Redescription of Inglisia vitrea Cockerell (Hemiptera, Coccidae) and its transfer to the genus Pseudokermes Cockerell

Th e soft scale insect Inglisia vitrea Cockerell is transferred to the genus Pseudokermes as P. vitreus (Cock-erell) comb. nov. Th e adult female is redescribed and illustrated. A lectotype is designated based on newly slide-mounted dry type material. New host-plant records and an updated geographical distribution for P. vitreus are given. Pseudokermes correntinus Granara de Willink is recognized as syn. nov. of I. vitrea .


Introduction
Th e coccid genus Inglisia Maskell was originally erected for a New Zealand species, I. patella Maskell. As discussed by Morrison and Morrison (1922), Hodgson (1994) and Hodgson and Henderson (2000), very few if any of the species currently included in the genus Inglisia are likely to be congeneric. Indeed, many changes have been made in the taxonomy of this genus in recent years. Prior to 1993, there were nineteen species included in Inglisia (Ben-Dov 1993). Hodgson (1994) erected the genus Vitrococcus to accommodate I. conchiniformis Newstead as V. conchiniformis (Newstead). Ben-Dov et al. (1997) transferred I. foraminifer major Maskell to the genus Cardiococcus Cockerell as C. major (Maskell). Later Hodgson and Henderson (2000) distributed many New Zealand species previously included in Inglisia into two genera, transferring I. fagi Maskell, I. leptospermi Maskell and I. ornata to their newly erected genus Crystal-lotesta as C. fagi (Maskell), C. leptospermi (Maskell) and C. ornata (Maskell), respectively, and transferred I. inconspicua Maskell to their newly erected genus Aphenochiton as A. inconspicuus (Maskell). Ben-Dov (1993) treated Cryptinglisia zizyphi (Brain) and C. elytropappi (Brain) as I. zizyphi [misspelled in Ben-Dov (1993) as I. zizyphy] and I. elytropappi. Hodgson (1994), in his identifi cation manual to genera of Coccidae, followed De Lotto's (1978) recombination in his discussion of the genus Cryptinglisia, and the two species are currently treated as C. zizyphi and C. elytropappi respectively by the scale insect database, ScaleNet (Ben-Dov et al. 2006).
Both Inglisia and Pseudokermes belong to the tribe Cardiococcini (Coccidae: Cardiococcinae) (Hodgson 1994). Inglisia patella, the type species of Inglisia diff ers from P. nitens, the type species of Pseudokermes, mainly by the following combination of features (features of P. nitens in parentheses): (1) Legs developed, with most segments present, although the tibia and tarsus are fused (legs reduced to small areas of sclerotization, occasionally with a claw with small digitules); (2) cribriform-like plates present on dorsum, at level of metathoracic legs (cribriform-like plates absent); (3) marginal setae of two types which alternate around margin: (i) a club-shape spine, blunt apically and narrowing abruptly at base, and (ii) a sharply spinose seta (marginal setae all spinose); (4) stigmatic spines not diff erentiated from marginal spines (with one large stigmatic spine, tapering to a blunt point or with a slightly fl attened apex); and (5) dorsum without bi-or trilocular pores (bi-or trilocular pores frequent along mid-dorsal line and in a narrow band just dorsad to marginal spines) (character states taken from Hodgson 1994). Inglisia vitrea does not share any of the fi ve characteristics listed above for I. patella but, on the other hand, it does share all the features listed for P. nitens. Clearly, Inglisia vitrea is not congeneric with I. patella and would appear to be congeneric with Pseudokermes nitens Cockerell. Based on these morphological affi nities of I. vitrea with P. nitens, we here transfer I. vitrea to Pseudokermes, as P. vitreus (Cockerell) comb. nov. and redescribe the species based on type and non-type material. We include additional features that were not described previously, e.g., distribution of dorsal microducts and simple pores, and the presence of vestigial legs represented by a pair of very short setae. We illustrate the adult female, designate a lectotype, and provide new host-plant and distribution data. In addition, we determine that Pseudokermes correntinus Granara de Willink, described from Argentina (Granara de Willink 1999), is morphologically indistinguishable from P. vitreus and thus it is here synonymized with the latter.

