Coccus pseudotumuliferus Gullan & Kondo, 2018

Gullan, Penny J., Kondo, Takumasa, Fiala, Brigitte & Quek, Swee-Peck, 2018, Taxonomy of coccids (Hemiptera: Coccidae: Coccus L.) associated with Crematogaster ants (Hymenoptera: Formicidae) in the stems of Macaranga plants (Euphorbiaceae) in Southeast Asia, Zootaxa 4521 (1), pp. 1-51 : 34-39

Coccus pseudotumuliferus Gullan & Kondo sp. n.

urn:lsid:zoobank.org:act:73044BE2-763D-494F-AA8B-0CCEF0859A1D

( Figs 2E , 11 A–C, 12)

Coccus “ tumuliferus var. C. 84”, Heckroth et al. 1998: 431, 432, 434, 436, 438 & 440.

Morphospecies C. 214, Heckroth et al. 1998: 431, 433, 436, 437 & 440.

Coccus "near tumuliferus ", Quek et al. 2017: 823.

Type material examined. Holotype: adult female, BORNEO: Sabah, Crocker Range, Tikolod, 650 m, ex hollow stem of Macaranga indistincta , 18 Oct. 1999, coll. S.-P. Quek, SPQ.125b, DNA voucher 1(1) ( FRIM). Paratypes: BORNEO: same data as holotype except ex M. pearsonii & M. glandibracteolata , SPQ.126a & SPQ.129b, DNA vouchers 2(2) ( ANIC); Sabah, Crocker Range, Keningau to Ulu Kimanis trail, ~ 5.28° N, ~ 116.05° E, 900–1200 m, ex M. angulata , M. puberula & M. glandibracteolata , 19 Oct. 1999, coll. S.-P. Quek, SPQ.131b, SPQ.136b, SPQ.139 & SPQ.147, DNA vouchers 4(4) (3 ANIC, 1 FRIM: SPQ.147); Sabah, Crocker Range, past Keningau, Senagang, 450 m, ex M. glandibracteolata & M. indistincta , 20 Oct. 1999, coll. S.-P. Quek, SPQ.148 & SPQ.150, DNA vouchers 2(2) (1 ANIC, 1 FRIM: SPQ.148); Sabah, Crocker Range, Mahua camp, 1000 m, ex M. puberula , 16 Oct. 1999, coll. S.-P. Quek, SPQ.101a, DNA vouchers 2(2); Sabah, Crocker Range, near Majora, Apin Apin, 500 m, ex M. motleyana , 16 Oct. 1999, coll. S.-P. Quek, SPQ.102 & SPQ.103b, DNA vouchers 2(2); Sabah, Crocker Range, near Majora, 500? m, ex M. glandibracteolata & M. indistincta , 16 Oct. 1999, coll. S.-P. Quek, SPQ. 108b, SPQ.110a & SPQ.111, DNA vouchers 4(4) (3 ANIC, 1 FRIM: SPQ.110a); Sabah, Crocker Range, Tambunan to Kota Kinabalu road, Rafflesia Reserve, 1200 m, ex M. petanostyla , 15 Oct. 1999, coll. S.-P. Quek, SPQ.100b, DNA voucher 1(1); Sabah, Crocker Range, Tambunan to Kota Kinabalu road, 200 m, ex M. bancana , 24 Oct. 1999, coll. S.-P. Quek, SPQ.174b, DNA voucher 1(1); Sabah, Crocker Range, Tambunan to Ranau road, 15 km from Ranau, ex M. indistincta , 22 Oct. 1999, coll. S.-P. Quek, SPQ.155, DNA voucher 1(1) ( FRIM); Sabah, Crocker Range, Tambunan to Trusmadi trail, 1200–1300 m, ex M. angulata , M. indistincta & M. puberula , 23–24 Oct. 1999, coll. S.-P. Quek, SPQ.157, SPQ.159, SPQ.160 & SPQ.161, DNA vouchers 4(4).

Note. The holotype is not a perfect specimen but was chosen for three reasons: (1) it shows the diagnostic features of the species, (2) it is a DNA voucher specimen included in the analysis of Quek et al. (2017), and (3) it is from northwest Borneo, a region where this species is well sampled. The collection site probably was along Jalan Kampung Tikolod at about 5° 38'18" N and 116° 16'15" E. Although the description below is based on measurements of specimens from a range of localities in Borneo, our type series is restricted to specimens from the Crocker Range in Sabah, for the following reasons. Subsequent, especially molecular, research on further samples of these coccids may show cryptic species or subsequent authors may have a more restrictive species concept. Species delineation is problematic because of parthenogenesis (as explained in the Introduction) and thus deciding whether there is one variable species or several more tightly defined species is highly subjective.

