A new genus and two new species of soft scale insect ( Sternorrhyncha , Coccoidea , Coccidae ) from Africa

A new soft scale insect genus, Sterculicoccus Hodgson, gen. n., is introduced to take a new species, Sterculicoccus tafoensis Hodgson, sp. n., off Triplochiton from Ghana. Sterculicoccus belongs to the Myzolecaniinae and is close to Alecanium Morrison. In addition, a new species of Hemilecanium Newstead, H. cedrelus Hodgson, sp. n., is also described off Cedrela toona from Zambia. Hemilecanium cedrelus is close to H. coriaceum Hall and H. uesatoi Kondo & Hardy (Saissetiinae). Th e adult females of these two species are described, along with the 1st-instar and the 2ndand 3rd-instar females of H. cedrelus.


Introduction
Whilst the Coccidae of some areas of Africa are quite well known [e.g.South Africa and Zimbabwe -both with more than 80 species (Scalenet, 2008)], others have had very little attention.Two such countries are Ghana (with 24 known species of Coccidae) and Zambia (with 16 species).Th e present paper describes a new species from each of the latter two countries, one of them in a new genus.

Materials, methods, and conventions
Several specimens were already mounted but others were mounted especially for this study in the usual way (Hodgson and Henderson, 2000), using acid fuchsin stain, except that the specimens were left in unheated KOH for about 72 h for clearing.Th e fi gures are typical scale insect diagrams, with the dorsal surface forming the left of the main drawing and the ventral surface forming the right; important structures within these central drawings are enlarged around the margins; these are not drawn to the same scale.Terminology follows that of Hodgson (1994).Specimens of both species will be deposited in Th e Natural History Museum, Cromwell Road, London (BMNH) and in the Coccoidea Collection of the Smithsonian Museum, Washington (USNM), kept in the US Department of Agriculture, Beltsville, Maryland.
Sterculicoccus Hodgson, gen.n. urn:lsid:zoobank.org:act:DBD73214-1639-45E6-8F3C-6618B8CDD3E0Type species: Sterculicoccus tafoensis Hodgson, sp.n. urn:lsid:zoobank.org:act:E30D4191-C87F-472A-A2FF-B745A2954F05Generic diagnosis.Body broadly oval.Anal cleft deep, about 1/3 rd -1/4th total-body length, with parallel margins; with a narrow sclerotisation around anterior margin of cleft and sclerotisations ventrally in each stigmatic cleft.Dorsal derm otherwise membranous with numerous areolations, those anterior to anal plates and dorsal to mouthparts much larger than elsewhere.Dorsal setae mainly short, but with a few long.Anal plates together oval, with polygonal reticulations on dorsal surface.Outer anal opening with an O-shaped sclerotised band.Marginal setae apparently ventral, short and spinose, in a band 2-3 setae wide; stigmatic clefts absent but with stigmatic sclerotisations in each stigmatic area.Venter with a marginal band of large tubular ducts plus a broad submarginal band of much smaller tubular ducts.Pregenital discpores each with mainly 5 loculi, in a small group beneath posterior end of anal plates.Antennae small, 6 segmented.Clypeolabral shield proportionately large.Spiracles very large.Legs much reduced.
Relationships.Sterculicoccus, gen.n., is clearly a member of the subfamily Myzolecaniinae as defi ned by Hodgson (1994) -absence of dorsal tubular ducts; absence of eyespots; rather large spiracles; pregenital setae represented by bands of setae rather than pairs of larger setae; legs and antennae reduced in size, and anal tube short.However, it is unusual within this group in having (i) 2 types of ventral tubular ducts (usually only of one type) and (ii) each anal plate with only 4 setae, all near the apex (typically with many setae).In the Key to Myzolecaniinae in Hodgson (1994, p. 91), Sterculicoccus keys out at Alecanium Morrison.Alecanium and Sterculicoccus are clearly closely related and share the following character states: (i) stigmatic clefts very shallow or absent, without stigmatic spines; (ii) each stigmatic area with a small sclerotisation enclosing some spiracular disc-pores; (iii) marginal setae spinose, in a marginal band several setae wide; (iv) some dorsal setae long and fl agellate; (v) anal plates elongate oval; (vi) pregenital disc-pores with mainly 5 loculi; (vi) legs very reduced; (vii) antennae reduced; (viii) spiracles large; (ix) ventral tubular ducts in a wide submarginal band and in transverse bands medially on meso-and metathorax; (x) ventral setae with a similar distribution (i.e. with a band across metathorax), and (xi) outer end of anal tube with an O-shaped sclerotisation.Sterculicoccus diff ers from Alecanium in having (character-states on Alecanium in brackets): (i) two types of ventral tubular ducts (only 1 type); (ii) only one type of marginal seta (two types); (iii) dorsal surface of anal plates with a polygonal reticulate pattern of microridges (absent); (iv) anal plates each with only four setae near apex (each with many setae distributed over most of dorsal surface), and (v) long dorsal setae restricted to dorsad to mouthparts (throughout dorsal surface).
