An account of the mealybug genus Paraputo Laing (Hemiptera: Coccoidea: Pseudococcidae) in the Pacific region

The mealybug genus Paraputo is discussed from the Pacific region and three new species, P. aracearum sp. nov., P. chimbuensis sp. nov., and P. guadalcanalae sp. nov., are described and illustrated. They are discussed with two species, P. kukumi Williams and P. leveri (Green), already known from the area. Most of the species are found on economic plants and one of the mealybugs, P. leveri, has been recorded as possibly invasive. A key to adult females is provided.


Introduction
Over 150 species of Pseudococcidae are known to occur in the Pacific region north of New Zealand, based on the published works by Zimmerman (1948) for Hawaii, Beardsley (1966) for Micronesia in its broad sense, Williams and Watson (1988) for the tropical South Pacific region, Ben-Dov (1994) in a catalogue of world species, and on a few shorter papers published recently. Despite the small size of some of the islands, there is remarkable endemicity and about 100 species have not been found in other regions so far. Almost 30 species recorded from the Pacific region, either endemic or otherwise, have been listed as invasive species that are pests or a threat to US agriculture (Miller et al. 2002).
The genus Paraputo Laing, is known to occur in the New World, southern Asia, China, and the Pacific region (Ben-Dov 1994) and, in an account of the mealybugs of southern Asia, Williams (2004) redefined the genus and discussed 44 species, commenting that the species show wide variation of characters. Only two species have been recorded so far from the entire Pacific region (P. kukumi and P. leveri), although Williams (2004) also recorded P. leveri from Java, Sabah, and Thailand. P. leveri often feeds on roots, sometimes causing damage, and appears to be extending its range. It is one of the invasive species listed by Miller et al. (2002) mentioned above and has been intercepted in the USA at port inspection. P. kukumi is also extending its range in the Pacific area and, already one of the new species, P. aracearum, has been intercepted on two occasions at ports in the USA.
Without adequate inspection services, these insects can easily be introduced accidentally on plant material.
Three additional species possessing the characteristics of Paraputo have been found in the Pacific region, mostly on economic plants, and are discussed herein, thus expanding our concepts of the genus. The description and illustration of the third-instar female of P. kukumi are also included to show that the cerarii can be counted without difficulty in this instar but appear to form a continuous row in the adult female when they are not so easily recognizable.
A key to all five species, based on the characters of the adult female, is now provided.

Methods
Descriptions are based on slide-mounted specimens. The specimens were prepared using the methods of Williams and Granara de Willink (1992) and Watson and Chandler (1999), and terminology follows that of Williams (2004). Each figure shows the entire insect with the dorsum on the left and the venter depicted on the right. Enlargements of some important characters are shown on each side of each illustration.
The abbreviations of depositories used in the text are: BMNH, The Natural History Museum, London, UK; USNM, The United States National Museum of Natural History (slide collections housed at USDA, Beltsville, Maryland, USA).
( Figure 1) Ventral surface with flagellate setae similar to those on dorsum. Multilocular disc pores, each about 7.5 mm in diameter, present posterior to vulva and in more or less double rows at medial posterior edges of abdominal segments V-VII, a few also present medially on abdominal segment IV. Trilocular pores evenly distributed, numerous. Discoidal pores same as on dorsum, scattered. Oral collar tubular ducts all about same size, each narrower than a trilocular pore, distributed across abdominal segments IV-VI and present in marginal groups on abdominal segments III-VII and on inner edges of anal lobes.

Etymology
The name is based on the botanical family name Araceae and is used in the genitive plural.