Other material studied. BORNEO: Brunei, Batu Apoi Forest Reserve, near KBFSC, 4° 33' N, 115°09' E, ex M. trachyphylla & Macaranga sp., 7, 9–11, 26–28 Aug. 1995, coll. P.J. Gullan, PJG-B15: 5(2 adult females & 16 first-instar nymphs), PJG-B22: 8(7 adult females, including 1 on slide with C. macarangae , & 6 first-instar nymphs), PJG-B23: 6(4 adult females & 28 first-instar nymphs) (coll. P.S. Cranston), PJG-B26: 1(1), PJG-B28: 2(1 adult female & 1 nymphal female), PJG-B51: 1(1), PJG-B52: 2(2), PJG-B56: 6(3 adult females, 2 nymphal females & 8 first-instar nymphs); East Kalimantan, Bukit Bangkirai, 1° 1.497' S, 118° 51.949' E, <100 m, ex M. pearsonii , 13 Feb. 2005, coll. S.-P. Quek, SPQ.727, DNA voucher 1(1); East Kalimantan, Samarinda, Tenggarong to Kota Bangun Road, <100 m, ex M. pearsonii , 11 June 2001, coll. S.-P. Quek, SPQ.321, DNA voucher 1(1); East Kalimantan, Wanariset to Bukit Bangkirai Road, 00° 59.29' S, 116° 55.47' E to 1° 0.72' S, 116° 52.04' E, <100 m, ex M. hosei , M. hypoleuca & M. pearsonii , 13 Feb. 2005, coll. S.-P. Quek, SPQ.722, SPQ.723 SPQ.724a & SPQ.725, DNA vouchers 4(4); North Kalimantan, Long Ampung, Sungai Anai trail, 1° 42.96' N, 114° 57.30' E & 1° 42.97' N, 114° 57.23' E, 700 m, ex M. glandibracteolata & M. beccariana , 9 Feb. 2005, coll. S.-P. Quek, SPQ.696, & SPQ.698a DNA vouchers 2(2); North Kalimantan, Long Ampung, Sungai Selungei trail, 1° 42.27' N, 114° 58.90' E, 700 m, ex M. glandibracteolata , 10 Feb. 2005, coll. S.-P. Quek, SPQ.714, DNA voucher 1(1); West Kalimantan, Siduk to Nanga Tayap, 1° 21.72' S, 110° 12.10' E & 1° 21.67' S, 110° 12.30' E, <100? m, ex M. indistincta / velutina , M. aethëadenia & M. hosei , 22 June 2001, coll. S.-P. Quek, SPQ.408a, SPQ.412a & SPQ.415, DNA vouchers 3(3); Sabah, Poring, ~ 6° N, 116° E, ex M. pearsonii & M. indistincta , 30.x.1992 & 25.xi.1992, coll. B. Fiala, #251a, 252a & 253a, 8(8); Sabah, Poring, ex M. pearsonii, Apr. 2001 , coll. B. Fiala, #120 (TK0036) & #128 (TK0038), DNA vouchers 3(3); Sabah, Poring, ex M. glandibracteolata , M. indistincta & M. pearsonii , 17 Apr. 2001, coll. B. Fiala, #39 (TK0021), #40 (TK0023), #48 (TK0043), #106b (TK0031), #110 (TK0035), DNA vouchers 5(5); Sabah, Poring, ex M. pearsonii & M. winkleri , 18 Apr. 2001, coll. B. Fiala, #159 (TK0042) & #158 (TK0044), DNA vouchers 2(2); Sabah, Poring, ex M. pearsonii , 24 Apr. 2002, coll. B. Fiala, #120 (TK0099), DNA voucher 1(1); Sabah, Poring Hot Springs, Kipungit waterfall, ex M. beccariana, Dec 1994 , coll. H.-P. Heckroth, #598, 1(2 adult females on slide with 5 adults of C. penangensis ); Sabah, Poring Hot Springs, ex M. indistincta , no date, coll. H.-P. Heckroth, #581 & 589, 2(3, including 1 on slide with 1 adult female of C. near circularis ) ( FRIM); Sabah, Ranau, Kota Kinabalu, roadside, ex M. beccariana , 27 Mar. 2002, coll. B. Fiala. #124 (TK0098), DNA voucher 1(1); Sabah, road from Ranau to Sandakan, ex M. pearsonii, Jan. 1995 , coll. H.-P. Heckroth, #631, 3(3); Sabah, road from Ranau to Sandakan, ex M. pearsonii , no date, coll. H.-P. Heckroth, #529, 1(4) ( FRIM); Sabah, 10 km south of Ranau, ex M. pearsonii, 1995 or no date, coll. H.-P. Heckroth, #525, 526, 527, 614, 635 & 1203, 6(23, including 2 on slide with 1 adult female of C. secretu s) ( FRIM); Sabah, 10 km south of Ranau, ex M. winkleri , no date, coll. H.