–35 [25], that of holotype 23; ASDS(1) 49–99 [72], holotype 93; ASDS(2) 34–94 [72], with that of holotype 76 placed just anterior to the eyespot; ASDS(3) 17–95 [65], holotype 84 located opposite the T2 pair of legs; and ASMe(1) 14–66 [31], holotype 20 located lateral to the anterior border of the rostrum. Abdominal (A1 to A8) pairs of anterior marginal, subdorsal (inner border of the subdorsal fold), and submedian setae

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Figure 2. Digital image composite of habitus and diagnostic characters of Aleurolobus tomkinsae. A) Habitus. (puparium). B) Habitus (nymph). C) Posterior abdominal setae, vasiform orifice, caudal furrow and cleft. D) Two anterior cephalic setae associated with fold and tracheal cleft. E) Thoracic tracheal cleft and subdorsal seta.

(dorsal and ventral) with the following lengths in mm: PMS 7–46 [30], holotype 14; PSMeS (A1) 11–87 [48], holotype 14; PSDS(4) 48–94 [72], holotype 94; PSDS (5) 36– 79 [59], holotype 63; PSMS(6) 12–88 [52], holotype 70; PSMS(7) 12–81 [39], holotype 54; PSMS(8) 4–38 [19], broken off on holotype; PSMeS(A1) 19–98 [53], holotype 60; PDSMe(A8) 7–28 [11], holotype 7; PVSMe(A8) 16–34 [26],holotype 28 and 29; CS 35–64 [52], holotype 52. Venter. Antennae reaching the anterior margin of the mesothoracic legs. Legs with adhesive pads. Tracheal furrows stippled forming a band along the outer margin of T2 and T3 legs; caudal furrow narrow and granular. Nymph (2nd instar) Habitus. Nymph light brown in color, Slide mounted puparium. Puparium 457 long by 316 mm wide. Lateral margin dentate with 12 teeth within 100 mm. Subdorsal fold absent. Vasiform orifice. Length 48 by 50 mm wide; distance from vasiform orifice to caudal margin 24 mm long. Pores. Pores and porettes distributed over cuticle. Chaetotaxy. ASMe 15 long, located lateral to the anterior margin of the rostrum, PSDS (A1) 12 mm long, CS 19 mm long. Lingula with apical pair of setae (19 mm long) extending beyond vasiform orifice. Venter. Legs one segmented, cone-shaped with a terminal adhesive pad. Distribution. Indomalayan region: India, Sri Lanka, and Thailand. Host. Bergera koenigii (Rutaceae), Magnolia sp. (Magnoliaceae), and undetermined leaves.

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Material Examined. Holotype: Puparium intercepted on Bergera koenigii (from India in passenger baggage at Chicago, Illinois Port of entry), on 3-iv-2012, M. Choudhary, deposited at the USNM (slide# apcil120943571001).

Paratypes: 17 Puparia (4th instar) and 1 nymph (2nd instar). 1)

2)

3)

4)

On Bergera koenigii leaves from India intercepted in passenger baggage on 21-v201, slide# apcil111426507001 deposited at the USNM: slide #apcil120943571002 (4 puparia) on 3-iv-2012 and apsca092385965001 (puparium), on 24-viii-2009 deposited at the PPQC; slide# apcil120957298002 (puparium) on 4-iv-2012 and apaga120946190002 (2 puparia) 3-iv-2012, deposited at CSCA; slide# apcil1686507002 (puparium) on 4-iv-2012 and apcil111386507010 (2 puparia) on 17-v-2011, deposited at CASC. On indeterminate leaf from India, intercepted in passenger baggage, 21.i.2004: slide# atlga041771019131 (puparium and 1 nymph), deposited at the PPQC and atlga041771019131, deposited at USNM. On Bergera koenigii leaves from Sri Lanka intercepted in passenger baggage in quarantine: slide# chiil031470240250 (2 puparia) on 15-v-2003, deposited at the USNM. On Magnolia sp. leaves from Thailand, intercepted in quarantine at the USDA Plant Inspection Station in San Francisco, CA. Coded as APWCA112442638002 (2 slides) on 25-viii-2011, deposited at the USNM.