Th e key in Hodgson (1994)    Derm membranous, apart from a narrow crescentic sclerotisation around anterior margin of anal cleft.Derm with abundant small areolations, each polygonal in shape, with a central dorsal microductule.Dorsal setae variable: (i) a larger slightly spinose seta with a sharp apex and well-developed basal socket; each 16-18 μm long; apparently restricted to a broad submarginal band, plus a few occasionally up margins of anal cleft; (ii) a slightly smaller, more setose seta, each 12-16 μm long, in a broad area around anal plates, and (iii) a short, fi ne seta, each about 8-15 μm long, frequent throughout rest of dorsum, but also with 0-3 setae up to 100 μm long on either side of large areolation dorsad to mouthparts.Dorsal pores of 2 types: (i) a small convex, closed pore, each about 3 μm wide, frequent throughout; and (ii) a small microductule with a narrow outer ductule about 5 μm long, with sclerotised pore about 1.0-1.5 μm wide and an inner ductule quite long and swollen at proximal end; abundant throughout with one in each areolation.Dorsal tubular ducts, dorsal tubercles and preopercular pores absent.Anal plates together approximately oval, each plate with a rounded outer angle and with dorsal surface covered in micro-ridges arranged in a polygonal pattern; without a small additional triangular anal plate between lateral plates anteriorly; each plate 185-210 μm long, combined width 140-165 μm; each plate with 4 setae; 2 along inner margin, 1 near apex (each 33-38 μm long) and one dorso-laterally 45-50 μm long.Ano-genital fold possibly absent; posterior opening of anal tube apparently with a sclerotised O-shaped margin, with a group of 10-18 short setae, each 25-35 μm long, medially between ventral arms of anal ring posteriorly.Anal plates without strong inner apodemes.Anal ring with 3 pairs of anal ring setae, each about 125 μm long, all setae attached to ventral half of anal ring; anal ring apparently without anal ring pores; anal tube quite short, anal ring lying beneath anal plates.Eyespots not detected.

Sterculicoccus tafoensis
Margin.Marginal setae all short and spinose with parallel margins, each 11-17 μm long, in a marginal band 2-3 setae wide; basal socket broad; with several hundred around margin, appearing to be on ventral surface; setae on margins of anal lobe not diff erentiated.Stigmatic clefts absent or extremely shallow, each with a small area of sclerotisation on venter which encloses outermost spiracular disc-pores; without stigmatic spines.
Venter.Derm entirely membranous.Pregenital disc-pores quite large, each 8-10 μm wide, with mainly 5 or 6 loculi; restricted to a small group of 18-30 (total) apparently posterior to anal opening, probably with 8-20 on each side.Spiracular disc-pores each mainly with 5 loculi, present in a wide band between spiracles and margin, with perhaps 45-55 in anterior band and 50-70 in posterior band.Ventral microducts, each with ductule about 5 μm long and pore perhaps 2 μm widest; abundant throughout but perhaps least frequent medially on abdomen.Other ventral pores, including preantennal pores, absent.Ventral tubular ducts of 2 types: (i) ducts with a long, narrow outer ductule (each about 25 μm long and 2 μm wide), with a narrow, much shorter inner ductule about 14 μm long, and with a fairly small glandular end; in a fairly broad submarginal band, which extends medially past spiracles; also with a few on either side of mouthparts; absent medially on thorax and abdomen; (ii) much larger ducts, each with a long, narrow outer ductule about 33-40 μm long and 3 μm wide, with a sclerotised outer orifi ce, and a narrow, much shorter inner ductule about 20-28 μm long, and a fairly small glandular end; present in a dense marginal band about 2-3 ducts wide, intermingled with marginal setae.Ventral setae abundant, particularly submarginally, where quite fi ne, each 7-9 μm long, with a wide basal socket; some medially rather longer, longest about 27 μm long; pregenital setae not diff erentiated; with 2 pairs of interantennal setae, these similar to ventral submarginal setae, each 16-28 μm long.
