Coccus penangensis Morrison

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 : 30-33

Coccus penangensis Morrison

urn:lsid:zoobank.org:act:D8A35CBE-4EB3-4492-9C2C-38AD45B77D04

( Figs 2C&D , 10 )

Coccus penangensis Morrison, 1921: 657 .

Type material examined. Holotype: adult female, PENINSULAR MALAYSIA: Penang Island, in hollow stems of Macaranga triloba [now correctly named M. bancana ], date not given, coll. I.H. Burkill, ( USNM). Paratypes: same data as holotype, 2(2 adult females + 3 first-instar nymphs) ( USNM). Morrison (1921: 659) states that this species was "described from two mounted adults, three mounted larvae, and few unmounted specimens, ...". One of his three slides has a single adult female and is labelled " Holotype " in Morrison's handwriting (on both the slide and its envelope). Another slide has two adult females and includes the word " Paratype ". We assume that Morrison's overlooked the fact that there were two adult females on the paratype slide when he stated that the description was based on two mounted adults. It seems that this species originally was to be named " Lecanium inquilinum " by Green as this name is crossed out on the slide labels and replaced by " Coccus penangensis ". The slide envelope for the holotype also includes the words: "Green, ms: (Part of type material)".

Examined non-type material from original collections. PENINSULAR MALAYSIA: Penang Island, in hollow stems of Macaranga triloba [now correctly named M. bancana ], coll. I.H. Burkill, No. 2693-b, also labelled: "(type material)", 4(6) ( BMNH); Penang Island, in hollow stems of Macaranga , coll. I.H. Burkill, 1(8) ( BMNH). These two specimen lots have the same data as the type collection listed above and the same data 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. BORNEO: Brunei, Batu Apoi Forest Reserve, 4° 33' N, 115° 09' E, ex M. beccariana , M. trachyphylla & Macaranga sp., 2–31 Aug. & 1 Sept. 1995, coll. P.J. Gullan, PJG-B1: 6(6 adult females), PJG-B6: 16(14 adult females & 2 second-instar females), PJG-B9: 6(4 adult females & 2 third-instar females), PJG-B14: 5(4 adult females & 5 first-instar nymphs), PJG-B28: 1(1), PJG-B30: 21(18 adult females, 1 pharate adult female, 5 first-instar nymphs), PJG-B33: 7(7), PJG-B43: 5(4 adult females & 1 third-instar female), PJG-B45: 5(4 adult females & 5 first-instar nymphs), PJG-B48: 7(7), PJG-B51: 9(9), PJG-B58: 3(3), PJG-B59: 5(5), PJG-B64: 6(6), PJG-B70: 8(7 adult females & 2 first-instar nymphs); Brunei, Batu Apoi Forest Reserve, near KBFSC, 4° 33' N, 115° 09' E, ex Macaranga sp., 23 Aug. 1995, coll. P.S. Cranston, PJG-B44: 5(5 adult females); Brunei, Batu Apoi Forest Reserve, near KBFSC, 4° 33' N, 115° 09' E, ex Macaranga sp., 12 Aug. 1995, coll. C. Maycock, PJG-B31: 3(2 adult females & 1 pharate adult female); North Kalimantan, Long Ampung, Sungei Ampung trail, 1° 43.367' N, 114° 59.838' E, 700 m, ex M. hullettii & M. indistincta , 8 Feb. 2005, coll. S.-P. Quek, SPQ.689 & SPQ.691, DNA vouchers 2(2); North Kalimantan, Long Ampung, Sungei Selungei trail, 1° 42.266' N, 114° 58.898' E, 700 m, ex M. indistincta , 10 Feb. 2005, coll. S.