243 research outputs found

    Coccus Linnaeus

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    Coccus Linnaeus urn:lsid:zoobank.org:act: D7AFF012-1B2A-4F9B-BA23-795DCC253C10 Coccus Linnaeus, 1758: 455. Type species: Coccus hesperidum L. Lecanium Burmeister, 1835: 69. Unavailable name. Taiwansaissetia Tao, Wong & Chang, 1983: 77; synonymy by Lin et al. 2013: 259. A full synonymy for Coccus is available from ScaleNet (García Morales et al. 2017). Adult females of all Coccus species from Macaranga share the features of areolations on the dorsal derm, a cluster of preopercular pores anterior to the anal plates, a broad submarginal band of a ventral tubular ducts, and pregenital disc-pores confined to the area lateral to the vulva. Prior to molecular phylogenetic studies, we considered erecting a new genus for these Macaranga -associated coccids but, as discussed in the Introduction above, their close genetic relationship to C. hesperidum (the type species of Coccus) made this action untenable, despite the morphological distinctness of the Macaranga coccid species from C. hesperidum. For example, adult females of C. hesperidum lack ventral tubular ducts in the submarginal area of the body (prominent in the Macaranga coccids) and possess dorsal tubercles (lacking in the Macaranga -associated species).Published as part of 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, pp. 1-51 in Zootaxa 4521 (1) on page 8, DOI: 10.11646/zootaxa.4521.1.1, http://zenodo.org/record/377024

    Coccus Gullan, Kondo, Fiala & Quek, 2018, sp. n.