-P. Heckroth, #646, 1(5) ( FRIM); Sabah, 60.5 km south of Ranau, ex M. hypoleuca, 1995 , coll. H.-P. Heckroth, #629, 1(1 on slide with 1 adult female of C. secretus ) ( FRIM); Sabah, Tawau, Air Panas, ex M. motleyana , 15 Mar. 2002, coll. B. Fiala, #55b (TK0110), DNA voucher 1(1); Sabah, Tawau Hills, ex M. glandibracteolata , M. indistincta & M. pearsonii , 5–7 Apr. 2001, coll. B. Fiala, #76 (TK0025), #78b (TK0029), #77 (TK0026), #90 (TK0027), #95 (TK0033), DNA vouchers 5(5); Sabah, Tawau, ex M. indistincta & M. umbrosa [latter now correctly identified as M. lamellata ], 12 & 20 Mar. 2002, coll. B. Fiala, #13 (TK0109) & #78b (TK0104), DNA vouchers 2(2); Sabah, Tawau, Bukit Bombalai, ex M. glandibracteolata , 13 Mar. 2002, coll. B. Fiala, #32 (TK0105), DNA voucher 1(1); Sabah, Tawau, ex M. pearsonii , 30 Aug. 2003, coll. B. Fiala, #64, 4(4); Sabah, Tawau Hills, ex M. triloba [now correctly named M. bancana ], coll. H.-P. Heckroth, #477, 1(3) ( FRIM); Sarawak, 2 km Lambir, ex M. hosei, Dec. 1992 , coll. H.-P. Heckroth, #47 & #85, 3(3); Sarawak, Kubah [national park near Kuching], ex M. aethëadenia, Dec. 1994 , coll. H.-P. Heckroth, #377, 1(3); Sarawak, Lambir, 150 m, ex M. beccariana & M. hosei , 3 Sept. 2001, coll. K. Murase, KM.s03, KM.s06, KM.s21 & KM.s24, DNA vouchers 4(4) ( FDS); Sarawak, Lambir, 150 m, ex M. beccariana & M. hullettii , 2–4 Aug. 2003, coll. T. Itioka, TI.s45, TI.s54 & TI.s58, DNA vouchers 3(3) ( FDS); Sarawak, 2 km Lambir NP, ex M. beccariana, Dec. 1992 , coll. H.-P. Heckroth, #30 & #73, 7(7); Sarawak, 2 km Lambir NP, ex M. hosei, Dec. 1992 , coll. H.-P. Heckroth, #47, 1(1 on slide with 1 adult female of C. heckrothi ) ( FRIM); Sarawak, 3 km Lambir NP, ex M. beccariana, Dec. 1992 , coll. H.P. Heckroth, #44, 1(1); Sarawak, 3 km Lambir NP, ex M. beccariana, Feb. 1993 , coll. H.P. Heckroth, #45, 1(3) ( FRIM); Sarawak, 8 km Lambir NP, ex M. lamellata, Dec. 1992 , coll. H.-P. Heckroth, #106, #107 & #113, 4(4); Sarawak, Serian, "Serian" waterfall [perhaps Ranchan Waterfall], ex M. hypoleuca, Dec. 1994 , coll. H.-P. Heckroth, #404, 1(7); South Kalimantan, Meratus Mts, Kapayang village to Loksado, 381 m & 500 m, 2° 48.86' S, 115° 30.70' E & 2° 48.98' S, 115° 31.13' E, ex M. bancana & M. motleyana , 18 June 2001, coll. S.-P. Quek, SPQ.346b & SPQ.340, DNA vouchers 2(2); South Kalimantan, Meratus Mts, Loksado to Kandangan, 170–211 m, 2° 47.48' S, 115° 25.94' E to 2° 48.21' S, 110° 25.52' E, ex M. motleyana , 18 June 2001, coll. S.-P. Quek, SPQ.356a, SPQ.358a & SPQ.362, DNA vouchers 3(3). PENINSULAR MALAYSIA: Gombak, lower logging road, ex M. hosei, Feb. 1993 , coll. H.-P. Heckroth, #214, 3(3); Gombak, lower logging road, ex M. hosei, Mar. 1993 , coll. H.-P. Heckroth, #270, 6(9) (5 slides in ANIC & 1 slide with 4 adult females in FRIM); Johor, just after Mersing towards Johor Bahru, lowland, ex M. griffithiana , 16 Sept. 1999, coll. S.-P. Quek, SPQ.068a, 2(2); Johor, Labis Air Panas, ex M. hosei , coll. H.-P. Heckroth, #1069, 1(3 on slide with 1 adult female of C. secretus ) ( FRIM); 6 km Rawang, ex M. griffithiana, Feb. 1993 , coll. H.-P. Heckroth, #206, 6(4 adult females + 2 probably third-instar females).