Comment. S. tafoensis new species is the only species currently known in this genus but is readily separated from other possibly similar species by the presence of: (i) a reticulate pattern of microridges on the dorsal surface of the anal plates, (ii) the presence of two types of ventral tubular ducts, the larger ducts forming a marginal band and the smaller ducts in a broad submarginal band, and (iii) marginal setae of one type only, all spinose, in a band 2 to 3 setae wide around the entire margin, and not diff erentiated into stigmatic spines at each stigmatic area.
Etymology.Th e specifi c name tafoensis is composed of Tafo, the name of the site in Ghana where this species was collected, and -ensis (Latin), a suffi x denoting place or locality.Hemilecanium Newstead urn:lsid:zoobank.org:author:2DEA755F-A46A-4203-B3B8-893324D22FAFurn:lsid:zoobank.org:pub:7FB67D8F-AAA7-427B-AF54-53D64A52B87ATh e genus Hemilecanium was introduced by Newstead (1908) to take H. theobromae, collected off cacao in Cameroon.Prior to 2005, Hemilecanium contained 4 species: H. coriaceum Hall, H. imbricans (Green), H. recurvatum Newstead and the type species.Since then however, there have been major changes in the species composition of Hemilecanium, most notably when Kondo and Hardy recently (2008) synonymised Etiennea Matile-Ferrero (type species E. villiersi Matile-Ferrero) with Hemilecanium, based on a comparative morphological study of the adult females, adult males and 1 st -instar nymphs of 4 species of Hemilecanium and 6 species of Etiennea.Th e study also included a phylogenetic analysis based on adult female and 1 st -instar nymphal characters.Th ese changes brought the total number of species in Hemilecanium to 26.Earlier, Hodgson (1994) had drawn attention to the morphological similarities of Etiennea and Hemilecanium (both genera having dorsal tubular ducts with funnel-like apertures and both with two centres of sclerotisation on the dorsum) but had considered that they could be easily separated due to the presence of cribriform plates on Hemilecanium (absent on Etiennea species).However, Kondo and Williams (2005) showed that 4 cribriform plates were present on the dorsum of the 1 st -instar nymphs of several Hemilecanium species so that the 1 st -instar nymphs of E. villiersi and H. theobromae were very similar.
Despite these changes, the species within the new concept of Hemilecanium can be divided into two groups based on the structure of the 1 st -instar nymphs.Th ese are referred to as the petasus and thoebromae groups by Kondo and Hardy (2008).Th e 1 st -instar nymphs are only known for nine of the species now included in Hemilecanium but these can be separated as follows: the petasus group is diagnosed by the presence of: (i) the dorsum without clusters of 4-locular pores; (ii) each spiracular pore band with 2-4 spiracular disc-pores; (iii) each femur with a very long seta near apex (only on the prothoracic leg of H. uesatoi), and (iv) each stigmatic cleft with 3 well-developed stigmatic spines.Th e petasus group includes H. ferox (Newstead), H. montricardiae (Newstead), H. multituberculatum (Hodgson), H. petasus (Hodgson), H. sinetuberculum (Hodgson) and H. uesatoi Kondo & Hardy.Th e theobromae group, on the other hand, has the following combination of characters: (i) dorsum with 4 clusters of 4-locular pores; (ii) each spiracular pore band with 1 spiracular disc-pore; (iii) very long setae absent from all femora, and (iv) stigmatic spines absent.It includes H. imbricans, H. mangiferae Kondo & Williams, H. theobromae and H. villiersi.As pointed out by Kondo and Hardy, these two groups are also supported by a single adult female character, with those in the theobromae group lacking stigmatic spines while those in the petasus group have well-developed stigmatic spines.Th is suggests that the present composition of Hemilecanium may be paraphyletic.Th is was understood by Kondo and Hardy who go on to say (2008, p. 212) "We need data on fi rst-instar nymphs of additional species to determine whether these two crawler types and their associated adult females will correspond to reciprocally monophyletic groups".

Hemilecanium cedrelus
Adult female (Fig. 2).Described from 5 specimens in good condition, and with reference to the remaining 10 specimens.