Etymology. This species is named after Marilyn Tomkins, plant safeguarding officer at USDA-APHIS-PPQ inspection station in South San Francisco, who found the species on magnolia plants imported from Thailand. Comments. Aleurolobus tomkinsae closely resembles A. marlatti, A. oplismeni (recorded on Oplismenus sp. Poaceae), and A. setigerus (recorded on Citrus, Rutaceae). Only A. marlatti has been reported on Bergera or Murraya. Aleurolobus marlatti, A. setigerus, and A. tomkinsae are similar in that they have a black cuticle with the submargin marked with suture-like lines that extend to the subdorsal fold, crescent-shaped eyespots, and the tracheal and caudal clefts each with 3 teeth. The submarginal setae are long in the drawing of A. niloticus (junior synonym of A. marlatti) from Egypt by Priesner & Hosny (1934), but those setae only extend no more than1/2 of the way towards but do not reach the margin. Aleurolobus oplismeni is similar to A. tomkinsii in that it is pigmented dark, possess a ring of setae that extend beyond the pupal margin, has tracheal cleft with teeth, and reniform eyspots. Aleurolobus oplismeni differs in that the long setae extending beyond the margin originate on the subdorsal fold only and not the submargin and that the tracheal thoracic clefts have 4–6 teeth and the caudal cleft with 4 teeth, not three. Aleurolobus tomkinsae differs from the other species by having three subdorsal setae (ASDS) on the cephalothorax, two abdominal setae along the subdorsal fold (at A4 and A5) that extend beyond the pupal margin, and three submarginal setae (PSMS at A6, A7 and A8 with A6 also extending beyond pupal margin); The length of the vasiform orifice is greater than the distance from the posterior margin of the vasiform orifice to the caudal margin. The average ratio is 0.9 (distance from the orifice to the margin proportional to the length of the vasiform orifice). Two exceptions are noted showing the distance and length being subequal: one specimen from Sri Lanka and one from India have a subequal ratio of 1.03 and 1.06 similar to A. marlatti where the length of the orifice is slightly less by 2 and 7 mm.

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Aleurolobus marlatti has the length of the vasiform orifice less than the distance from the posterior border of the vasiform orifice to the caudal margin (Dubey & Ko 2009) as opposed to greater than in A. tomkinsae. This taxon has minute subdorsal and submarginal setae except for the long pair on the last abdominal segment and the long caudal setae, and lacks the submedial pair of cephalothoracic (T1) setae. Aleurolobus setigerus differs from the other two taxa by the presence of two pairs of cephalic subdorsal setae, two pairs of submarginal thoracic setae (the posterior one adjacent to the transverse suture), and six pairs of submarginal abdominal setae that are centered between the margin and the dorsal fold. The subdorsal and submarginal setae extend beyond the pupal margin. The shape of the operculum is consistent with the other species, except the apex is acute whereas in the other two taxa the apex of the operculum is slightly rounded. Measurements of the puparium and other structures of A. tomkinsae indicate that specimens from different countries and host plants vary in size. Puparia collected on B. koenigii from Sri Lanka were the largest, ranging from 1029–1082 (2 puparia) mm in length; those on this host from India were 739–1005 mm (11 puparia); and those on Magnolia sp. from Thailand were the smallest, ranging from 626–721 mm (2 puparia) in length. Asialeyrodes Corbett 1935 Diagnosis. Puparium habitus. Broadly oval in shape, all known species pale white or yellow, except A. meghalayensis Regu & David 1991, which is dark brown. Slide mounted puparium. Submarginal area wide and separated from the dorsum by a subdorsal fold. Dorsum. Conspicuous papillae absent; tracheal and caudal clefts and folds evident. Chaetotaxy. ASMeS and PSMeS(A1) present. Vasiform orifice. Vasiform orifice small, subcordate and without teeth, operculum similar in shape to vasiform orifice, filling vasiform orifice or nearly so, lingula obscured; orifice not surrounded by a trilobed area. Comments. This is an Old World genus (Indomalayan, Austro-Indomalayan regions) with 15 described species. The genus Asialeyrodes is similar to Cockerelliella Sundararaj & David 1991 and Pseudcockerelliella Sundararaj 2007; however, the latter 2 genera have the cephalothoracic fold joining the transverse and longitudinal sutures into one structure forming an arched, trapdoor- like structure on the cephalothorax, and the submargin is narrow; whereas, Asialeyrodes has a wide submargin and a complete submarginal fold that separates the entire submargin from the dorsal disk that is interrupted only by the caudal furrow. The genus Asialeyrodes was first described as having no distinct pores or papillae and with distinct tracheal and caudal folds. This description was also cited and accepted by Sundararaj & David (1991). Only four species in the genus are now known to have rows or clusters (groups) of papillae present on the cephalon or thorax extending in some species to the vasiform orifice. Several species are described with distinct pores and porettes and few without distinct tracheal folds. Several authors use papillae and tubercles to define the same structures. Also one species, Asialeyrodes maesae Takahashi 1934, is described as having both the longitudinal and transverse molting sutures not reaching the margin of the subdorsal disk. Key to the puparia of the species of Asialeyrodes 1.