-P. Quek, SPQ.712, DNA voucher 1(1); Sabah, Crocker Range, Keningau, 1200 m, ex M. angulata , 17 Oct. 1999, coll. S.-P. Quek, SPQ.114, DNA voucher 1(1); Sabah, Crocker Range, Keningau to Ulu Kimanis trail, 5.28° N, 115.05° E, 280–1100 m, ex M. angulata , M. bancana & M. indistincta , 19 Oct. 1999, coll. S.-P. Quek, SPQ.130, SPQ.137a, SPQ.141 & SPQ.145, DNA vouchers 4(4); Sabah, Crocker Range, near Majora, 500? m, ex M. indistincta , 16 Oct. 1999, coll. S.-P. Quek, SPQ.105, DNA voucher 1(1); Sabah, Crocker Range, Senagang, past Keningau, ≤ 450 m, ex M. glandibracteolata & M. indistincta , 20 Oct. 1999, coll. S.-P. Quek, SPQ.149 & SPQ.151, DNA vouchers 2(2); Sabah, Crocker Range, forest behind Tambunan, 600 m, ex M. hypoleuca , 18 Oct. 1999, coll. S.-P. Quek, SPQ.119, DNA voucher 1(1); Sabah, Crocker Range, Tambunan to Kota Kinabalu road, 200–1300 m, ex M. angulata , M. bancana , M. petanostyla & M. trachyphylla , 24 Oct. 1999, coll. S.-P. Quek, SPQ.162, SPQ.164, SPQ.165, SPQ.166, SPQ.168, SPQ.170, SPQ.172, SPQ.173a & SPQ.174a, DNA vouchers 9(9); Sabah, Crocker Range, Tambunan to Trusmadi trail, 1300 m, ex M. angulata , 10 Oct. 1999, coll. S.-P. Quek, SPQ.158 & SPQ.158b, DNA vouchers 2(2); Sabah, Crocker Range, Tikolod, 650 m, ex M. indistincta , 18 Oct. 1999, coll. S.-P. Quek, SPQ.127, DNA voucher 1(1); Sabah, Crocker Range, ex M. angulata , 14 Apr. 2001, coll. B. Fiala, #20 = TK0020, DNA vouchers 2(2); Sabah, Crocker Range, ex M. angulata , 14 Apr. 2001, coll. B. Fiala, #31, 4(4, including DNA voucher NH5) (2 ANIC, 2 FRIM); Sabah, Crocker Range, Rafflesia Centre, ex M. angulata , 28 Mar. 2002, coll. B. Fiala, #126 (TK0097) & #134 (TK0096), DNA voucher 2(2); Sabah, Poring, ex M.? depressa [now identified as M. angulata ], Feb. 1992, coll. B. Fiala, #29a, 4(4); Sabah, Poring, ex Macaranga sp. [now identified as M. motleyana ], 16 Apr. 2001, coll. B. Fiala, #36 (TK0034), DNA voucher 1(1); Sabah, Poring, ex Macaranga indistincta , 12 Apr. 2001, coll. B. Fiala, #137 (TK0037), DNA voucher 1(1); Sabah, Poring, ex Macaranga glandibracteolata , 24 Mar. 2002, coll. B. Fiala, #103b (TK0101), DNA voucher 1(1); Sabah, Poring, ex M. glandibracteolata , 24 Aug. 2003, coll. B. Fiala, #2, 3(3) (1 ANIC, 2 FRIM); Sabah, Poring Hot Springs, Kipungit waterfall, ex M. beccariana, Dec. 1994 , coll. H.-P. Heckroth, #598, 1(5 adult females plus 2 adult females of C. pseudotumuliferus ); Sabah, Poring Hot Springs, ex M. indistincta , no date, coll. H.-P. Heckroth, #550, 1(2 adult females, probably C. penangensis but poorly cleared, + 2 nymphs) ( FRIM); Sabah, Tawau, ex M. umbrosa [now identified as M. lamellata ], 15 Mar. 2002, coll. B. Fiala, #19 (TK0106) & #65b (TK0103), DNA vouchers 2(2); Sarawak, Lambir, ex M. bancana , 20 Jan. 1999, coll. K. Murase, KM.s27, DNA voucher 1(1) ( FDS); Sarawak, Lambir, ex M. kingii [now named M. umbrosa ] & M. winkleri , 2–4 Aug. 2003, coll. T. Itioka, TI.s56, TI.s57a & TI.s57b, DNA vouchers 3(3) ( FDS); Sarawak, Lambir, ex M. lamellata , 24 Feb. 1992, coll. B. Fiala, #38a, 2(2); Sarawak, Lambir, ex M. kingii var. platyphylla [now named M. umbrosa ], 24 Feb. 1992, coll. B. Fiala, #44a, 3(3); Sarawak, 8 km Lambir NP, ex M. lamellata, Dec. 1992 , coll. H.