Materials and methods
Specimens were slide-mounted using the method described by Williams and Granara de Willink (1992), except that xylene was used instead of clove oil. Photographs of the dry type specimens and original labels (Figs 1A and B) were taken with a Nikon COOLPIX 3100 digital camera; Fig. 1C was taken with an Olympus OM4 camera, and were processed using the computer program Adobe Photoshop® 5.0. Afterwards, most of the dry type specimens were slide-mounted by T. Kondo. Th e illustration of the adult female ( Fig. 2) shows the dorsum on the left and the venter on the right. Th e redescription of I. vitrea [now P. vitreus] is based on measurements and observations on the type material and other specimens, as listed. Th e terms for external morphology is taken from Cockerell's (1894) original description of the species, with some additional observations. Only specimens of P. vitreus were available in the present study, and material of other species of Inglisia and Pseudokermes were not seen, and so comparisons to other species mentioned in this manuscript were made using published literature.
Measurements of the lectotype are given in parentheses. Th e material studied is deposited at the Auburn University Coccoidea Collection, Auburn, Alabama, U.S.A. (AUCC); the National Museum of Natural History Entomological Collection, Washington, DC., U.S.A., Coccoidea collection held at USDA, Beltsville, Maryland (USNM); and the Bohart Museum of Entomology, University of California, Davis, U.S.A. (BME).
Type material studied. Th e type material consisted of one slide of 2 third-instar nymphs, one uncleared, with label "6409 / Inglisia / vitrea Ckll. / Acacia sp. / Port of Spain, / Trinidad / F.W. Urich #16 / TYPE", plus dry syntype coccids on two small pieces of twig, housed in a small cardboard box with two original labels inside, as follows: "Ckll. Coccidae 25. TYPE / Inglisia vitrea, Ckll / on Acacia sp. Port of Spain, Trinidad / Coll. F.W. Urich. (no. 16)" and "6049 / on / Acacia / Port of Spain / Trinidad / W.I.". Th ese data exactly match those given in Cockerell's (1894) original description. In order to preserve stability and the nomenclatural status of this species, a lectotype is here designated from the syntypes. Th e examined type material is as follows: Adult female, lectotype of Inglisia vitrea Cockerell; paralectotypes: 4 adult females on 4 slides, 3 third-instar nymphs on 2 slides, 1 second-instar nymph on 1 slide, plus several dry specimens still in box (USNM).

A B C
about one-third total length of scale. Surface of scale strongly but fi nely striate radiately on sides; the apparent striations, highly magnifi ed, resolve themselves into rows of small dots (air cells?). Th e dorsal portion of this scale is divided into testudinoid plates; there are apparently six series of such plates, three along each side, but the two middle series are the sides of a single median row, divided only by the keel or ridge. Beyond these the broad margin is not divided into plates. Th e plates are concentrically striate, with a granular patch in the middle of each. Th e median row of plates consists of two large dorsal ones, and two smaller posterior ones. Th e second row consists of three plates on each side, and the third row of seven on each side. Margin with a fringe of rods at rather distant intervals. Th e subtransparent scale removed leaves the insect shiny red-brown, looking like a Lecanium. Derm with large gland pits near the margin; margin with simple spines, varying in size." Th e insects are commonly fl at and almost round when infesting the leaves [ Fig. 1C; see also round-shaped specimen (rshs), upper right of Fig. 2, small inset drawing after Hamon and Williams (1984)], but are more convex and generally longer than broad when infesting twigs (pers. obs.). Slide-mounted material. Body 1.72-4.70 mm long, 1.28-4.84 mm wide. Lectotype damaged, 2.04 mm long, approximately 1.30 mm wide. Note. Th e lectotype and adult female paralectotypes are much smaller than the rest of the adult specimens studied (except for the Florida specimen) because the types (and the Florida specimen) are young adult females.