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    Diagnosis for Coccus species associated with Macaranga. Adult female. Unmounted material (Fig. 2). Insects oval to elongate oval, generally less than 3.0 mm at widest point. Colour in life variable, from white, brown, yellowish-brown to red; covered in a thin layer of wax, with texture either smooth or somewhat finely granulose. Insects rather flat, not becoming convex, often with dorsal elevations. Generally associated with tending ants, and living within hollow stems of Macaranga plants. Slide-mounted material. Body circular, oval or elongate oval, up to 4.5 mm long, 3.1 mm wide at maturity. Dorsum: Derm membranous, generally with conspicuous areolations covering whole dorsum; derm not becoming heavily sclerotised at maturity. Dorsal tubercles and pocket-like sclerotisations absent. Dorsal setae variable, each sharply spinose, with a clubbed or rounded apex, or very short and about the same length as diameter of its setal socket. Dorsal tubular ducts absent. Dorsal microducts small, appearing bilocular under high magnification, each situated in an areolation, and scattered rather evenly throughout dorsum (since only found in areolations). Simple pores present, scattered evenly on dorsum. Preopercular pores scarce to numerous, generally small (each 5–10 µm diameter), present in a small to large, usually elongate, cluster anterior to anal plates, numbering 1–35 pores. Anal plates together quadrate to pyriform, inner lobes generally well developed, with a membranous fold of tessellated texture; each anal plate with 3–26 dorsal setae, 2 hypopygial setae, 2 ventral fringe setae and 1–4 ventral subapical setae. Anal ring almost entire, bearing a total of 8 or 10 setae, with 1 pair (if 8 setae) or 2 pairs (if 10 setae) thinner than the rest (setae difficult to count as obscured by ano-genital fold and easily confused with anterior margin setae). Sclerotised rim around anal plates absent. Margin: Eyespots present or absent, when present located on margin or slightly on dorsal submargin. Marginal setae variable, each usually flagellate or sharply spinose with apex either tapering to a point, bifid or fimbriate, present in 1–3 rows. Stigmatic setae distinct from marginal setae, usually well developed, generally totalling 3, rarely 1 but sometimes up to 4 setae per stigmatic cleft; each seta sharply to bluntly spinose, median seta generally longer than lateral setae, but sometimes all setae subequal in length. Stigmatic clefts generally well defined. Spiracular furrow (or stigmatic groove) with a band of spiracular pores 1–4 pores wide, totalling usually 20–40 pores (number of pores difficult to count on most specimens), each pore 4–6 µm wide and with 2–7 (mostly 5) loculi. Venter: Ventral derm membranous. Ventral setae slender, longest on posterior abdominal segments anterior to vulva. Interantennal setae numbering 2 or 3 pairs. Ventral tubular ducts present in a narrow or broad submarginal band, with tubular ducts also present on both sides of mouthparts (but usually absent anteriorly), across prothoracic and mesothoracic segments, often also present across metathorax or around metathoracic coxae. Ventral microducts present, scattered evenly on venter. Pregenital disc-pores each with 5–10 loculi, restricted to a small area on each side of genital opening. Antennae each 5–8 segmented. Mouthparts normal for coccids; clypeolabral shield with 1 pair of setae; labium with 4 pairs of setae. Legs normal in structure but often small in relation to body size, without tibiotarsal sclerotisation; tarsal digitules thin, one of each pair slightly thicker than the other, each with apex knobbed; claw digitules both knobbed, one thicker than other; claw denticle absent. Spiracles normal, often each set in a spiracular depression. Spiracular pores each with 3–7 (mostly 5, rarely an occasional pore with 2) loculi. First-instar nymph. We were able to examine the first-instar nymphs of three species of Coccus, C. macarangae, C. penangenis and C. pseudotumuliferus, collected from Macaranga at one locality in Brunei. These nymphs could not be identified to species, although there may be small differences in body size, but our sample sizes were small and all of the nymphs were from one locality. We identified the first-instar nymphs as belonging to a particular species if they were collected from under the venter of an identified adult female. This was important because often more than one Coccus species occurred inside each Macaranga plant. Only the first-instar nymph of C. macarangae has been described and illustrated in the taxonomic section below. There is no modern and detailed description of the first-instar nymph of the type species of Coccus, C. hesperidum, although Borchsenius (1957) has a short description and Annecke (1966) provides a taxonomic illustration with reasonable detail. The available first-instar nymphs of Coccus species from Macaranga resemble the nymph of C. hesperidum illustrated by Annecke (1966) except that the latter is shown with spiracular pores of five loculi (four loculi in Macaranga nymphs) and quite straight marginal setae (there is a distinct bend at half-length in the marginal setae of Macaranga nymphs). Slide-mounted material. Body oval to elongate oval, 330–490 µm long, 190–330 µm wide; stigmatic areas distinct, but sclerotised clefts absent. Dorsum. Derm membranous. Dorsal setae minute, present in 2 submedial longitudinal rows on thorax and first abdominal segment. Trilocular pores on head not detected, but instead a pair of tiny simple pores present. Dorsal microducts and preopercular pores absent. Anal plates each longer than wide, elongate triangular with anterolateral margin subequal in length to posterolateral margin, and anterior margin rounded; each plate with 2 small dorsal setae near apex, 1 inner margin seta and an apical seta up to 150 µm long. Anal ring bearing 6 setae. Margin. Eyespots present as pigmented spots on margin. Marginal setae robust flagellate and curved to bend posteriorly, present in 1 row, with always 2 setae between anterior and posterior stigmatic areas, 12 (rarely 11 or 13) on head between anterior stigmatic areas, and 8 on each side of abdomen. Stigmatic setae numbering 3 in each cleft, thicker than marginal setae; median seta longest, 2.0– 2.4 µm thick, spinose, slightly curved towards apex and apically rounded; lateral setae small, each ≤ 5 µm long, apically rounded. Venter. Derm membranous. Ventral setae slender, 3 pairs of pregenital setae longest, other setae mostly minute, present in a marginal row around body and a submarginal row on each side of abdomen. Interantennal setae numbering 1 pair consisting of a single seta near base of each antenna. Ventral tubular ducts absent. Ventral microducts sparse, present submarginally with each side of body having 1 microduct between ventral setae on most abdominal segments, 2 microducts between anterior and posterior stigmatic areas, and 1 present between base of antenna and body margin. Disc-pores, except spiracular pores, absent. Antennae each 6 segmented; fleshy setae present on last 3 segments. Mouthparts typical of coccid nymphs. Legs normal; tarsal and claw digitules each with a small knobbed apex; claw with small denticle present. Spiracles normal; spiracular furrows each with 3 (anterior furrow) or 4 (posterior furrow) spiracular pores in a line. Male instars. No adult or immature males were collected. One second-instar male and one pupa with a pharate adult male plus its prepupal exuviae were collected by PJG in Brunei from two different Macaranga plants but these belong to an undescribed species of Myzolecanium Beccari (Gullan et al. 1993; Hodgson 1994). No other collections examined for this study contained any male coccids, and no publications on the Macaranga coccids report males; therefore, it is assumed that these Coccus species are parthenogenetic. Species recognition. Delimitation of the Coccus species from Macaranga is confounded by geographic variation, probably exacerbated by lack of gene flow due parthenogenetic reproduction (as discussed in the Introduction). Several morphological features of adult females that typically are used to separate other Coccus species can vary substantially within a species in the Macaranga coccids, e.g., the shape and length of the marginal body setae and the number and size of the preopercular pores found anterior to the anal plates. The recent availability of a phylogenetic reconstruction for these coccids based on nuclear genes (Quek et al. 2017) greatly assisted in recognising which morphological character states are taxonomically informative; thus, for this revision, we recognise each of the numbered clades presented in figure 3 of Quek et al. (2017) as a species, although four of them are based on very limited specimen sampling. In the case of clade 4 (C. near circularis) and clade 9 (C. near macarangicolus), we believe that these specimens are geographic variants of their respective species. Coccus circularis was described from a few specimens from Singapore (Morrison 1921) and C. macarangicolus was based on just a few specimens from Kuala Lumpur (Takahashi 1952), whereas the sequenced specimens are all from Borneo. We describe specimens of clade 3 (C. near tumuliferus), clade 7 (C. sp. Y) and clade 8 (C. sp. X) as three new species, below. Specimens of one new species represented in clade 3 of Quek et al. (2017) exhibit morphological variation of cuticular features and their live appearance also varies (see under C. pseudotumuliferus below). This may have led H.-P. Heckroth (Heckroth et al. 1998; Heckroth, unpublished data) to recognise two morphospecies (C. tumuliferus var. C. 84 and C. 214) for specimens that we are assigning to a single species. As typical for all coccids, species of Coccus from Macaranga have been described based solely on the morphology of the adult females. Third-instar females of these Coccus species can be very difficult to distinguish from their adult females because they can be similar in size and have similar body setae. However, immature female soft scales rarely have multilocular pores near the vulva and their antennae each can have one or more fewer segments than those of adult females. Morphospecies C. 41. Specimens referred to by this code number in Heckroth et al. (1998) were reported as found almost exclusively on the Bornean endemic M. winkleri (section Winklerianae), which was inhabited by Crematogaster morphospecies 8. This ant species is not part of the Crematogaster borneensis -group and its placement is unclear (Feldhaar et al. 2016). All specimens of C. 41 (Heckroth collections numbers 41, 153 and 155) were collected in December 1992 in Lambir Hills National Park. All examined individuals of C. 41 are unusual in having the diagnostic features of a third-instar female (no multilocular pores near the vulva and 6- segmented antennae) but usually a body of the typical size of an adult. We hypothesise that they are probably immature specimens of C. penangensis, which may have failed to moult quickly to the adult due to some unsuitability of the host M. winkleri. Heckroth et al. (1998) reported that C. 41 never occurred in the same plant as C. penangensis (see their table 3, p. 436), although C. penangensis did occur on M. winkleri (fig. 1b, p. 434), and that C. 41 was uncommon. An immature female of C. macarangae (ID based in 12S DNA) sent by B. Fiala (No. 96) collected from M. winkleri at Tawau Hills, Sabah, on 7 April 2001, looks adult-like due to its body size (more than 2 mm long) and well-developed dorsal areolations; however, it also has no multilocular pores near the vulva and has 6-segmented antennae. This suggests that M. winkleri may generally be a poor host for the development of these Coccus species.Published as part of 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, pp. 1-51 in Zootaxa 4521 (1) on pages 9-11, DOI: 10.11646/zootaxa.4521.1.1, http://zenodo.org/record/377024

    Coccus pseudotumuliferus Gullan & Kondo 2018, sp. n.