Note. This species was recognised as a variety as well as a separate morphospecies and referred to as “ C. tumuliferus var. C. 84” and "morphospecies C. 214" by Heckroth et al. (1998). Specimens of this new species form Clade 3, referred to as "near tumuliferus ", in Quek et al. (2017, fig. 3). This clade has several very closely related subgroups, which we consider to be geographic variants, which mostly do not appear to exhibit consistent morphological differences (see below under Comments); thus measurements for the description below were based on adult females from across Borneo, despite some variation in live appearance among females (note that live appearance is unknown for most specimens of this species).

Etymology. Early in our study we confused specimens of this species with those of C. tumuliferus , with which it shares many morphological similarities, including the raised areas on the dorsum; hence, we have named this new species "pseudotumuliferus" from the Latin " pseudo -" meaning "false".

Adult female. Unmounted material. In life, adult females varied from bright, shining pink to red ( Fig. 2E ) to brownish-yellow, depending on collection locality and perhaps age. Available adult females preserved in ethanol were bright pinkish red in colour. Body broadly oval to almost circular, having a fairly definite arrangement of dorsal humps ( Fig. 11 A–C) with the marginal row of humps most obvious and the central area of dorsum varying from having humps to almost flat; mature females covered dorsally with a brittle, whitish, glassy secretion, easily broken, moulded into elevations corresponding to those of the body, with secretion in the central part of dorsum variable among specimens from different collections ( Fig. 11 A–C).

Slide-mounted adult female (n=19, all from Borneo and including holotype and 8 paratypes; Fig. 12 ). Body oval to circular, 1.7–3.6 (holotype 2.45) mm long, 1.2–3.2 (holotype 2.06) mm wide.

Dorsum. Derm (dd) completely membranous, weakly areolated, with clear area of each areolation usually 4–7 µm in widest dimension; with a series of humps (oval, elongate to irregularly circular raised areas of cuticle) present in a submarginal row of 23–25 (11 or more rarely 12 on each side, plus always 1 medially on head), 3 or 4 on each side submedially, and sometimes 3 or 4 slightly raised areas medially but poorly defined. Dorsal setae (dset) very short, each 2.5–3.0 µm long with rounded apex, scattered on dorsum; humps difficult to discern on younger slide-mounted specimens. Simple pores (sp) each 2.5 µm wide, scattered evenly on dorsum. Preopercular pores (pop) each 2.5–6.3 (mostly 3.0–5.0) µm wide, often scarce and easily confused with simple pores, present in a small group of 3–18 anterior to anal plates. Dorsal microducts (dmic) in areolations each about 2.0 µm wide, appearing bilocular under high magnification. Anal plates (anplt) each broadly triangular with anterolateral margin usually slightly convex and slightly longer than posterolateral margin, length of each plate 1.2–1.8 times width, inner lobes fairly well developed and with a tessellated texture but often difficult to see, each plate 148–183 µm long, 90–140 µm wide, anterolateral margin 135–170 µm long, posterolateral margin 98–133 µm long; each plate with 12–20 dorsal setae (except for specimens from the Meratus Mountains and some from Gombak; see Comments section for variation), each seta 10–23 µm long. Anal ring (ar) bearing 10 setae, each 85–125 µm long.