-P. Heckroth, #106, #107 &#138, 5(5); Sarawak, 10 km Lambir NP, ex M. hulletti, Dec. 1992 , coll. H.-P. Heckroth, #148, 1(2) ( FRIM); South Kalimantan, Meratus Mts, Loksado area, 2° 48.856' N, 115° 30.695 E, 380 m, ex M. bancana , 18 June 2001, coll. S.-P. Quek, SPQ.345, DNA voucher 1(1); South Kalimantan, Meratus Mts, Loksado to Kandangan, 2° 47.670' N, 115° 28.712' E, 170–200 m, ex M. hullettii , M. indistinct / velutina & M. motleyana , 18 June 2001, coll. S.-P. Quek, SPQ.351, SPQ.352, SPQ.353, SPQ.354, SPQ.356b & SPQ.361a, DNA vouchers 6(6); West Kalimantan, Siduk to Nanga Tayap, 1° 14.672' S, 109° 59.184' E to 1° 22.360' S, 110° 13.686' E, <100 m, ex M. aethëadenia & M. indistinct / velutina , 21 & 22 June 2002, coll. S.-P. Quek, SPQ.393, SPQ.396, SPQ.420 & SPQ.412b, DNA vouchers 4(4). PENINSULAR MALAYSIA: Genting, 900 m, ex M. hullettii, Mar. 1993 , coll. H.-P. Heckroth, #222, 2(1 adult female & 1 first-instar nymph); Gombak Field Studies Centre, ex M. triloba [now correctly named M. bancana ], Mar. 1993, H.P. Heckroth, #269, 1(1) ( BMNH); Gombak, ex M. triloba [= M. bancana ] & M. hullettii , 22 Mar. 1992, coll. B. Fiala, #174a & #175a, 4(4); Gombak, lower logging road, ex M. triloba [= M. bancana ], Feb. & Mar. 1993, coll. H.-P. Heckroth, #251, 5(5) (3 ANIC, 2 FRIM) & #298, 2(2) ( ANIC); Johor, Mawai camp, Ginger Hill, ~ 1.871° N, ~ 103.954° E, <100 m, ex M. bancana , 7 Sept. 1999, coll. S.- P. Quek, SPQ.031, DNA voucher 1(1); Pahang, Beserah Hill nr Kuantan, ~ 3.867° N, ~ 103.347° E, <300 m, ex M. bancana , 14 Sept. 1999, coll. S.-P. Quek, SPQ.047, DNA voucher 1(1); Pahang, Tioman Island, Tekek-Juara trail, 2.805° N, 104.117° E, ex M. bancana , 21 Sept. 1999, coll. S.-P. Quek, SPQ.084, DNA voucher 1(1); Terengganu, Bauk Hill, 200–300 m, ex M. bancana , 12 Sept. 1999, S.-P. Quek, SPQ.037, SPQ.037b, SPQ.040 & SPQ.041, DNA vouchers 4(4); Perak [state], Taiping Hills, in stems of M. triloba [now correctly named M. bancana ], attended by Crematogaster bourensis [sic], 2.iv.1924 (letter), coll. I.H. Burkill, Singapore Field No. 13055, 1(3) ( USNM); Perak [state], Straits Settlement, near Tanjong Malim, on Macaranga triloba [now correctly named M. bancana ], 5.vii.1924 (letter), Singapore Field No. 13489, coll. I.H. Burkill, 1(3) ( USNM). SINGAPORE: Central Catchment, Sime Road, 1° 21'35" N, 103° 48'30" E, ex M. bancana , 18 Mar. 2009, coll. P.S. Cranston, 3(3); Old Upper Thompson Road, ~ 1.381° N, ~ 103.813° E, <100 m, ex M. bancana , 4 Oct. 1999, coll. S.-P. Quek, SPQ.088 & SPQ.089, DNA vouchers 2(2); in hollow stems of Macaranga , coll. I.H. Burkill or C.F. Baker, 3(17 adult females & 1 nymphal female) ( BMNH). SUMATRA: Ketambe, ~ 3° 50' N, 7° 40' E, ex M. triloba [now correctly named M. bancana ], 30 Oct. 1992, coll. U. Maschwitz, sent by B. Fiala, #8a, 2(2); Medan, ~ 3° 40' N, 98° 20' E, ex M. triloba [= M. bancana ], 20 Oct. 1992, coll. U. Maschwitz, sent by B. Fiala, #4a, 3(3); Sibolangit, ~ 3° 40' N, 98° 20' E, ex M. hullettii , 20 Oct. 1992, coll. U. Maschwitz, sent by B. Fiala, #5a & #9a, 8(6 adult females, 1 pharate adult female & 1 third-instar female); Sibolangit, ~ 3° 40' N, 98° 20' E, ex M. triloba [= M. bancana ], 23 Oct. 1992, coll. U. Maschwitz, sent by B. Fiala, #6a, 2(2).