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    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.Published as part of 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, pp. 1-51 in Zootaxa 4521 (1) on pages 34-39, DOI: 10.11646/zootaxa.4521.1.1, http://zenodo.org/record/377024

    Coccus heckrothi Gullan & Kondo 2018, sp. n.

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    Coccus heckrothi Gullan & Kondo sp. n. urn:lsid:zoobank.org:act: 86F500BF-88D0-44F0-AE47-F4E76687E9AB (Fig. 5) “ Coccus sp. Y”, Quek et al. 2017: 823. Type material examined. Holotype: adult female, BORNEO: Sarawak, 2 km from Lambir Hills National Park in direction of Miri, in hollow stem of Macaranga hosei, Dec. 1992, coll. H.-P. Heckroth, #85, 1(1) (FRIM). Paratypes: BORNEO: same data as holotype, 12(12) and 2(2 third-instar females) (ANIC except 4 adult females to FRIM); 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. pseudotumuliferus) (FRIM); Sarawak, 3 km Lambir, in hollow stem of Macaranga trachyphylla with C. secretus, Dec. 1992, coll. H.-P. Heckroth, #92, 3(3) (ANIC); Sarawak, 3 km Lambir, in hollow stem of Macaranga hosei, Feb. 1993, coll. H.-P. Heckroth, #50, 1(4) (FRIM); Sarawak, Lambir, ex M. rufescens & M. hosei, 3 Sept. 2001, coll. K. Murase, KM.s11 & KM.s22, DNA vouchers 2(2) (FDS); Sarawak: Lambir, ex M. rufescens (TI.s42 only) & M. hosei, 2–4 Aug. 2003, coll. T. Itioka, TI.s42, TI.s49, TIs.50, TI.s51 & TI.s52a, DNA vouchers 5(5) (FDS). Other material examined. BORNEO: Sabah, Crocker Range, Keningau, 1100 m, ex M. puberula, 17 Oct. 1999, coll. S.-P. Quek, SPQ.113, DNA voucher 1(1); Sabah, Crocker Range, ex M. pearsonii, 13 Apr. 2001, coll. B. Fiala, #8 (TK0016), DNA voucher 1(1); North Kalimantan, Long Ampung, Sungai Anai trail, 700 m, 1° 42.964' N, 114° 56.943' E, ex M. aëtheadenia & M. bancana / M. indistincta, 9 Feb. 2005, coll. S.-P. Quek, SPQ.701, SPQ.704 & SPQ.705, DNA vouchers 3(3). Note. Specimens of this new species form Clade 7, which is referred to as “ Coccus sp. Y” in Quek et al. (2017, fig. 3). This clade has two very closely related subgroups, which are considered to be geographically separated populations—one from Sarawak and Sabah, and the other from North Kalimantan. This species appears to be common in and near Lambir Hills National Park in Sarawak. Etymology. During his Ph.D. studies, Hans-Peter Heckroth recognised this species as Coccus morphospecies 85, but did not specifically refer to it in his publications on the Macaranga coccids (Heckroth et al. 1998, 2001). Here we name the species after him in recognition of his pioneering research on the Macaranga coccids. Adult female. Unmounted material. Not seen. Slide-mounted adult female (n = 10, all from Lambir and including holotype; Fig. 5). Body elongate oval, 1.9–3.5 (holotype 1.9) mm long, 1.4–2.9 (holotype 1.4) mm wide. Dorsum. Derm (dd) with circular to oval areolations, mostly each 10–40 µm in greatest dimension, and with lightly sclerotised submarginal lines radiating inwards at right angles to margin, often obvious only on more mature females. Dorsal setae (dset) slender, each 6–23 (mostly 10–15) µm long, sparsely scattered on dorsum. Simple pores (sp) each 2.5–3.2 µm wide, scattered evenly on dorsum. Preopercular pores (pop) each 4–7 (mostly 5) µm wide, present in an elongate cluster of 8–16 pores anterior to anal plates. Dorsal microducts (dmic) in areolations each 1.8–2.2 µm wide, appearing bilocular under high magnification. Anal plates (anplt) each triangular and typically with slightly convex margins, anterolateral margin always slightly longer than posterolateral margin, length of each plate 1.5–1.9 times width, inner lobes swollen, with a tessellated texture, each plate 165–183 µm long, 85–118 µm wide, anterolateral margin 125–158 µm long, posterolateral margin 105–133 µm long; each plate with 12–23 dorsal setae, each 18–25 µm long. Anal ring (ar) with 10 setae, each 90–120 µm long. Margin. Eyespots, if visible, each 20–35 µm in maximum dimension, on margin but difficult to discern. Marginal setae (mset) in 1 row, most setae sharply spinose, rarely with apices fimbriate or bifurcate, each 12–45 (mostly 20–40) µm long, setae near anal cleft with tendency to be shorter and more fimbriate at apices than setae of rest of margin; with 10–21 setae between anterior and posterior stigmatic areas on side of body. Stigmatic setae (stgset) well developed, sharply spinose, typically numbering 3 per cleft, median setae longest, each 22–43 µm long, lateral setae each 12–28 (mostly>15) µm long. Venter. Derm membranous. Ventral setae (vset) slender, with posterior-most prevulvar pair longest (each seta up to 55 µm long), elsewhere shorter, each 7–20 µm long. Interantennal setae numbering 3 pairs, each seta ≤ 20 µm long. Ventral tubular ducts (vtd) present in a broad submarginal band; each duct with outer ductule 15–20 µm long, inner ductule usually 15–18 µm long, and duct opening about 2 µm wide. Ventral microducts (vmic) each about 2 µm wide, scattered fairly evenly on venter. Pregenital disc-pores (pgp) each with 8–10 loculi, each pore 6–7 µm wide. Antennae (ant) 7 (rarely 6 or 8) segmented, each 225–270 µm long, fleshy setae present on last 3 segments. Clypeolabral shield 258–280 µm long, 205–255 µm wide; labium 110–120 µm long, 130–150 µm wide. Legs with hind trochanter + femur 150–185 µm long; hind tibia + tarsus 150–190 µm long; all tarsal digitules each 30–40 µm long; claw digitules each 20–30 µm long, claws each 24–32 µm long. Spiracles normal: anterior peritremes each 60–80 µm wide; posterior peritremes each 70–88 µm wide. Spiracular pores (spp) each 4–5 µm wide, typically with 5 loculi. Comments. Adult females of C. heckrothi can be distinguished from all the other species of Coccus known from Macaranga by having the combination of (i) anal plates each with 12–23 dorsal setae, each seta 18–25 µm long; (ii) short, slender dorsal setae, each mostly 10–15 µm long and tapering to apex; and (iii) marginal setae in a single row and with each seta sharply spinose, mostly 20–40 µm long and rarely with apex fimbriate or bifurcate. The adult females of C. heckrothi are most similar to those of C. penangensis in the number of dorsal setae on each anal plate (11–23 in C. penangensis) and in having short and slender dorsal body setae (mostly 15 µm (usually 20) preopercular pores. Mealybugs (Pseudococcidae) and coccids have been reported associated with Cladomyrma ants in the hollowed-out stems of Crypteronia griffithii in Peninsular Malaysia (Maschwitz et al. 1991) but the identity of those coccids is unknown.Published as part of 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, pp. 1-51 in Zootaxa 4521 (1) on pages 18-20, DOI: 10.11646/zootaxa.4521.1.1, http://zenodo.org/record/377024