Margin. Eyespots present slightly removed from dorsal margin, each 17.5–22.5 µm in maximum dimension, often difficult to detect. Marginal setae (mset) variable in length and robustness among specimens, sharply spinose to flagellate, usually present in 2 or 3 rows, rarely (Meratus Mountains specimens only) in an irregular single row on part of margin, each seta 12–65 µm long, with 25–63 setae between anterior and posterior stigmatic areas on each side of body. Stigmatic setae (stgset) shorter than marginal setae, usually numbering 3 per cleft, occasionally fewer (but often lost from DNA vouchers), spinose with rounded apices, all setae subequal in length or median seta slightly longer, each 3.0–18.0 (mostly 7.5–12.5) µm long.

Venter. Derm membranous. Ventral setae (vset) slender, longest submedially on posterior abdominal segments, each 15–80 µm long, elsewhere shorter, 10–30 (mostly ≤20) µm long. Interantennal setae numbering 2 pairs, each seta 12–15 µm long. Ventral tubular ducts (vtd) present in a broad submarginal band; each duct with outer ductule 17–20 µm long, inner ductule 15–20 µm long, and duct opening about 2 µm wide. Ventral microducts (vmic) each 2.0– 2.5 µm wide, scattered fairly evenly on venter. Pregenital disc-pores (pgp) each with 7–11 loculi, each pore 5– 7 µm wide. Antennae (ant) 7 segmented, each 215–265 µm long; apical antennal segment 37.5–52.5 µm long, with apical prolongation 6–15 µm long and usually ≥ 10 µm on at least 1 antenna (rarely absent); fleshy setae present on last 3 segments. Mouthparts normal; clypeolabral shield 200–258 µm long, 170–228 µm wide; labium 90–115 µm long, 120–150 µm wide. Legs with hind trochanter + femur 148–175 µm long; hind tibia + tarsus 140–188 µm long; all tarsal digitules each 35–45 µm long; claw digitules each 22–30 µm long, claws each 20–27 µm long. Spiracles normal: anterior peritremes each 55–78 µm wide; posterior peritremes each 65–83 µm wide. Spiracular pores (spp) each 4–5 µm wide, almost all with 5 loculi.

Comments. Heckroth et al. (1998) pointed out that C. tumulifer us and C. tumuliferus var. C. 84 were morphologically very similar and presumably closely related, and although they thought that these two Coccus species shared the same ant partner, the associations now are known to be more complex ( Quek et al. 2017). Heckroth et al. (1998) recorded C. pseudotumuliferus (as var. C. 84) only on Borneo, in both secondary and primary forest, from 14 species of Macaranga but most abundant on M. beccariana . Also, as stated in Quek et al. (2017, table S7), morphospecies C. 214 of Heckroth et al. (1998) resembles C. pseudotumuliferus . PJG examined three Heckroth specimens (collection #214 from M. hosei at Gombak, Peninsular Malaysia) of this morphospecies and considered them to be identifical to adult females from Heckroth collection #270 (also from M. hosei at Gombak), as well as sufficiently similar to Bornean collections of C. pseudotumuliferus to be treated as a geographic form of this species, at least until further information is available. On Borneo, Heckroth et al. (1998) recorded morphospecies C. 214 exclusively from primary forest, but on Peninsular Malaysia, where there is little primary forest left, the few collections were from secondary forest. The only Heckroth specimens from Borneo slide-mounted and identified by him as C. 214 are apparently C. near circularis (#522, 573, 640, 1140 & 1410) and probably C. penangensis (#550), all from Poring Hot Springs (difficult to identify as many specimens are poorly cleared). We assume that Heckroth first recognised his morphospecies C. 214 based on females from Gombak that have this 214 collection number, but later decided that the morphospecies also occurred on Borneo. Probably on Borneo it may have been confused with C. near circularis and C. penangensis because Heckroth's doctoral thesis states that C. 214 is very similar to C. penangensis and cannot be distinguished without slide preparation. As discussed above in the note following the list of specimens examined, there is a small amount of genetic variation among specimens of C. pseudotumuliferus from different areas of Borneo as well as some variation in live appearance (although live appearance is poorly recorded). Morphological variation across the geographic range of this species is discussed in the third paragraph below. Our type series is restricted to specimens from the Crocker Range in Sabah largely because of variation across the range of specimens that we consider to be this species (see Note after the listing of type specimens above).