Instar diagnosis.Dried material with many clear, round, brown-ochre-coloured spots present throughout dorsum indicating position of dorsal tubercles.Mounted material oval to almost round.Dorsum membranous apart from a narrow sclerotisation around anterior margin of anal cleft; derm with a reticulate pattern of areolations; also with about 150-200 randomly distributed dorsal tubercles, distributed over entire dorsum, each without satellite tubular ducts; also "scars" (see discussion below) in position of dorsal tubercles of 3rd-instar.Conical preopercular pores and dorsal setae present.Anal plates roughly quadrate with 4 setae near apex.Margin with a single line of sharply spinose setae; with 3 stigmatic spines each clearly diff erentiated from marginal spinose setae, median stigmatic spines much longer than lateral spines.Venter with abundant 10-locular disc-pores on all abdominal and thoracic segments.Ventral microducts abundant throughout.Tubular ducts of probably 1 type, present in a wide submarginal band.Antennae 6 segmented.Limbs fully developed but relatively small; tibio-tarsal articulation without a sclerosis; claw digitules diff erent; claw with a small denticle.Spiracles proportionately very large, width of peritreme much wider than width of coxae.Mouthparts relatively large.
Unmounted material.Dried material quite dark brown; younger specimens with a distinct shallow longitudinal dorsal ridge and a clear, shelf-like margin; older specimens becoming strongly convex with two strong shoulders medially; venter becoming highly concave, forming an inner egg chamber.Positions of dorsal tubercles indicated by clear round areas of more brown-ochre-coloured derm scattered over dorsum.
Dorsum.Derm mainly membranous but with a heavy sclerotised band around anterior margin of anal cleft, which expands anteriorly with age.Derm of available specimens with polygonal reticulations throughout, each reticulation with an inner areolation and a dorsal microductule; perhaps becoming sclerotised at maturity.Dorsal setae all rather spinose, quite sharply pointed, with parallel sides; frequent laterally and anteriorly to anal plates, but absent submarginally; those anterior to anal plates each 33-36 μm long, those more anteriorly and laterally smaller, down to 16-24 μm long.Preopercular pores present in an elongate triangular group anterior to anal plates, each pore conical and 8-11 μm wide; each group with 85-140 pores, extending anteriorly to about metathorax.Dorsal microductules oval, appearing bilocular but with single inner ductule arising medially, each about 2.5-3.0 μm widest, most ductules swollen proximally; abundant, present in each dorsal reticulation.Other dorsal pores absent.Dorsal tubercles large and convex, but sunken into derm; each 16-26 μm wide, with a heavily sclerotised outer cone plus 2 inner chimney-like tubes (one inside other); outermost tube with a ring of about 10-12 vertical ridges (see Discussion below adult female); innermost tube long, extending some way above tubercle; dorsum with a total of about 140-200 tubercles, randomly distributed throughout; also with about 16 submarginal "scars" and 2 pairs of submedial "scars" (in approximate positions of dorsal tubercles of 3 rd instar).Anal plates each about 275-330 μm long, width of single plate 115-170 μm; each plate triangular, with 4 apical setae, both inner margin setae and subapical seta 25-30 μm long, other seta appearing dorsal, about 60-72 μm long.Anogenital fold with a line of 6-8 setae along anterior margin, each fairly short but with a long seta at each corner, latter about 60-75 μm long; each lateral margin with 3 setose setae.Anal ring well developed, with many pores and probably 5 pairs of setae, each 270-360 μm long; anal tube about as long as anal plates.Eyespots not detected.
Venter.Derm membranous.Spiracular disc-pores each mainly with 5 loculi, in broad groups near margin and each peritreme but very few or even sometimes absent in between; with about 45-50 in each anterior band and 60-85 in each posterior band, latter with a small group of multilocular pores near each spiracle.Multilocular disc-pores each about 8-10 μm wide, mainly with 10 loculi, abundant across all abdominal segments and across meso-and metathorax; scarce on prothorax and head.Ventral microducts each about 3 μm wide, abundant throughout, except marginally.Ventral tubular ducts slightly variable but probably all of one type, each with an outer ductule 17-30 μm long, inner ductule 13-20 μm long, with or without a glandular end; abundant in a broad submarginal band and rather less frequently in bands across each thoracic segment; with 1 or 2 present medially on abdomen among multilocular disc-pores.Other pores types absent.Ventral setae mainly rather spinose, most about 20-26 μm long, present across each abdominal and thoracic segment but most abundant in a submedial band just laterad to spiracles and legs; with about 5 pairs of rather short inter-antennal setae, longest 40 μm long; abdominal segments V, VI and VII each with a pair of longer setae, longest on VI and VII, each about 150 μm long; submarginal setae frequent, each about 25 μm long.