    Coccus

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    Key to adult females of Coccus species associated with Macaranga (Note: this key may be difficult to use unless the specimens are cleared thoroughly of body contents and their cuticle is well stained) 1. Dorsal setae (excluding marginal row) appearing absent but present and very short, most setae less than 2 times as long as width of setal socket, rarely setae near body margin up to 3 times as long as setal socket width, never as long as ventral setae..................................................................................................... 2 - Dorsal setae clearly evident, most setae more than 3 times as long as setal socket width, as long or longer than most ventral setae, but may be shorter than long interantennal and pregenital setae............................................ 5 2. Marginal setae mostly fimbriate at apices. Anal plates with dorsal setae usually confined to posterior half of each plate........................................................................................ caviramicolus Morrison - Marginal setae mostly tapering to a point, apices rarely bifurcate or fimbriate (except on some individuals of C. tumuliferus). Anal plates with dorsal setae usually present on posterior two-thirds of each plate................................... 3 3. Legs reduced, each smaller than mouthparts, hind trochanter + femur 130 µm long. Dorsum covered with oval or circular dermal elevations (Fig. 11), sometimes difficult to see on slide-mounted specimens. Anal plate setae tapered, even if robust, and 8–23 µm long. Antennae 7, rarely 6, segmented.................................................... 4 4. Dorsal submarginal raised areas (humps) very rounded, almost always numbering 8 on each side of body plus 1 medially on head. Stigmatic clefts often each with just 1 long robust seta (≤ 25 µm long, frequently damaged or missing), sometimes 2 lateral setae (each 3–8 µm long), rarely 3 subequal very short setae (≤ 5 µm long). Apical antennal segment with a short (2–8 µm) or often no apical prolongation.......................................................... tumuliferus Morrison - Dorsal submarginal raised areas oval or elongate, usually numbering 11, rarely 12, on each side of body plus 1 medially on head. Stigmatic clefts each usually with 3 setae of subequal length (mostly 7–18 µm long, often damaged or missing). Apical antennal segment with an apical prolongation almost always ≥ 10 µm long on at least 1 antenna............................................................................................ pseudotumuliferus Gullan & Kondo sp. n. 5. Dorsal setae knobbed, or with rounded apices, not tapering to a point. Marginal setae sharply spinose, apices never bifurcate or fimbriate. Hind trochanter + femur 100 µm long.......................................... 6 6. Marginal setae in 2 rows, numerous, with 41–45 between stigmatic areas on each side of thorax, each seta sharply spinose with slightly bent tip. Dorsal setae long and flagellate, each 35–90 µm long................. lambirensis Gullan & Kondo sp. n. - Marginal setae in 1 row, not numerous, with 9 and mostly 15 µm long................................................................... heckrothi Gullan & Kondo sp. n. - Marginal setae with apices mostly bifurcate or fimbriate, rarely sharply spinose. Lateral stigmatic setae each usually <15 µm long............................................................................... penangensis Morrison 9. Dorsal setae rather abundant, sharply spinose, mostly with straight or bent apices, occasionally a few setae with apices bifurcate or fimbriate, each seta 15–40 µm long. Preopercular pores 5–11 µm wide, each generally larger than a pregenital disc-pore................................................................................... macarangae Morrison - Dorsal setae rather sparse, slender, with a flagellate apex, each 15–30 µm long. Preopercular pores usually ≤ 6 µm wide, each about same size as a pregenital disc-pore.................................................... circularis MorrisonPublished as part of 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, pp. 1-51 in Zootaxa 4521 (1) on page 12, DOI: 10.11646/zootaxa.4521.1.1, http://zenodo.org/record/377024

    Coccus lambirensis Gullan & Kondo 2018, sp. n.