Puparium pale; lacking a submarginal row of setae; tracheal pore deeply invaginated to indistinct; with submedian depressions on thorax and abdomen or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

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2(1). 3(2).

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4(3).

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5(2). 6(5).

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7(5). 8(7).

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9(7). 10(1). -


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Puparium pale to black; with a submarginal row of setae; tracheal pore indistinct to marginal but not deeply invaginated; submedian depressions absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Submedian papillae-like structures present; submedian depressions absent or present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Submedian papillae and submedian depressions absent. . . . . . . . . . . . . 5 Submedian papillae in groups present from T1 to A8 with T1 and T3 submedian papillae enlarged; submedian row of depressions absent; PSMeS(A8) 9.25 mm long; CS absent; tracheal and caudal furrows distinct with polygonal markings; caudal and tracheal ventral folds stippled . . . . . . . . . . .Asialeyrodes splendens Meganathan & David 1994 Submedian row of papillae present from C1 to A8 of subequal size in a uniform row; submedian depressions present or absent; PSMeS(A8) present or absent; CS present; caudal and tracheal furrows absent; ventral caudal and tracheal folds either absent or smooth, not marked. . . 4 Submedian row of depressions present from C1 to A8; PSMeS(A8) absent; tracheal pore deeply invaginated with internal teeth; thoracic and caudal furrows not indicated; ventral caudal fold smooth (not marked); ventral tracheal fold not indicated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes dubius Martin & Mound 2007 Submedian row of depressions absent; PSMeS(A8) present; tracheal pores not deeply invaginated (within margin) without teeth; caudal furrow marked with stipples; tracheal folds and furrows not indicated . . . . . . . Asialeyrodes lateropilliformis Dooley & Smith-Pardo sp. nov. Ventral caudal fold not defined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Ventral caudal fold faint to marked with dots or polygonal structures . . . 7 Tracheal pores indistinct; dorsal pores scattered but not paired; vasiform orifice with operculum covering half the orifice, lingula with knobbed apex that extends to the posterior margin of the orifice . . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes euphoriae Takahashi 1942 Tracheal pores round, chitinized and without teeth; pores paired; vasiform orifice with operculum covering the orifice and obscuring the lingula . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes lushanensis Ko 1993 Ventral caudal fold marked with polygonal structures . . . . . . . . . . . . . 8 Ventral caudal fold marked with dots . . . . . . . . . . . . . . . . . . . . . . . . . 9 Ventral thoracic tracheal folds distinct; ventral fold faint; dorsal caudal furrow with polygonal sculpturing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes menoni Meganathan & David 1994 Ventral thoracic tracheal folds absent; ventral caudal fold marked with polygonal sculpturing but not on dorsal caudal furrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes dorsidemarcata (Singh 1932) Ventral thoracic tracheal fold distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes multipori Takahashi 1942 Ventral thoracic tracheal fold indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Asialeyrodes corbetti Takahashi 1949 Submarginal furrow with 3–4 rows of papillae; A1 seta absent . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes papillatus Regu & David 1991 Submarginal furrow lacking papillae; A1 seta present. . . . . . . . . . . . . 11

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Figure 3. Illustration of habitus and diagnostic characters of Asialeyrodes lateropapilliformis. A) Habitus. B) Thoracic spiracle. C) Detail of sculpturing. D) Operculum and caudal furrow.