Comment.Th e adult females of this species are superfi cially similar to those of Hemilecanium coriaceum Hall and H. uesatoi Kondo & Hardy, which also have dor-sal tubercles randomly distributed throughout the dorsum,.However, H. cedrelus differs from H. coriaceum as follows (characters-states on H. coriaceum in brackets): (i) presence of 3 clearly diff erentiated stigmatic spines (absent or 1 barely diff erentiated); (ii) multilocular disc-pores abundant across all abdominal segments and also across meso-and metathorax (many fewer, restricted to abdomen); (iii) preopercular pores in an elongate group anterior to anal plates (in a broad group incorporating some dorsal tubercles anterior to anal plates); and (iv) large size of spiracular peritremes (small).Adult female H. cedrelus are also similar to the newly described H. uesatoi from the Ryukyu Archipelago, Japan, but the latter diff ers in having normal-sized spiracles; pocket-like sclerotisations restricted to the submargin, preopercular pores extending anteriorly onto head; 8-segmented antennae, and no denticle on the claw.In addition, the 1 st -instar nymphs are rather diff erent (see under that stage below).
Initially it was assumed that the ring of sclerotised spots on each dorsal tubercle, which are clearly visible in dorsal views of each tubercle, were satellite tubular ducts similar to those on some Hemilecanium species.However, true satellite ducts have the structure of a small tubular duct, i.e. with a long outer ductule, a small cup-shaped invagination and sometimes an inner ductule, each duct opening onto the surface of the tubercle through an aperture some distance from the funnel-like central cone (see Etiennea halli, E. kellyi, E. petasus and E. villiersi in Hodgson, 1991 (now all in Hemilecanium)).Th is is quite diff erent from what we see on H. cedrelus where, when seen from the side, these "sclerotised spots" appear to refer to vertical sclerotised ridges on the outer funnel-shaped tube; none of which have either an associated ductule or an outer aperture.It is therefore considered that these structures are not satellite tubular ducts.
Distribution.Hemilecanium cedrelus is currently only known from Zambia.
Th ird-instar female (Fig. 3) Described from 1 specimen in good condition.(Note: the data in brackets for the number of dorsal tubercles are from the pharate 3rd-instar nymph -see discussion under 2nd-instar nymph below.) Instar diagnosis.Oval and rather fl at.Dorsum with a submarginal ring of large dorsal tubercles plus 2/3 submedially on thorax.Margin with a single line of sharply spinose setae; with 3 stigmatic spines clearly diff erentiated from marginal spinose setae.Venter with a small group of 5-locular disc-pores posterior to anal opening; also with a sparse submarginal band of tubular ducts.Antennae 5 or 6 segmented.Legs fully developed; claw digitules dissimilar; claw with a small denticle.Spiracles of normal proportions, width of peritreme smaller than width of coxae.Mouthparts relatively large.
Unmounted material.Dried material pale brown; oval, rather fl at, with a few shallow, radial ridges.No sign of a wax test.
Mounted material.As in instar diagnosis.Body 1.33 mm long and 0.9 mm wide; anal cleft about 1/5th body length.Dorsum with a submarginal ring of about 30 large dorsal tubercles plus 2(3) submedially on thorax, plus "scars" left by the 12 dorsal tubercles of 2 nd -instar nymph.Dorsum.Derm mainly membranous, without a sclerotised band around anterior margin of anal cleft and without a reticulate pattern of areolations.Dorsal setae very few, each short and fi nely spinose, each about 10 μm long; distribution uncertain but very sparse.Preopercular pores absent.Dorsal microductules small, each about 1.5 μm wide, with an inner ductule about 8 μm long, most ductules swollen proximally; abundant.Other dorsal pores absent.Dorsal tubercles of more or less two types: (i) large and convex, each about 20-24 μm wide, basically similar to those on adult female; with (on each side in a submarginal ring) 5 (5-7) on abdomen; 2 (3) between stigmatic clefts, 2 (3) between anterior cleft and eyespot and 4 anteriorly between eyespots; and (ii) what are here considered to be remains of dorsal tubercles of 2nd-instar female, structure very unclear (but probably similar to "scars" on adult female); with 3 on each side of abdomen, 1 between stigmatic clefts, between anterior cleft and eyespot and 2 anteriorly between eyespots.Anal plates each about 110 μm long, width of single plate about 55 μm; each plate triangular, with 4 setae: 2 inner margin setae, both short, and two other apical setae, positions uncertain, each possibly 16-23 μm long.Anogenital fold with a line of perhaps 8 setae along anterior margin, each about 28 μm long; each lateral margin with 2 setae.Anal ring well developed, with 4 pairs of setae, each about 100 μm long; anal tube about twice length of anal plates.Eyespots oval, 21-23 μm widest.