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    Coccus lambirensis Gullan & Kondo sp. n. urn:lsid:zoobank.org:act: CBBDC76D-0BBC-4E83-86C3-9BB452F41D79 (Fig. 6) “ Coccus sp. X”, Quek et al. 2017: 823 This species was referred to as “ Coccus sp. X” in Quek et al. (2017) (use of this name is not intended to be for nomenclatural purposes). The holotype listed below is the only known specimen and is a DNA voucher. Its nucleotide sequence data place it as sister to a specimen identified as Coccus near macarangicolus in figure 3 of Quek et al. (2017). Coccus X is sufficiently distinct morphologically to be described below as a new species. Type material examined. Holotype: young adult female, BORNEO: Sarawak, Lambir Hills National Park, ex Macaranga beccariana, 11–15 Aug. 2002, coll. T. Itioka, TI.s39, DNA voucher 1(1) (FDS). Etymology. This species is named after its collection locality, “Lambir”, referring to a site in primary forest in Lambir Hills National Park, called Taman Negara Bukit Lambir in Malay. Adult female. Unmounted material. Unknown. Slide-mounted adult female (n=1; Fig. 6). Body oval, 2.83 mm long, 2.56 mm wide. Dorsum. Derm (dd) membranous, with irregular polygonal areas abutting each other, each with a central areolation, and with faint submarginal wrinkle lines radiating inwards from margin. Dorsal setae (dset) slender, flagellate, each 35–88 µm long, longest from head to anterior abdomen and shortest on area lateral and posterior to anal plates. Simple pores (sp) each about 2.5 µm wide, scattered evenly on dorsum. Preopercular pores (pop) each 3.8–7.5 µm wide, 5 in number, present in a medial cluster anterior to anal plates. Dorsal microducts (dmic) in areolations each about 2.0– 2.5 µm wide, appearing bilocular under high magnification. Anal plates (anplt) each triangular but outer apex of each plate distorted (perhaps during slide preparation), anterolateral margin 1.3 times longer than posterolateral margin and both margins slightly convex, length of each plate 1.8 times width, inner lobes normal, each plate 200 µm long, 110 µm wide, anterolateral margin 162–163 µm long, posterolateral margin 125 µm long; each plate with 9 flagellate dorsal setae, each 10–40 µm long. Anal ring (ar) probably bearing 10 setae (difficult to see), each 75–90 µm long. Margin. Eyespots not detected. Marginal setae (mset) in 2 rows, each seta fairly robust and almost always flagellate at apex, 27–58 µm long, with 41–45 setae between anterior and posterior stigmatic areas on each of body. Stigmatic setae (stgset) well developed, spinose with rounded apices, apparently 3 in number, median setae longest, each 25–45 µm long, lateral setae each 15–35 µm long. Venter (partially missing on holotype). Derm membranous. Ventral setae (vset) slender, mostly each 10–25 µm long, except prevulvar setae each up to 88 µm long. Interantennal setae missing due to damage to ventral cuticle. Ventral tubular ducts (vtd) present in a broad submarginal band; each duct with outer ductule 17–18 µm long, inner ductule about 20 µm long, and duct opening about 2 µm wide. Ventral microducts (vmic) each about 2 µm wide, scattered fairly evenly on venter. Pregenital disc-pores (pgp) each 6–8 µm wide and mostly with 10 (occasionally 6, 8 or 9) loculi. Antennae (ant) 7 segmented, each 320 µm long; fleshy setae present on last 2 segments. Clypeolabral shield missing; labium ~ 100 µm long, ~ 100 µm wide. Legs with hind trochanter + femur 172–175 µm long; hind tibia + tarsus 195–200 µm long; all tarsal digitules each 47–58 µm long; claw digitules each 20–23 µm long, claws each 27–28 µm long. Spiracles normal: anterior peritremes each 62–63 µm wide; posterior peritremes each 72–73 µm wide. Spiracular pores (spp) each 4–5 µm wide, usually with 5 (rarely 3 or 4) loculi. Comments. The adult female of C. lambirensis can be distinguished from all other species of Coccus found on Macaranga by its possession of numerous marginal setae in two rows and its long flagellate dorsal setae, each mostly 40–80 µm long, that are longer than the dorsal setae of all other Macaranga coccids. Although adult females of C. pseudotumuliferus and C. tumuliferus group can have marginal setae in two rows, these two species have very short dorsal setae, each of about the same length as the diameter of its setal socket. In addition, in C. lambirensis, each dorsal areolation (clear area with microduct at centre) is situated on an irregularly shaped polygonal area with the abutting polygonal areas giving a cellular appearance to the derm, whereas in other species of Coccus from Macaranga the dorsal areolations are situated on uniform derm. Although molecular data placed C. lambirensis as sister to a specimen (SPQ.392 from Siduk in West Kalimantan) identified as C. near macarangicolus, it differs from that specimen and the examined syntype of C. macarangicolus (from Kuala Lumpur) in the length and shape of its dorsal setae as well as in the number of marginal setae (one row in C. macarangicolus). An unidentified adult female from Poring in Sabah (B. Fiala No. 30a) is similar to C. lambirensis in having a double marginal row of setae but its dorsal setae are shorter and slightly more robust and it has more than 30 preopercular pores anterior to the anal plates. The single adult female most similar to the holotype of C. lambirensis is from Panga in Thailand (probably Phang Nga province) on Xylocarpus granatum (Meliaceae), collected 8 March 2006 by Numakura and sent to TK. The latter unidentified specimen has similar dorsal and marginal setae to the holotype of C. lambirensis, with 45–57 marginal setae between the anterior and posterior stigmatic furrows on each side of body, but has more than 30 preopercular pores anterior to the anal plates. It is possible that Macaranga is not the usual host-plant genus of C. lambirensi s.Published as part of 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, pp. 1-51 in Zootaxa 4521 (1) on pages 20-22, DOI: 10.11646/zootaxa.4521.1.1, http://zenodo.org/record/377024