Dorsum. Derm membranous, becoming entirely sclerotized on older adult females. Dorsal setae completely absent. Dorsal microducts (dmic) oval, bilocular, each about 5 μm wide, with a long terminal fi lament; present marginally, along inner margins of anal lobes and longitudinally in a mid-dorsal line; absent elsewhere. Simple pores (sp) each 2 μm wide, closely associated with dorsal microducts and showing a similar distribution; outline of simple pores and dorsal microducts on mid-dorsal line, especially anterior to anal plates, becoming sclerotized and thus resembling preopercular pores. Dorsal tubular ducts, preopercular pores, dorsal tubercles and pocket-like sclerotizations absent. Anal plates (aplt) together quadrate, with smooth rounded outer angles, plates located at about 1/4 to 1/3 of body length from posterior margin, each plate 133-160 (155) μm long, 70-95 (90) μm wide, anterolateral margin 93-113 (113) μm long, posterolateral margin 90-113 (110) μm long, with 6-10 setae on dorsal surface, plus 1 pair of fringe setae anteriorly and about 3 ventral subapical setae; hypopygial setae not detected. Anal ring (ar) with 6 setae. A narrow sclerotic area present around anal plates, in a narrow mid-dorsal line beginning on area anterior to anal plates and extending anteriorly to level of mouthparts along mid-dorsal line of microducts; on more mature specimens, sclerotizated area on mid-dorsum may be broader but generally confi ned to mid-dorsal areas (as illustrated in Hamon and Williams, 1984), and in older specimens the whole dorsum may become sclerotized.
Diagnosis. Th e adult female of P. vitreus can be diagnosed by the combination of the following features: (1) insect in life covered in a glassy wax cover; (2) mid-dorsal line becoming sclerotized, often with margins also becoming sclerotized; (3) dorsal setae completely absent; (4) dorsal tubercles and dorsal tubular ducts absent; (5) stigmatic setae sharply spinose, one per stigmatic cleft; (6) antennae greatly reduced, one segmented; (7) legs vestigial, represented by a pair of very short setae; (8) ventral tubular ducts in a submarginal band, absent elsewhere; (9) spiracular pores in a narrow band, present in a line from each stigmatic margin towards each spiracular peritreme; and (10) perivulvar pores present in a small group around vulva.
Morphological variation. Th e specimens from Costa Rica and Florida had longer marginal setae (around 55 μm on specimens from those locations versus approximately 30 μm on specimens from elsewhere). Compared to the type material, in the specimens from Barbados, Brazil, and the Dominican Republic, the spiracular pore band becomes broader near the margin. Th is is, no doubt, in part due to the large size of the latter specimens. On the lectotype, a tight band of ca. 30 spiracular pores extends from each spiracle to the body margin. Th e Florida specimen has ca. 15 spiracular pores in a band from spiracle to margin. Th e margins of the insect are often as sclerotic as its midline. According to Dr Chris Hodgson (pers. comm.), specimens found on the leaves are often wider than long; older specimens can be entirely sclerotized dorsally, and often have a wide submarginal band of quite large areolations. Specimens found on twigs, including, the type material, are generally longer than wide (pers. obs.).
Biology. Th e insects can be found on twigs and on leaves, with specimens on twigs being often associated with tending ants (T. Kondo, personal observation). Cockerell (1894) reported a specimen infested by a hymenopterous parasite.
Taxonomic notes. Th e specifi c epithet "vitrea" is amended to "vitreus" in order to match the gender of the genus "Pseudokermes".
In the descriptions of Pseudokermes spp. by Hodgson (1994) and Granara de Willink (1999), all species are described as having preopercular pores. In younger specimens of P. vitreus, the area anterior to the anal plates clearly shows no preopercular pores, but instead there is a mid-dorsal line of bilocular microducts associated with simple pores in the same way that they are present around the body margin. In older specimens, a linear area anterior to the anal plates becomes sclerotized up to level dorsad to mouthparts, and at this point, the structure of the bilocular microducts and simple pores become unrecognizable. Th icker areolations are formed on this sclerotized area (Fig. 2), giving the dorsal microducts and simple pores the appearance of preopercular pores. It is possible that this may be the same situation observed in other species of Pseudokermes. However, this will only be confi rmed by observing teneral adult females, and further studies are needed to test whether this may be the case in species of Pseudokermes in general.
Pseudokermes vitreus appears similar to P. nitens (as described by Hodgson 1994), but the two diff er in the following features (character states of P. nitens in parentheses): (1) spiracular pores present in a line extending from stigmatic areas to spiracles (spiracular pores present in a small group around stigmatic area only and not extending towards spiracle); (2) ventral tubular ducts restricted to a submarginal band (ventral tubular ducts present in a submarginal band with a few present also around mouthparts); and (3) perivulvar pores restricted to a small group around vulva (perivulvar pores present around vulva with some pores also present posteriorly on submedial areas of posterior abdominal segments).