Margin.Marginal setae all sharply spinose, each 12-16 μm long, with a broad base, rather straight sides and narrow basal sockets; with a total of 26 anteriorly between eyespots, and (on each side) 15-19 between eyespots and anterior stigmatic areas, 20-22 laterally between stigmatic areas and 58-64 on each side of abdomen; each anal lobe with a group of 2 or 3 longer, slightly curved setae, longest 53-66 μm long.Stigmatic clefts shallow, each with 3 stigmatic spines, clearly diff erentiated from marginal spines, slightly curved and with a less pointed apex than marginal setae; median spine longest, 70-90 μm long, each lateral spine 20-40 μm long.
Venter.Derm membranous.Spiracular disc-pores, each with a broad sclerotised margin and perhaps mainly with 5 or 6 loculi, in broad bands between margin and each peritreme; with 20 or 21 in each anterior band and 24-26 in each posterior band.Preanal multilocular disc-pores each about 5 μm wide with 5 loculi, in a small group of 3 each side beneath anal plates.Ventral microducts each about 3 μm wide, abundant throughout apart from marginally, where absent.Ventral tubular ducts of one type, each with an outer ductule 15 μm long, inner ductule 10-12 μm long, with a glandular end; in a mainly narrow submarginal band but tending to be most concentrated on either side of spiracular disc-pore bands, with 3 anteriorly on head, 1-3 between eyespots and anterior stigmatic clefts, 11-12 on each side between stigmatic clefts and 11-18 on each side of abdomen.Other pore types absent.Ventral setae perhaps slightly larger medially than laterally, most about 10-12 μm long; with one longer pair and one shorter pair of inter-antennal setae, longest about 60 μm; with long setae medially on abdominal segments V-VII, longest about 85 μm; with 2 or 3 short setae associated with each coxa; other setae fairly frequent in a broad submarginal band, most abundant on abdomen, each about 6-7 μm long.
Comment.Th is is the only immature stage of any Coccidae known to the author with preanal multilocular disc-pores.
Second-instar female (Fig. 4) Described from a single specimen containing a pharate 3 rd -instar nymph.Instar diagnosis.Oval.Similar to 3 rd -instar but venter without disc-pores posterior to anal opening and without tubular ducts; marginal setae and spiracular discpores fewer.
Unmounted material.Dried material pale brown; oval, rather fl at, with a few shallow, radial ridges.No sign of a wax test.
Mounted material.As for instar diagnosis.Body 1.5 mm long and 1.08 mm wide; anal cleft about 1/6th body length.Submarginal ring of 12 dorsal tubercles.
Dorsum.Derm mainly membranous, without a sclerotised band around anterior margin of anal cleft and without a reticulate pattern of areolations.Dorsal setae possibly absent.Dorsal microductules small, each about 1.5 μm wide, with or without an inner ductule, most ductules swollen proximally; sparse.Other dorsal pores absent.Dorsal tubercles of 1 type, each 11-15 μm wide; structure basically similar to those on adult female but smaller, each with about 6 vertical ridges on outer inner tube; with 1 pair of tubercles anteriorly, and (on each side) 1 between eyespots and anterior stigmatic cleft, 1 laterally between stigmatic clefts and 3 or 4 on abdomen (plus dorsal tubercles of pharate 3rdinstar nymph (as indicated in brackets in the description above).Anal plates each about 90 μm long, width of single plate about 42 μm; each plate triangular, with 4 setae, all broken.Anogenital fold with 2 pairs of setae anteriorly, longest setae about 33 μm long; each lateral margin with 1 seta.Anal ring well developed, with 3 pairs of setae, each about 110 μm long; anal tube about twice length of anal plates.Eyespots oval, 15 μm widest.