Adult females of C. pseudotumuliferus from Borneo can be distinguished from all other species of Coccus known from Macaranga by having the combination of (i) very short dorsal setae that can appear to be absent; (ii) 11, rarely 12, dorsal submarginal raised areas (humps) on each side of body plus 1 medially on head (most obvious on non-slide-mounted specimens); (iii) usually 12–20 setae per anal plate; and (iv) one or both apical antennal segments with a prolongation typically ≥ 10 µm long. Adult females of C. pseudotumuliferus are most similar to the adult females of C. caviramicolus , C. secretus and C. tumuliferus , which also have extremely short dorsal setae, but adult females of C. pseudotumuliferus differ from those of C. caviramicolus in having marginal setae mostly tapering to a point (fimbriate in C. caviramicolus ); from C. secretus in having dorsal setae rounded at the apices (tapering to a point in C. secretus ) and the dorsal setae of the anal plates much shorter (15–45 µm long in C. secretus as compared with 10–23 µm long in C. pseudotumuliferus ); and from C. tumuliferus in the number of submarginal raised areas (8 on each side plus 1 medially on head in C. tumuliferus as compared with 11, rarely 12, in C. pseudotumuliferus ), the shape of these raised areas (usually oval to elongate in C. pseudotumuliferus but more circular in C. tumuliferus ), and the number of stigmatic setae (often only 1 per cleft in C. tumuliferus as compared with mostly 3 in C. pseudotumuliferus ).

There is a lot of variation in the length and robustness of the marginal setae, which range from short (15–25 mm long) rather robust setae to longer (50–80 µm) slender and usuallly flagellate setae, even among females collected from different plants in a single locality. However the molecular data ( Quek et al. 2017) strongly suggest that this variation in the marginal setae is not indicative of cryptic species because specimens that are identical in the nuclear genes sequenced can have different marginal setae. However, five specimens from the Meratus Mountains in the Indonesian province of South Kalimantan, that form a poorly supported subclade in the data of Quek et al. (2017), have fewer setae on the anal plates (7–10 per plate compared with the usual 12–20) and their marginal setae tend to be in a single row, especially on the head and thorax. A number of adult females from Peninsular Malaysia (see list in 'Material examined' above) represent the only record of this species from outside of Borneo. However, they differ from Bornean collections of C. pseudotumuliferus in having their marginal setae mostly in a single row. Furthermore, specimens from near Mersing in the Johor area (SPQ.068a) and from near Rawang (Heckroth #206) have marginal setae mostly with fimbriate apices, whereas specimens from Gombak (Heckroth #214 and #270) have mostly flagellate marginal setae (occasionally each up to 60 µm long) with just a few fimbriations on some setae. Additionally, the anal plate setae on specimens from Gombak number only 7–13 and these females also have an indistinct pattern of dorsal raised areas. Unfortunately there are no DNA data available for any of these Peninsular Malaysian specimens to suggest whether the morphological differences reflect genetic distinctness from the Bornean populations of C. pseudotumuliferus .

Coccus pseudotumuliferus may occur on rare occasions inside the hollow stems of non- Macaranga host plants (see Heckroth et al. (1998) for morphospecies C. 214), but PJG and TK have not seen these coccid specimens to verify their identity.

Figures

FIGURE 2. Live coccids and ants inside the cut stems of Macaranga plants: A, C. macarangae Morrison, in M. winkleri, Luasong, Sabah, Borneo (E. Linsenmair, 2003 BF #96); B, C. macarangae, in M. beccariana, Temburong, Brunei, Borneo (P.J. Gullan, PJG-B16); C, C. penangensis Morrison, in M. glandibracteolata, Poring, Sabah, Borneo (B. Fiala, 2003 BF #2); D, C. penangensis, in Macaranga sp., Temburong, Brunei, Borneo (P.J. Gullan, PJG-B31); E, C. pseudotumuliferus Gullan & Kondo sp. n., in M. pearsonii, Tawau, Sabah, Borneo (B. Fiala, 2003 BF #64); F, C. secretus Morrison, in M. indistincta, Tawau, Sabah, Borneo (B. Fiala, 2003 #95). All photographs taken by the collector specified in parentheses.

FIGURE 11. Photographs of the dorsum of adult females preserved in ethanol: A–C, C. pseudotumuliferus Gullan & Kondo sp. n., showing variation in the numbers and shapes of dorsal elevations or humps; D, C. tumuliferus Morrison.

FIGURE 12. Adult female of Coccus pseudotumuliferus Gullan & Kondo sp. n. Abbreviations as in Fig. 3.