    Coccus circularis Morrison

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    Coccus circularis Morrison urn:lsid:zoobank.org:act: DD61FCBF-DC4A-4EDD-818E-9E3875A97319 (Fig. 4) Coccus circularis Morrison, 1921: 665. Type material examined. Holotype: adult female, SINGAPORE: in hollow stems of Macaranga sp., date not given, coll. I.H. Burkill, Holotype 1(1) (USNM). Paratypes: SINGAPORE: same data as holotype, 2(1 adult female + 1 first-instar nymph) (USNM); in hollow stems of Macaranga triloba [now correctly named M. bancana], date not given, coll. I.H. Burkill, 1396, 1(1) (USNM). Examined non-type material from original collection. SINGAPORE: in hollow stems of Macaranga griffithiana, date not given, coll. I.H. Burkill, 1389, 1(2 adult females + 2 nymphs) (BMNH). These four BMNH specimens have similar data to the type collections listed above and one of the same Burkill collection numbers as provided by Morrison (1921) in his original description of this species, but they are not type specimens, as explained in the Materials and methods. Other material examined: PENINSULAR MALAYSIA: Negeri Sembilan, Felda Pasoh, 10 µm long and with a flagellate apex; (ii) marginal setae in a single row and with each seta spinose, 17–38 µm long and with a pointed, divided or fimbriate apex; and (iii) anal plates each with 4–7 setae, each seta ≤ 30 µm long. Adult females of C. circularis are most similar to the adult females of C. macarangae in having fewer than 10 setae on each anal plate and some or many marginal setae that are apically fimbriate or bifurcate, but differ from C. macarangae in that the dorsal setae are sparser, generally shorter (≤ 30 µm long) and more slender, and the preopercular pores are usually smaller (5–6 µm wide) and scarce (≤10) in C. circularis (C. macarangae with some dorsal setae up to 40 µm long and 20–50 preopercular pores each 5–11 µm wide). Burkill's collections of this species were from two Macaranga species from Singapore. No specific locality was provided on the labels but, during Burkill's time, collections may have been made at various places on the island because there would have been more native vegetation present. Heckroth et al. (1998) mentioned only a single specimen of this species and provided no further information on it. Only specimens from three collections from the Crocker Range in Sabah on Borneo were sequenced for the phylogenetic reconstruction of Quek et al. (2017, fig. 3, Clade 4). These three adult females, referred to as C. near circularis, are very similar morphologically to specimens from Singapore and one from Peninsular Malaysia, but have a greater number (19–27) of marginal setae between the anterior and posterior spiracular clefts on each side of the body compared with the Singaporean and Peninsular Malaysian specimens (5–12 setae), generally shorter marginal setae (12–25 µm long, but mostly 13–20 µm, versus 17– 38 µm long), shorter dorsal setae (10–15 µm versus 15–30 µm long) and often more dorsal setae on each anal plate (6–11 versus 4–7). The specimens from the Crocker Range were collected at higher altitude and are geographically separated from populations found in Singapore and Peninsular Malaysia. There are a number of other specimens from Sabah (from Poring Hot Springs and near Ranau) that seem to be C. near circularis, but most of the specimens are mature, damaged or heavily stained. The latter, however, appear similar to the three females from the Crocker Range, although their marginal setae appear shorter (generally 10–18 µm long). The single specimen from Brunei is also morphologically similar to the type specimens of C. circularis except that it has very few ventral tubular ducts on the head or the posterior abdomen. More samples are required for molecular analysis to determine the extent of any genetic variation among the regions.Published as part of 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, pp. 1-51 in Zootaxa 4521 (1) on pages 14-18, DOI: 10.11646/zootaxa.4521.1.1, http://zenodo.org/record/377024