11(10).

Longitudinal and transverse sutures either terminate before or extend to the submarginal fold; submedian papillae absent; ASMeS and PSMeS (A1) with either pointed or capitate apices; submarginal setae less than 49 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Longitudinal and transverse sutures terminate at the submarginal fold; submedian papillae present; C1 and A1 setae with pointed apices; submarginal setae from 49 to 70 mm long . . . . . . . . . . . . . . . . . . . . . 13 12(11). ASMeS and PSMeS (A1) with pointed apices; transverse and longitudinal sutures terminate well before submarginal fold; cephalothorax with no . . . . . . . . . . . . . . Asialeyrodes maesae (Takahashi 1934) ASMeS and PSMeS (A1) with capitate apices; transverse and longitudinal sutures reaches submarginal fold; cephalothorax with sutures . . . . . . . . . . . . . . . Asialeyrodes indica Sundararaj & David 1991 13(11). Puparium pale and oval with a smooth lateral margin; peripheral row of tubercles extend from T1 to A7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes elegans Meganathan & David 1994 Puparium dark brown and subcircular with a crenulate lateral margin; peripheral row of tubercles absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asialeyrodes meghalayensis Regu & David 1991

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Figure 4. Digital image composite of habitus and diagnostic characters of Asialeyrodes lateropapilliformis. A) Habitus. B) Series of papillae and legs. C) Caudal region with marginal seta and caudal furrow. D) Vasiform orifice. Caudal cleft and furrow. E) Cephalothorax with dorsal fold, series of papillae and marginal setae. F) Abdomen with dorsal fold and series of papillae.

Asialeyrodes lateropapilliformis Dooley & Smith-Pardo sp. nov. Figs. 3 and 4. Description. Puparium habitus (Figs. 3 and 4). Puparium broadly oval in shape and pale in color. Slide mounted puparium. Dimensions are taken from 4 specimens of the type series in microns (mm). Pupal case dimensions ranged from 799–991 long [925] by 630–809 wide [743, that of holotype measuring 991 long by 809 wide. Margin crenulated; tracheal and caudal clefts present appearing as pores; tracheal furrows indistinct, caudal furrow narrow and marked with granules, distance from posterior margin of vasiform orifice to posterior margin of puparium 138–163 with that of the holotype 43 long. The ratio of the orifice length to the distance from the posterior orifice margin to the caudal margin is 0.16–0.19 [0.18] with that of the holotype 0.18. Dorsum. Longitudinal molting suture terminates at the subdorsal fold not reaching margin; transverse suture terminating before the subdorsal fold, just lateral to area above the 3rd pair of legs. Subdorsal fold wide (125–142) and composed of overlapping, scale-like structures separating the dorsum from the wide submargin and forming a complete, radially concentric ring that encompasses the puparium, only interrupted posteriorly by the narrow caudal furrow. Dorsum with a narrow band of papillae extending from the area on the dorsal surface above and lateral to the mouthparts, which then overlaps the legs, and extends posteriorly to the vasiform orifice. Pores. Paired geminate pore with associated porette present with sclerotized margin and separated by no more than the diameter of the pore; scattered on dorsum with submarginal series in 2–3 irregular rows from the cephalon to below the vasiform orifice. Vasiform orifice. Orifice subcordate with lateral and posterior thick margins but not notched on the posterior margin, wider than long, ranging from 25– 26 long by 34–39 wide with that of the holotype measuring 26 long and 38 wide. Caudal fold marked with granules, area below the posterior margin of the vasiform orifice wide, and narrow at its posterior apex. Distance from the posterior margin of