Margin.Marginal setae all sharply spinose, each 8-20 μm long, with a broad base, slightly curved sides and narrow basal sockets; with 12 anteriorly between eyespots, and (on each side) 6 between eyespots and anterior stigmatic areas, 6 or 7 laterally between stigmatic areas and 19-22 on each side of abdomen; each anal lobe probably with a group of 2 or 3 longer setae but all broken.Stigmatic clefts shallow, each with 3 stigmatic spines, clearly diff erentiated from marginal spines, slightly curved and with a less pointed apex than marginal setae; median spine longest, 60 μm long (only one present), each lateral spine 15-18 μm long.
Venter.Derm membranous.Spiracular disc-pores each with a broad sclerotised margin and perhaps mainly with 5 loculi, in narrow bands between margin and each peritreme; with 7 in each anterior band and 10-12 in each posterior band.Preanal multilocular disc-pores absent.Ventral microducts each about 3 μm wide, sparse in a broad submarginal band and also occasional medially on head, thorax and abdomen.Ventral tubular ducts absent.Other pores types absent.Ventral setae few, with two pairs of interantennal setae, longest 33+ μm; with long setae medially on abdominal segments V-VII, longest about 80+ μm; each coxa with 1 minute associated seta; other setae in a submarginal line (1 laterally between stigmatic clefts), and an inner submarginal line on abdomen; each about 6-7 μm long.
Comment.Despite having a slightly larger body size than the 3 rd -instar nymph described above, this is clearly a female 2 nd -instar nymph.Th is is shown not only by the smaller limbs etc, smaller number of spiracular disc-pores and absence of preanal disc-pores, but also by the presence of the pharate 3 rd -instar and the distribution of the latter's dorsal tubercles, which are clearly visible inside its cuticle.
Because this specimen had a pharate 3 rd -instar nymph within, it was possible to study the number and distribution of the dorsal tubercles, which were well developed and these data are given in brackets in the description of the 3 rd -instar nymph above.Th e number and distribution of the "scars" on the derm of the 3 rd -instar nymph agree with the number and positions of the dorsal tubercles on the 2 nd -instar nymph and therefore clearly refer to these (see also the Discussion beneath description of 1 st -instar nymph of H. cedrelus).
First-instar (Fig. 5) Instar diagnosis.Oval.Dorsum membranous but with a series of 1-4 large, triangular or cone-shaped protuberances medially on most segments.Dorsal setae absent.Margin with small spinose setae.Each stigmatic cleft with 3 stigmatic spines, median spine long.Venter with three pairs of long preanal setae.Ventral microducts in a sparse submarginal line.Legs well developed; each femur with an exceptionally long seta on anterior margin; long setae also present on tibia and tarsus; claw digitules diff erent; claw with a small denticle.
Mounted material.As for instar diagnosis.Body 0.5-0.53mm long and 0.26-0.30mm wide; anal cleft very short.
Dorsum.Derm mainly membranous, but with large triangular or cone-shaped protuberances medially, each margin of protuberance about 33-35 μm long and each 20-22 μm wide at base, distributed as follows: none on abdominal segments V-VII, pairs on abdominal segments III & IV, singles on abdominal segments I & II, each thoracic segment plus a pair posteriorly on head and 2 pairs together more anteriorly on head.Dorsal setae absent.Dorsal microductules small, each about 1.5 μm wide with a long inner ductule, most ductules swollen proximally; mainly in 2 pairs of longitudinal lines, one pair of lines medially (with 5 pores on abdomen, probably 3 on thorax and 1 on head) and other lines submarginal (with 7 pores on abdomen, 2 between stigmatic clefts and 4 anteriorly); also with single pores submedially in each thoracic segment.A pair of trilocular pores present on head some distance from anterior margin, each about 3.0 μm wide.Other dorsal pores absent.Anal plates each quite elongate, about 60-65 μm long, with a few shallow longitudinal ridges; each with a small spine on inner margin, 2 short setae along inner margin, each about 8 μm long; a very long apical seta, each 260-310 μm long, and a single seta on posterior margin, about 36 μm long.Anogenital fold with a single long seta at each corner, each 26-28 μm long, and a single similar seta on each lateral margin, 23-28 μm long.Anal ring with 2 rows of pores, each with 5-7 pores, plus 6 anal ring setae, each about 80-85 μm long; anal tube extending anterior to anal plates.Eyespots each 15-18 μm wide.