Comments
This species is almost identical to P. leveri but all the dorsal setae of the adult females are noticeably longer, 40-75 mm long, whereas in P. leveri the dorsal setae are 17.5-25.0 mm long. Furthermore, the translucent pores in the adult females of P. aracearum are present only on the hind coxae, whereas there are also translucent pores on the hind femora and tibiae in P. leveri. The holotype is mounted on the same slide as three paratypes and is clearly marked and mapped on the left label. long, 6.25 mm wide at base, with a concentration of trilocular pores, on a membranous area. Anterior cerarii similar but each cerarius with anterior and posterior groups of conical setae and trilocular pores so that cerarii appear as a continuous row. Dorsal surface with numerous flagellate setae present, mostly 30-60 mm long, except for long setae 50-75 mm long, concentrated medially on abdominal segment VIII and straddling anal ring. Multilocular disc pores absent. Trilocular pores abundant, evenly dispersed. Discoidal pores, each smaller than a trilocular pore and with a thick sclerotized rim, scattered.
Ventral surface with slender flagellate setae. Multilocular disc pores, each about 7.5 mm in diameter, present medially posterior to vulva and on abdominal segments VI and VII. Trilocular pores evenly distributed, not as numerous as those on dorsum. Discoidal pores same as on dorsum, scattered. Oral collar tubular ducts present, of two sizes. A small type of duct, narrower than a trilocular pore and about as long as diameter of a multilocular disc pore, situated across medial areas of abdominal segments IV and V and submedially on abdominal segment VI, a group also present posterior to vulva between anal lobes, and one or two present on abdominal segment III. A larger type of duct, about as wide as a trilocular pore, present in small marginal groups on abdominal segments and extending anteriorly around body margins to area between antennae.

Material examined
Holotype: adult R, Papua New Guinea, Chimbu Province, Karimui, on rhizomes and inside stems of Elettaria cardamomum (cardamon) (Zingiberaceae), 22 October 1982 (coll. B. M. Thistleton) (BMNH). Paratype: Papua New Guinea, same data as holotype, one adult R (the left specimen on the same slide as the holotype and clearly marked and mapped on the right label) (BMNH).

Etymology
The name is based on the locality ''Chimbu'' with the Latin suffix ''-ensis'' indicating origin.

Comments
In possessing 18 pairs of cerarii but the cerarii appearing as a continuous row with the addition of anterior and posterior groups of conical setae and trilocular pores to each cerarius, and in possessing short dorsal setae, P. chimbuensis is related to P. danzigae Williams described from Sabah. P. chimbuensis differs from P. danzigae in possessing ventral oral collar tubular ducts around the entire margin but in P. danzigae, there are groups of oral collar tubular ducts on the posterior abdominal segments only. Moreover, there are numerous translucent pores on the hind coxae in P. danzigae and they are lacking on the hind femora and tibiae. In P. chimbuensis, they are absent on the hind coxae and are present on the hind femora and tibiae. In the key to species of Paraputo of southern Asia presented by Williams (2004), P. chimbuensis keys to P. glycosmis Williams but the cerarii of P. glycosmis are distinct and do not appear as a continuous row. Also there are noticeable long dorsal setae, many 200 mm long, in P. glycosmis, whereas in P. chimbuensis they are mostly only 30-60 mm long. Dorsal surface with numerous, long flagellate setae present, 50-80 mm long; some noticeably longer setae, mostly 125 mm long, present on each side of anal ring and a few setae, each about 40 mm long, situated anterior to anal ring. Multilocular disc pores absent. Trilocular pores numerous, evenly distributed. Discoidal pores, each a little smaller than a trilocular pore and with wide sclerotized rim, scattered.
Ventral surface with slender flagellate setae, similar in length to those on dorsum. Multilocular disc pores absent. Trilocular pores not as numerous as those on dorsum, evenly dispersed. Discoidal pores, same as on dorsum, scattered. Oral collar tubular ducts, each narrower than a trilocular pore, present medially near posterior edges of abdominal segments V and VI and in small marginal groups between abdominal segments VI and VII and segments VII and VIII.

Etymology
The name is based on the locality ''Guadalcanal'' with the Latin genitive indicating ''of'' or ''from''.

Comments
This is an unusual species of Paraputo in lacking multilocular disc pores entirely on the venter and in possessing only nine pairs of cerarii. There is a single frontal pair of cerarii and the other eight pairs occur on the abdomen. P. guadalcanalae is probably related to P. szemaoensis (Borchsenius) described from China by Borchsenius (1960), a species with only abdominal cerarii but lacking the frontal pair. Besides, P. szemaoensis possesses ventral multilocular disc pores but these are absent in P. guadalcanalae. Both species possess numerous anal ring setae.
The type species of Paraputo. P. ritchiei Laing, an African species described by Laing (1929) and discussed by Ferris (1955) (5Ripersia anomala Newstead), also lacks cerarii on the thorax, as in P. guadalcanalae, but differs in possessing two pairs of cerarii on the head and multilocular disc pores on the venter.