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the orifice to the submarginal band (29) is about half the distance from the caudal pore to the submarginal fold (73). Distance from the caudal fold to the posterior margin of the vasiform orifice ranges from 137–163, that of holotype measuring 143 long. Operculum fills more than 70% of orifice. Lingula obscured by operculum. Chaetotaxy. Cephalic ASMeS, PSMeS(A1), PSMeS(A8), and CS present with the CS adjacent to the caudal furrow. ASMS broken off of all specimens. PSMeS(A1) ranging from 8–13 [10.5] long on paratype with PSMeS(A1) broken off on holotype; PSMeS(A8) is 10.2 long on paratype and PSMeS(A1) broken off on holotype , and CS ranged from 4.8–12 [7.3] long with the holotype measuring 4.8 and 7.2 long and a distance from the caudal margin of 40.6; AMS 14–17 [15] long with holotype 14 long; PMS adjacent to the caudal furrow is 22–23 [22.5], those of holotype are broken off. Venter. Submedian pair of mesothoracic papillae present; caudal furrow stippled, wide basally just below that posterior margin of the vasiform orifice tapering narrowly from the subdorsal fold to the caudal cleft; legs with adhesive pads. Nymph (2nd instar). Habitus: Oval in shape and pale in color. Slide mounted puparium. Dimensions: 510 long by 359 wide. Margin crenulated; tracheal cleft appearing as a closed pore and caudal cleft appearing as an elongated plate with caudal and tracheal furrows indistinct. Dorsum. Patterned with roughened cuticle giving a scalloped appearance. Longitudinal and transverse molting suture terminates at margin just below tracheal pore. Subdorsal fold absent. Vasiform orifice (19 long and 28 wide) small with lateral and posterior thick margins, subcordate, not notched on the posterior margin. Lingula obscured, operculum 15 long by 22 wide; ratio of length of orifice to distance from vasiform orifice to caudal margin is 0.35. Chaetotaxy. All setae broken off; ASMeS and PSMeS(A1) AMS, and PMS setal bases present. Venter. Legs one-segmented, cone-shaped each with a terminal adhesive pad; oriented toward the margin. Comments. Asialeyrodes lateropapilliformis is similar to A. dubius in that both species lack the submarginal row of setae, and differs from A. dubius by having lacking the submedian row of depressions that are sunken or depressed below the surrounding area A8 seta present, and the tracheal pore pores not deeply invaginated and without teeth. Asialeyrodes lateropapilliformis is similar to A. papillatus. Both species have tracheal pores with a chitinized rim, molting sutures terminating at the submarginal fold, a chain of papillae that extends from the cephalon to A1 segment overlapping the legs, submargin and dorsal disk with numerous pairs of pores and associated porettes, and the caudal furrow granular and narrower than the width of the orifice A. lateropapilliformis differs from A. papillatus by having a uniform chain of papillae that extends from the cephalon to the vasiform orifice, submarginal row of setae absent, rows of papillae forming within the submarginal fold absent, PSMeS(A1) pair present, and the abdominal sutures 1–6 not marked with papillae-like structures; the latter has a uniform chain of papillae terminating at A1, submarginal row of setae present, the submarginal fold with 3–4 rows of papillae-like structures, PSMeS(A1) setae absent, and abdominal sutures 1–6 marked with papillae-like structures. Asialeyrodes lateropapilliformis is similar to A. splendens in that both species have a row of papillae extending to the vasiform orifice overlapping the legs, lack a row of submarginal setae, caudal furrow marked (granulated), and the presence of ASMeS, PSMeS(A1), and PSMeS(A8) setae. The former differs from the latter in that the papillae are in a uniform row, a pair of enlarged papillae on the prothorax and

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metathorax absent, tracheal dorsal furrows/ventral folds not differentiated, and a pair of CS present adjacent to the caudal furrow; whereas the latter has papillae grouped in clusters (not a uniform row), with a pair of enlarged papillae on the prothorax and metathorax, tracheal and caudal folds stippled and tracheal furrows marked with distinct polygonal structures. Asialeyrodes lateropapilliformis differs from A. elegans and A. meghalayensis, with the former lacking a row of submarginal setae whereas the latter two species have a row of submarginal setae. In addition, A. meghalayensis puparia are brown to black whereas the other two species are pale. Distribution. The species is currently only known to occur in Indonesia (Austrooriental region). Host. The only recorded host for this species is Syzygium sp. (Myrtaceae). Material Examined. Holotype: puparium, intercepted in quarantine at the USDA Plant Inspection Station in San Francisco, CA. 23-viii-2011, on Syzygium sp. from Indonesia, deposited at the USNM (Slide# APWCA112366214004a),