Margin.Marginal setae all fi nely spinose, most 6-8 μm long (that on each anal lobe 13-15 μm long), with well-developed socket, distributed as follows: with 8 anteriorly between eyespots, and (on each side) 2 between eyespots and anterior stigmatic cleft, 2 laterally between stigmatic clefts and 8 on abdomen.Stigmatic clefts shallow, each with 3 stigmatic spines clearly diff erentiated from marginal setae; median spinose seta very long, rather parallel sided and possibly with a slightly fl attened apex, each 40-45 μm long, with a broad basal socket; anterior lateral spine shortest, about 5 μm long, posterior spine about 6.5 μm long.
Venter.Derm membranous.Spiracular disc-pores each with very thick margins and perhaps with mainly 3 or 4 loculi (occasionally 5?), with 3 pores in each anterior pore band and 4 in each posterior band.Ventral microducts each about 1.5 μm wide, present in a submarginal line, with (on each side) 1 on head, 2 on thorax and probably 6 on abdomen.Ventral setae few; with 1 pair of interantennal setae, each about 40 μm long, and with pairs of long setae medially in abdominal segments V-VII, longest about 40-45 μm long; short setae in a submarginal line, with (on each side) 7 on abdomen, 1 on thorax and 1 anteriorly on head; also with an inner submarginal line of 7 setae on abdomen.
Comment.Th e 1 st -instar nymph of H. cedrelus is distinctive due to the presence of the triangular or cone-shaped protuberances medially on the dorsum of most seg-ments, unknown on any other 1 st -instar nymphs as far as the author is aware, including those of H. uesatoi.However, Hodgson (1993), when describing the dorsum of Etiennea (Hemilecanium) petasus wrote "Derm entirely membranous, but thrown into small dermal nodules -in some specimens, these are rather pronounced and found throughout, in others they are few, but are always present around the margin and in pairs medially, probably one pair per segment" (my italics).In seems possible, therefore, that these "nodules" are just more pronounced on H. cedrelus.In addition, the presence of very long setae on each femoral segment of H. cedrelus is unusual, although similar setae are known on other species (Hodgson, 1993) in the petasus group as defi ned by Kondo and Hardy (2008).Th ey are also known on Protopulvinaria pyriformis (Cockerell) (Ray & Williams, 1982) and Kilifi a De Lotto (Ray & Williams, 1982).In addition to the triangular or cone-shaped protuberances medially, which are absent on the 1 st -instar nymph of H. uesatoi, the 1 st -instar nymphs of H. cedrelus diff er from those of H. uesatoi in having (character-states on H. uesatoi (from Kondo & Hardy, 2008) in brackets): (i) long femoral setae on all femora (restricted to the metafemur only); and (ii) claw digitules dissimilar (similar).
Etymology.Th e specifi c name cedrelus is taken from the generic name of the host plant, Cedrela toona (Meliaceae).
A key to the adult females of Hemilecanium Newstead was included in Kondo and Hardy (2008, p. 195).Th is key can be modifi ed to include H. cedrelus as follows: General discussion Boratynski (1970), when describing the immature stages of Parthenolecanium persicae (Fabricius), showed that the positions of the dorsal tubercles (which he called marginal bicylindrical ducts) on immature instars was indicated in the following instar (i.e. the positions of the tubercles of the 2 nd -instar nymphs on the derm of the 3 rd -instar nymph) by atrophied ducts.Th is is a fairly common phenomenon in the Coccidae, where the term "pocket-like sclerotisations" has been frequently used to describe the atrophied tubercles.In H. cedrelus, the atrophied tubercles do not look like the pocket-like sclerotizations found on other Coccidae, although their exact structure is hard to determine, and so are referred to here simply as "scars".Th us, the position of the dorsal tubercles on the 2 nd -instar nymph is shown by "scars" on the dorsum of the 3 rd -instar nymph, and those of the 3 rd -instar nymph by "scars" on the dorsum of the adult female.It appears that the "scars" on the 3 rd -instar nymph (left from the tubercles of the 2 nd -instar nymph) disappear at the last moult into the adult female as none could be detected.