Description
Body of adult female (Figure 4) on microscope slide, broadly oval, 1.85-2.40 mm long, 1.45-2.50 mm wide. Anal lobes poorly developed, ventral surface of each lobe with a small sclerotized area, occasionally barely perceptible, and an apical seta 70-125 mm long, sometimes difficult to distinguish from surrounding setae. Antennae each 300-360 mm long, usually with six segments. Legs well developed; hind trochanter+femur 260-300 mm long, hind tibia+tarsus 210-250 mm long; claw stout, 50-55 mm long. Ratio of lengths of hind tibia+tarsus to hind trochanter+femur 0.70-0.92. Ratio of lengths of hind tibia to tarsus 1.08-2.33. Translucent pores present on anterior and posterior surfaces of hind coxa and on posterior surface of hind tibia. Tarsal digitules pointed. Labium 250-280 mm long, 160-180 mm wide, about same length as clypeolabral shield. Circulus 90-95 mm wide, with faint intersegmental line. Ostioles present, well developed, with inner edges of lips sclerotized, each lip with concentrations of setae and trilocular pores. Anal ring 100 mm long, 80-90 mm wide, usually with six setae, each 80-100 mm long, and sometimes accompanied by as many as two shorter setae. Cerarii numbering 17 basic pairs. Anal lobe cerarii each with numerous conical setae of different sizes occupying most of lobe margin, usually two setae longer than others, longest setae each 20 mm long, 5 mm wide at base, accompanied by numerous trilocular pores on a membranous area. Anterior cerarii similar to anal lobe pair, sometimes with long flagellate setae in addition and with anterior and posterior groups of conical setae; trilocular pores present in a band around entire margin so that cerarii appear as a continuous row.
Dorsal surface with long flagellate setae present, mostly about 70 mm long except for setae flanking anal ring where longest 90 mm long. Multilocular disc pores absent. Trilocular pores fairly numerous, evenly distributed. Discoidal pores minute, about same size as a single loculus of a trilocular pore.  Ventral surface with similar setae present to those on dorsum. Multilocular disc pores, each about 7.5 mm in diameter, situated posterior to vulva, in single medial rows at posterior edges of abdominal segments V and VI and in double row at posterior edge of abdominal segment VII. Trilocular pores evenly distributed. Discoidal pores, same as those on dorsum, scattered. Oral collar tubular ducts, each conspicuously wider than a trilocular pore, about 7.5 mm long, 5 mm wide, present across medial areas of abdominal segments III-VI and sparse on abdominal segment VII; present also in marginal groups on abdominal segments II-VII and sparsely distributed around anterior margins of body to area between antennae; one or two ducts sometimes present also between anal lobes.

Comments
This species was described originally from the Solomon Islands on the roots and aerial shoots of coconut. The present record from Fiji, also on coconut, suggests that P. kukumi is extending its range. No damage has been reported so far. The large oral collar tubular ducts on the venter of the adult female, all noticeably wider than the trilocular pores, distinguish this species from all others in the Pacific region. Although the numbers of cerarii in the adult female are sometimes difficult to determine because there are additional anterior and posterior groups of conical setae and trilocular pores to each cerarius, the exact number of 17 pairs can be determined more easily in the immature instars as shown in the third-instar female ( Figure 5).

Description
Body of adult female broadly oval. Antennae six-segmented, rarely seven-segmented. Legs well developed, with translucent pores on anterior and posterior surfaces of each hind coxa and others present on posterior surfaces of each hind femur and tibia. Cerarii numbering 18 distinct pairs, each cerarius with multiple slender conical setae, cerarii on thorax and anterior abdominal segments usually with fewer conical setae; sometimes, in any cerarius, one or two conical setae replaced by slender setae. Anal ring situated 1.0-1.5 times its length from apex of abdomen, bearing six setae. Obanal and cisanal setae stout, situated posterior to anal ring. Dorsal setae short and stiff, often curved, 17.5-25 mm long, except for longer setae 75-85 mm flanking anal ring. Multilocular disc pores present on venter only posterior to vulva and medially across posterior edges of abdominal segments V-VII in single to double rows. Oral collar tubular ducts present on venter, of two main sizes. A small type of duct, distributed medially on abdominal segments IV-VII. A slightly larger type of duct, present in marginal groups on abdominal segments IV-VII and posterior to vulva. A small sclerotized patch present on venter of each anal lobe. Circulus divided by intersegmental line.

Material examined
National Museum of Natural History, and for arranging the loan of specimens for study. I am grateful to Jon H. Martin, Department of Entomology, The Natural History Museum London, for kindly reading the manuscript and commenting on it.