    Coccus tumuliferus Morrison. All 1921

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    Coccus tumuliferus Morrison urn:lsid:zoobank.org:act: 537C7FF2-8E3B-423A-ADE9-96E5AD7DD249 (Figs 11D, 14) Coccus tumuliferus Morrison, 1921: 655. Coccus tumulifer Lindinger, 1932f: 197. Unjustified emendation; discovered by Williams & Ben-Dov, 2009: 46. Type material examined. Holotype: adult female, SINGAPORE: on Macaranga hypotema [sic, see below], E.E. Green letter dated 8.vi.1916, coll. I.H. Burkill, 1(2, holotype clearly marked and 1 paratype adult female) (USNM). Paratypes: same data as holotype but two slides specify "in hollow stems of M. hypotema ", 4(4 adult females, including 1 on slide with holotype, + 2 slides with a number of first-instar nymphs) (USNM; the 1 slide with 3 paratype females not seen). Morrison's original description refers to five adult females and several mounted "larvae", and this accords with the slides listed above, all of which have Morrison type labels on both the slides and their envelopes. However there is another USNM slide of this species which appears to have been prepared and labelled by Green from original Burkill material as its label starts: "Ctenochiton / tumuliferum / Green, ms. / Part of type / material.)" (details listed immediately below), but these specimens are not mentioned by Morrison in the original description of this species. Examined non-type material from original collection. SINGAPORE: in hollow stems of Macaranga hypotema [sic], coll. I.H. Burkill, 1(2 adult females + 1 third-instar female) (USNM; slide labelled by Green and discussed above); in hollow stems of Macaranga hypoleuca, attended by ants, coll. I.H. Burkill, 2(2) (BMNH). The BMNH also has another 10 slides, with a total of 29 adult females and nymphs, of this Singapore collection from Burkill; PJG examined these slides in 1994 and made notes but did not take measurements. All of these BMNH slides have similar data to the type collection listed above, but they are not type specimens, as explained in the Materials and methods. Note. The USNM type slides give the host plant as M. hypotema and Morrison (1921) listed the host as M. hypolema, but there is no such species. The BMNH slides give the host as M. hypoleuca, which is a member of the Pachystemon section of Macaranga (Davies et al. 2001). Other material examined. BORNEO: Sarawak, Santubong, ex M. hypoleuca, Dec. 1994, coll. H.-P. Heckroth, 445, 447, 450, 451 & 453, 5(14 + 1 adult female of C. macarangae) (2 slides ANIC, 3 slides FRIM). PENINSULAR MALAYSIA: Fraser’s Hill, ex M. hypoleuca, Mar. 1993, coll. H.-P. Heckroth, #284, 3(3); Gombak, lower logging road, ex M. hosei, Feb. 1992, coll. H.-P. Heckroth, #230, 2(2; 1 female taken from the mandibles of an ant); Johor, Kota Tinggi to falls, lowland, ex M. hypoleuca, 5 Sept. 1999, coll. S.-P. Quek, SPQ.018, DNA voucher 1(1); Johor, Mawai camp, Ginger Hill, ~ 1.871 N, ~ 103.954 E, <100 m, ex M. hypoleuca, 7 Sept. 1999, coll. S.-P. Quek, SPQ.027, SQP.030 & SPQ.032, DNA vouchers 3(2 adult females + 1 immature female); Johor, Sedili, 14 km from Route 3, <100 m, ex M. hypoleuca, 5 Dec. 1999, coll. S.-P. Quek, SPQ.182, DNA voucher 1(1); Pahang, near Kuantan on road to Pancing Falls, <100 m, ex M. hypoleuca, 15 Sept. 1999, coll. S.-P. Quek, SPQ.054, DNA voucher 1(1); Pahang, Kuantan to Johor Road, 299 km to Johor, lowland, ex M. pruinosa, 16 Sept. 1999, coll. S.-P. Quek, SPQ.062, SPQ.063a & SPQ.064, DNA vouchers 3(3); Pahang, Kuantan to Johor Road, 299 km to Johor, <100 m, ex M. hypoleuca, 16 Sept. 1999, coll. S.-P. Quek, SPQ.066 & SPQ.067, DNA vouchers 2(2); Terengganu, Bauk Hill, <100 m, ex M. hypoleuca, 12 Sept 1999, S.-P. Quek, SPQ.038, DNA voucher 1(1). SINGAPORE: Old Upper Thompson Road, ~ 1.381 N, ~ 103.813 E, <100 m, ex M. hypoleuca, 4 Oct. 1999, coll. S.-P. Quek, SPQ.090, DNA voucher 1(1). Adult female. Unmounted material. “.. rarely broad oval, but usually broadened behind and triangular with angles rounded; plane of dorsal surface flat, but in dried specimens covered with relatively large knobs having a fairly definite arrangement of a median longitudinal single row and on each side of this two other rows, the outer one forming a continuous row around the body at the margin; dorsally covered with a thin, brittle, whitish but more or less translucent, glassy secretion, very easily broken and usually more or less wanting, molded into elevations and depressions corresponding to those of the body, this covering normally wanting over the flattened extreme margin of the body; body color dull brown, of secretionary covering, as stated, translucent whitish;.. (Morrison 1921: 655). For the present revision, the available adult females preserved in ethanol were pinkish red in colour (Fig. 11D). Although Morrison recorded the body of dry adult females as dull brown, but this is unlikely to reflect the colour in life. Slide-mounted adult female (n=16, including holotype and paratypes; specimens from Santubong in Sarawak excluded; Fig. 14). Body oval to broadly oval, 1.5–2.8 mm long, 1.2–2.4 mm wide, widest in posterior half. Dorsum. Derm (dd) completely membranous, areolated; with a series of humps (oval to circular raised areas of cuticle) present in a submarginal row of 17 (typically 8 on each side plus 1 medially on head), 3 on each side submedially, and usually 3 medially but these often poorly defined. Dorsal setae (dset) very short, each 2–3 µm long with rounded apex, scattered on dorsum. Simple pores (sp) each 2.5–3.0 µm wide, scattered evenly on dorsum. Preopercular pores (pop) each 3.8–5.0 µm wide, scarce, present in a small group of 1–5 anterior to anal plates. Dorsal microducts (dmic) in areolations each 2.0– 2.5 µm wide, appearing bilocular under high magnification. Anal plates (anplt) each triangular with anterolateral margin longer than posterolateral margin, inner lobes fairly developed, with a tessellated texture, each plate 150–198 µm long, 100–125 µm wide, anterolateral margin 140–163 µm long, posterolateral margin 90–128 µm long; each plate with 14–22 dorsal setae, slender spinose, each 8–23 µm long. Anal ring (ar) probably always bearing 10 setae [Morrison (1921: 657) says 8 setae, but the thinner setae are difficult to see], each 100–155 µm long. Margin. Eyespots present slightly removed from dorsal margin, each 20–28 µm in maximum dimension. Marginal setae (mset) of 2 broad types usually not found together on a specimen (but female SPQ.030 with both types): (1) long (20–110 µm) and flagellate, present in 1 or 2 