Adult female. Unmounted material. “Normally short oval, flat, dorsal surface dull, naked, wrinkled radially near margin, outer portion light brown, central disk usually much darker brown to blackish; ..” ( Morrison 1921: 657). Morrison’s description of adult females from Penang Island differs in body colour from specimens

photographed in Borneo ( Fig. 2C & D ) probably because Morrison’s specimens were dead and dried. Females from both Brunei and Sabah were pale ochre to salmon pink with a thin dorsal secretion that varied in whiteness among females from different collections. Ridges radiating inwards from the body margin were apparent in all specimens.

Slide-mounted adult female (n=27, including holotype and paratypes; Fig. 10 ). Body oval to elongate oval, 1.6–3.0 mm long, 1.2–2.5 mm wide.

Dorsum. Derm (dd) membranous, with numerous areolations and lightly sclerotised irregular submarginal lines radiating inwards at right angles to margin. Dorsal setae (dset) slender, each 7–38 (mostly ≤20) µm long, scattered on dorsum. Simple pores (sp) each 2.5–3.0 µm wide, scattered evenly on dorsum. Preopercular pores (pop) each 3.5–7.5 (rarely up to 10) µm wide, scarce, present in an elongate group of 6–25 (usually 10–17) 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 broadly triangular with anterolateral margin usually slightly longer than posterolateral margin and latter often slightly convex, length of each plate 1.2–1.7 times width, inner lobes well developed, with a tessellated texture, each plate 120–170 µm long, 85–125 µm wide, anterolateral margin 95–140 µm long, posterolateral margin 85–115 µm long; each plate with 11–23 dorsal setae, each seta 8–20 µm long (12– 18 µm long on holotype). Anal ring (ar) probably always bearing 10 setae [ Morrison (1921: 659) said 8 setae, but the thinner ones are difficult to see], each 75–120 µm long.