Paratypes (3 slides): 4 puparia and 1 nymph (2nd instar nymph, same data as holotype, deposited at the PPQC (Slide# APWCA112366214004b-d). Etymology. This species is named for its narrow band of tubercles that extends from the cephalothorax to the area laterad of the vasiform orifice. CONCLUSIONS These newly described species (Aleyrodinae) further support the fact that whitefly diversity may be greater in tropical areas compared to that in more temperate and cooler regions (Bink-Moenen & Mound 1990). Whiteflies are frequently intercepted in quarantine on shipments of plant material originating from countries throughout the world, but especially from tropical countries. The introduction of an exotic species into areas outside its native range, separated from its natural enemies, has often led to pest outbreaks that threaten agricultural crops and international trade of propagative commodities. ACKNOWLEDGMENTS We would like to recognize Marilyn Tomkins, safeguard specialist at the USDA Plant Inspection Station, for intercepting A. tomkinsae in quarantine and Mr. Arthur Berlowitz, Officer-In-Charge of the Plant Inspection Station in San Francisco, California, for his continuous support and encouragement to describe the two new quarantine species. We are grateful to Dr. Greg Evans, Mr. Ray Gill and Luis Valencia for reviewing this manuscript. LITERATURE CITED Ashby, S. F. 1915. Notes on diseases of cultivated crops observed in 1913–1914. Bulletin of the Department of Agriculture, Jamaica 2:299–327. Bink-Moenen, R. & L. Mound. 1990. Whiteflies: diversity, biosystematics and evolutionary patterns, pp. 1–11. In: D. Gerling (Ed.). Whiteflies their Bionomics, Pest Status and Management. Intercept Ltd., United Kingdom, 348 pp. Cohic, F. 1966. Contribution a l’etude des aleurodes africains (1e Note). Cahiers de l’Office de la Recherche Scientifique et Technique Outre-Mer (Serie Biologie) 1:3–59. Corbett, G. 1935. Malaysian Aleyrodidae. Journal of the Federated Malay States Museum 18(4):733–739, 841–842. David, B. V. & T. R. Subramaniam. 1976. Studies on some India Aleyrodidae. Records of the Zoological Survey of India 70:133–233.