rows (rarely up to 3 rows on part of margin) (as illustrated in Fig. 14); (2) short (12–28 µm long) and with apices mostly fimbriate (or at least divided), usually present in a single row, rarely in 2 rows on parts of margin; with about 7–30 marginal setae between anterior and posterior stigmatic areas on each side of body. Stigmatic setae (stgset) of variable length and development, numbering 1–3 but usually 1 (rarely absent) per cleft, median seta 3–25 µm long, lateral ones if present each 3.0– 7.5 µm long, spinose and tapering to a rounded point, rarely 3 very short setae (≤ 5 µm) present. Venter. Derm membranous. Ventral setae (vset) slender, longest submedially on posterior abdominal segments, each 20–88 µm long, elsewhere shorter, 7.5–40 µm long. Interantennal setae in 2 pairs, each seta 50–70 µm long. Ventral tubular ducts (vtd) present in a broad submarginal band; each duct with outer ductule about 15 µm long, inner ductule about 17.5 µm long, and duct opening about 2.5 µm wide. Ventral microducts (vmic) each about 2 µm wide, scattered fairly evenly on venter. Pregenital disc-pores (pgp) each with 7–9 loculi, each pore 5–7 µm wide. Antennae (ant) usually 7, rarely 6, segmented [Morrison (1921) recorded 8 segments but this appears to be erroneous], each antenna 210–270 µm long; apical antennal segment 30–43 mm long, with apical prolongation 2–8 µm long on at least 1 antenna (often absent on 1 or both antennae of pair); fleshy setae present on last 3 segments when 7 segmented, and on last 2 segments when 6 segmented. Mouthparts normal; clypeolabral shield 210–270 µm long, 170–235 µm wide; labium 80–100 µm long, 120–150 µm wide. Legs with hind trochanter + femur 155– 195 µm long; hind tibia + tarsus 155–200 µm long; all tarsal digitules each 28–40 µm long; claw digitules each 25– 30 µm long, claws each about 30 µm long. Spiracles normal: anterior peritremes each 45–58 µm wide; posterior peritremes each 50–65 µm wide. Spiracular pores (spp) each 4–5 µm wide, mostly with 5 loculi, rarely 4. Comments. Adult females of C. tumuliferus 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) almost always 8 dorsal submarginal raised areas on each side of body plus 1 medially on head (most obvious on live or ethanolpreserved specimens); and (iii) usually 14–22 dorsal setae per anal plate, each slender spinose and mostly ≤ 20 µm long. Adult females of C. tumuliferus are most similar to the adult females of C. caviramicolus, C. pseudotumuliferus and C. secretus, which all also have extremely short dorsal setae, but they differ from females of C. caviramicolus in having marginal setae either flagellate or with few fimbriations (strongly fimbriate in C. caviramicolus) and anal plates with a ratio for the length of the anterolateral margin to posterolateral margin of 1.12 to 1.38 (ratio mostly 1.41 to 2.06 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 are usually much shorter (each 8–23 µm long in C. tumuliferus compared with 15–45 µm long in C. secretus); and from C. pseudotumuliferus in the number of submarginal raised areas (typically 8 on each side plus 1 medially on head in C. tumuliferus compared with usually 11 on each side plus 1 medially on head in C. pseudotumuliferus), the shape of these raised areas (usually circular in C. tumuliferus but oval to elongate in C. pseudotumuliferus), and the number of stigmatic setae (0–3 but usually 1 per cleft in C. tumuliferus compared with usually 3 per cleft in C. pseudotumuliferus). Heckroth et al. (1998) recognised C. tumuliferus from both Borneo and Peninsular Malaysia and recorded it as most common on M. hypoleuca, which is a common and widespread tree in both regions (Davies 2001). We had available for study 16 slide preparations, each containing from one to six adult females, made by Heckroth and identified by him as C. tumuliferus from Borneo. Five slides (#445, 447, 450, 451 and 453, ex M. hypoleuca, Santubong [probably on the lower slopes of Mt Santubong], Sarawak) have adult females of C. tumuliferus, but the other 11 slides (#495, 525, 526, 527, 581, 589, 598, 629, 631, 635 and 614, ex either M. bancana, M. hypoleuca, M. indistincta, M. beccariana or M. pearsonii, from three areas in Sabah) contain specimens of C. pseudotumuliferus. Other than the specimens from Santubong, all other records for C. tumuliferus are from Singapore and Peninsular Malaysia and, given that it is unlikely that Heckroth's specimens of C. tumuliferus are mislabelled, the explanation for this distribution is probably geographic. Santubong is in the region of Sarawak closest to Singapore and West Malaysia and may have been isolated from the rest of Sarawak by past geological and environmental conditions. The Kuching area (including Mt Santubong) of western Sarawak is south of the Lupar Valley and a geological fault called the Lupar Line. This fault marks the southwestern boundary of the rock formation called the Rajang Group in Sarawak (Moss 1998; Wang et al. 2016). The Lupar Valley contains a large river, the Batang Lupar, surrounded by extensive swamp forests, which may have created a barrier to the dispersal of some taxa. A barrier effect of the Lupar Valley has been hypothesised for a species of Malaysian frog for which populations from Peninsular Malaysia and western Sarawak were more genetically similar than populations in western Sarawak were to those in northeastern Sarawak (Zainudin et al. 2010). The adult females of C. tumuliferus from Santubong have 7–9 dorsal submarginal humps on each side of the body and of the typical shape for C. tumuliferus; 12–16 dorsal anal plate setae on the six females for which they are visible clearly; flagellate marginal setae up to 90 µm long (but usually <60 µm) in 1 or 2 rows; 1–4 stigmatic setae per cleft with each seta usually 10–25 (rarely up to 36) µm long; and an apical antennal prolongation 5–8 µm long (rarely 10 µm long on one antenna of a pair); thus these females barely differ in key morphological features from females collected from Singapore and Peninsular Malaysia. Further collecting and molecular assessment of C. tumuliferus from the Kuching region (i.e., south of the Lupar Line) would be valuable. Furthermore, C. tumuliferus may occur in the northwestern part of West Kalimantan but no collections have been made in that region.Published as part of 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, pp. 1-51 in Zootaxa 4521 (1) on pages 43-46, DOI: 10.11646/zootaxa.4521.1.1, http://zenodo.org/record/377024
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