Th e morphological features of P. correntinus Granara de Willink are almost identical to those of P. vitreus and thus P. correntinus is considered here to be junior synonym. All measurements and morphological features of P. correntinus agree with the description of P. vitreus, including the distribution of the spiracular pores, absence of ventral tubular ducts around the mouthparts, number of setae in the anal plates, and the length of marginal and stigmatic setae. Granara de Willink (1999) considered the antennae of P. correntinus to be composed of two segments, however, she also counted two segments in P. eugenium [Hodgson's (1994) P. nitens according to Granara de Willink (1999)] for which species Hodgson (1994) only counted one antennal segment. Furthermore, in P. vitreus there exists a single seta at the base of the one-segmented antennae and, in P. correntinus, Granara de Willink (1999) counted this seta as part of the fi rst segment. Th e apparent diff erence in the number of antennal segments may be due to diff erences in interpretation. Th e legs in P. vitreus are vestigial, and represented by a pair of very short setae (each 3 μm long) and are not easy to detect. Indeed, Hamon and Williams (1984) in their redescription of the species considered the legs to be absent, but in their illustration showed a seta at the position of each leg.

Discussion
Th e type species of Pseudokermes, P. nitens was redescribed by Hodgson (1994) based on non-type material from Argentina and Brazil. Hodgson (1994) considered the material he studied to be conspecifi c with the type material and gave the following differences between his redescription and the original description of Cockerell (1895b): (1) Cockerell (1895b) considered that the antennae and legs in P. nitens were absent but they were present though minute, and (2) the length of the posterior margin of the anal plates was longer than that of the anterior margin in his redescription (vice versa in Cockerell's description). Hodgson (1994) also indicated that the material he studied from San Paulo, Brazil, diff ered from the original description in having more spiracular pores.
Granara de Willink (1999) described three new species from Argentina and considered that Hodgson's (1994) redescription of P. nitens, based on material collected in Misiones, Argentina, corresponded to her newly described species, P. eugenium. Granara de Willink (1999) did not examine the type material of P. nitens and based her decisions entirely on the original description by Cockerell (1985b) and a redescription of the species by Hempel (1900). Granara de Willink (1999) stated that, because Cockerell and Hempel had not mentioned the presence of diff erentiated stigmatic spines in their descriptions of P. nitens which are present on the species from Argentina, she assumed that P. nitens does not have diff erentiated stigmatic spines, and thus decided that none of the Argentinean species where conspecifi c with P. nitens. Granara de Willink (1999) also indicated that P. nitens does not occur in Argentina because all the material she studied from Argentina has reduced antennae, traces of legs and undiff erentiated stigmatic setae (antennae and legs described as absent, and stigmatic spines not mentioned by either Cockerell (1895b) or Hempel (1900)). Microscopes in the early 1900's were not as powerful as those available at present, and on many occasions reduced morphological features were not observed by earlier coccidologists, including T. D. A. Cockerell. Th e legs in all species studied by Granara de Willink (1999) are greatly reduced and are represented by either a small sclerotization or a tiny claw with digitules. With the microscopes available in the late 1800's and early 1900's, it would have been very diffi cult to see such reduced features, which may have gone undetected. For example, Cockerell (1902) describes Akermes colimae Cockerell [now Aztecalecanium colimae (Cockerell)] as having no antennae and legs; however, a recent redescription by Kondo and Williams (2004) of this species clearly shows a pair of reduced antennae and small legs with all their segments (coxa, trochanter, femur, tibia, tarsus and claw). Th us, it is critical to compare the type material of P. nitens and P. eugenium in order to determine if they are conspecifi c.
As a result of this study, the composition of the genus Pseudokermes is changed to include 8 species: P. armatus, P cooleyi, P. eugenium, P. geoff roeum, P. marginatus, P. nitens, P. palmae and P. vitreus. Pseudokermes may include more synonyms because of the lack of studies based on type material. Furthermore, it is doubtful whether all species currently included in Pseudokermes are congeneric with the type species of the genus, P. nitens, and further studies are needed in order to elucidate their relationships.