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Dooley, J. 2011. Whitefly Pupa of the World. Plant Protection and Quarantine, United States Department of Agriculture. Available from http://keys.lucidcentral.org/keys/v3/whitefly/ Default.htm (accessed 6 May 2012) Dubey, A. K. & R. Sundararaj. 2006. Key to whiteflies of the tribe Aleurolobini (Hemiptera: Aleyrodidae) of India with description of five new species. Oriental Insects 40:33–60. Dubey, A. K. & C. C. Ko. 2009. A review of the genus Aleurolobus Quaintance & Baker (Hemiptera: Aleyrodidae), based mainly on pupal morphology with a description of a new species. Entomological Science 12:51–66. Evans, G. A. 2008. Whitefly Taxonomic and Ecological Website. Plant Protection and Quarantine, United States Department of Agriculture. Available from http://www.sel.barc.usda.gov:8080/ 1WF/whitefly_catalog.htm (accessed 12 October 2012) Gill, R. 2012. A preliminary report on the World species of Bemisia Quaintance & Baker and its congeners (Hemiptera: Aleyrodidae) with a comparative analysis of morphological variation and its role in the recognition of species. Insecta Mundi 0219:1–99. Kinard, G. 2009. Grin Taxonomy for Plants. United States Department of Agriculture (USDA), Agricultural Research Service (ARS). Available from http://www.ars-grin.gov/cgi-bin/npgs/ html/taxgenform.pl (accessed 1 October 2012) Ko, C. C. 1993. Aleyrodidae of Taiwan. Part II. Asialeyrodes Corbett. Japanese Journal of Entomology 61(3):613–618. Kotinsky, J. 1907. Aleyrodidae of Hawaii and Fiji with descriptions of new species. Bulletin, Board of Commissioners of Agriculture and Forestry Hawaii, Division of Entomology 2:93–102. Kuwana, I. 1927. On the genus Bemisia [Family Aleyrodidae] found in Japan, with description of a new species. Annotationes Zoologicae Japonensis 11:245–253. Martin, J. 2007. Giant whiteflies (Sternorrhyncha, Aleyrodidae): a discussion of their taxonomic and evolutionary significance, with the description of a new species of Udamoselis Enderlein from Ecuador. Nederlandse Entomologische Vereniging 150:13–29. Martin, J. H. 1985. The whitefly of New Guinea. Bulletin of the British Museum (Natural History) (Entomology) 50(3):303–351. Martin, J. H. 1988. Whitefly of northern Sulawesi, including new species from clove and avocado (Homoptera: Aleyrodidae). Indo-Malayan Zoology 5:57–85. Martin, J. H. 1999. The whitefly fauna of Australia (Sternorrhyncha: Aleyrodidae). A taxonomic account and identification guide. Technical Paper, Division of Entomology, Commonwealth Scientific and Industrial Research Organization, Canberra 38:1–197. Maskell, W. M. 1895. Contributions towards a monograph of the Aleurodidae, a family of Hemiptera -Homoptera. Transactions of the New Zealand Institute 28:411–449. Meganathan, P. & B. V. David. 1994. Aleyrodidae fauna (Aleyrodidae: Hemiptera) of Silent Valley, a tropical evergreen rain-forest, in Kerala, India. Fippat Entomology 5:1–83. Mifsud, D. & V. Palmeri. 1996. A new species of Aleurolobus Quaintance et Baker (Homoptera, Aleyrodidae) from southern Europe. Bollettino Del Laboratorio Di Entomologia Agraria Filippo Silvestri 52:89–95. Misra, C. S. 1924. The citrus whitefly, Dialeurodes citri in India and its parasite, together with the life history of Aleurodes ricini, n. sp. Proceedings Fifth Entomology Meeting, Pusa, 1923. Indian Agriculture, Pusa, Sir-Lanka: 129–135. Priesner, H. & M. Hosny. 1934. Contributions to a knowledge of the whiteflies (Aleurodidae) of Egypt (III). Bulletin of the Ministry of Agriculture. Egypt. Technical & Scientific Services 145:1–11. Pushpa, R. & R. Sundararaj. 2009. Whiteflies of the genus Cockerelliella Sundararaj and David (Hemiptera). Biosystematica 3(2):29–36. Regu, K. & B. V. David. 1993. Taxonomic studies on Indian aleyrodids of the tribe Aleurolobini (Aleyrodinae: Aleyrodidae: Homoptera). Frederick Institute of Plant Protection & Technology, Entomology Series 4:1–17. Russell, L. M. 1965. A new species of Aleurodicus Douglas and two close relatives (Homoptera: Aleyrodidae). Florida Entomologist 48:47–55. Quaintance, A. L. 1903. New Oriental Aleurodidae. Canadian Entomologist 35:61–63. Quaintance, A. L. & A. C. Baker. 1914. Classification of the Aleyrodidae. Part II. United States Department of Agriculture Technical Series Bulletin 27:95–109. Sundararaj, R. & B. David.V. 1991. On the genera Asialeyrodes Corbett and Cockerelliella n. gen. from India. Journal of the Bombay Natural History Society 88:82–87.

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Takahashi, R. 1931. Some white-flies of Formosa (Part I.). Transactions of the Natural Histtory Society of Formosa 21:203–209. Takahashi, R. 1932. Aleyrodidae of Formosa, Part I. Report. Department of Agriculture. Government Research Institute, Formosa 59:1–57. Takahashi, R. 1933. Aleyrodidae of Formosa Part II Report, Department of Agriculture Government Research Institute, Formosa 60:20–21. Takahashi, R. 1934. Aleyrodidae of Formosa, Part III. Department of Agriculture Government Research Institute, Formosa 63:39–71. Takahashi, R. 1942. Some foreign Aleyrodidae (Homoptera) VI. Species from Thailand and French Indo-China). Transactions of the Natural History Society of Formosa 32:204–216. Takahashi, R. 1949. Some Aleyrodidae from Riouw Islands (Homoptera). Mushi 20:47–53.

Received 21 Dec 2012; Accepted 13 Mar 2013 by J. N. Zahniser; Publication date 9 July 2013.