Margin. Eyespots indistinct, present on dorsal margin, each 15–28 µm wide, but not detected on some specimens. Marginal setae (mset) in 1 irregular row, each seta short and usually fairly stout, sometimes slender, each usually 7–20 (rarely up to 40) µm long, with apex bifurcate or fimbriate, rarely sharply spinose, with 12–31 setae between anterior and posterior stigmatic areas on each side of body. Stigmatic setae (stgset) well developed, spinose and tapering to pointed or rounded apices, usually numbering 3, median setae usually longest, each 10–28 (mostly 13–20) µm long, lateral setae each 5–18 (mostly 7–15) µm long.

Venter. Derm membranous. Ventral setae (vset) slender, with prevulvar setae each 18–75 µm long, elsewhere shorter, each 8–18 µm long. Interantennal setae numbering 2 or 3 pairs, each seta 7–15 µm long. Ventral tubular ducts (vtd) present in a broad submarginal band; each duct with outer ductule 15–22 µm long, inner ductule 17–20 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 5–9 (mostly 7 and 8) loculi, each pore 5–6 µm wide. Antennae (ant) 7 or 8 (mostly 7, very rarely 6) segmented, each 200–265 µm long; fleshy setae present on last 3 segments. Clypeolabral shield 180–258 µm long, 165–210 µm wide; labium 75–100 µm long, 103–140 µm wide. Legs with hind trochanter + femur 140–170 µm long; hind tibia + tarsus 145–185 µm long; all tarsal digitules each 27–42 µm long; claw digitules each 23–33 µm long, claws each 25–30 µm long. Spiracles normal: anterior peritremes each 45–77 µm wide; posterior peritremes each 55–83 µm wide. Spiracular pores (spp) each 4–5 µm wide, with 3–6 (mostly 5) loculi.

Comments. Adult females of C. penangensis can be distinguished from all other species of Coccus known from Macaranga by having the combination of (i) anal plates each with 11–23 setae, each seta ≤ 20 µm long; (ii) short, slender dorsal setae, each mostly ≤ 20 µm long and tapering to apex; and (iii) marginal setae in a single row with apex of each seta usually bifurcate or fimbriate, and mostly 7–20 µm long. The adult females of C. penangensis are most similar to those of C. heckrothi in the number of dorsal setae on each anal plate (although the setae of C. heckrothi are often slightly longer) and in the shape and length of the dorsal body setae, but differ in that the marginal setae of C. penangensis mostly have bifurcate or fimbriate apices (mostly with sharply pointed apices in C. heckrothi ) and the lateral stigmatic setae are usually <15 µm long (mostly> 15 µm in C. heckrothi ).

Two of five adult females of C. penangensis from Poring Hot Springs in Sabah, all mounted on one slide prepared by H.-P. Heckroth (slide also with 2 adult females of C. pseudotumuliferus ), have extremely few and very short (≤ 5 µm long) dorsal setae. The other three females are poor mounts but a few slightly longer dorsal setae are visible on two of them. Thus at least two of these females could erroneously key to the C. tumuliferus group or C. secretus because of these very short and sparse dorsal setae.

Coccus penangensis is a very widespread species, occurring in Borneo, Peninsular Malaysia, Singapore and Sumatra. Heckroth et al. (1998) recorded this species as common in both primary and secondary forest on Borneo (inside the stems of 17 Macaranga species) and Peninsular Malaysia plus Sumatra (in five Macaranga species). There is a large amount of morphological variation among adult females of C. penangensis from different localities, but there do not appear to be consistent differences related to geography. However, specimens from Burkill's collections on Penang Island appear to have longer dorsal setae than specimens from most other localities. Coccus penangensis was well represented (over 40 specimens) in the molecular phylogeny of Quek et al. (2017), with specimens from Bauk Hill on the east coast of Peninsular Malaysia and from near Siduk in West Kalimantan each forming a small cluster distinct from the remainder of the specimens. The adult females of these two groups did not appear to differ morphologically from specimens collected elsewhere but, unfortunately, the condition of molecular vouchers often makes detailed morphological comparisons difficult.

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 10. Adult female of Coccus penangensis Morrison. Abbreviations as in Fig. 3.