Research Article |
Corresponding author: Seunghwan Lee ( seung@snu.ac.kr ) Academic editor: Roger Blackman
© 2018 Jinyeong Choi, Seunghwan Lee.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Choi J, Lee S (2018) Review of the genus Coccus Linnaeus from Korea, with description of a new species (Hemiptera, Coccomorpha, Coccidae). ZooKeys 734: 121-135. https://doi.org/10.3897/zookeys.734.22774
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The genus Coccus from Korea is reviewed, including a new species, C. ficicola sp. n., and a first record of C. malloti (Takahashi, 1956). The new species is characterized by a distinctive pattern of ventral tubular ducts on medial area of head and thorax, and submarginal area of abdomen. The adult female of Coccus ficicola sp. n. is described and illustrated, and a morphological comparison is given with congeners. Coccus malloti is redescribed and illustrated based on the adult female specimens from Korea. A key to the four species of Coccus known from Korea is provided with diagnoses and photographs.
Coccinae , Coccini , soft scale insect, taxonomy
The genus Coccus Linnaeus, 1758, which is a species-rich group in the family Coccidae, comprises approximately 111 species worldwide (
The specimens were mounted on microscope slides using the method of
Coccus hesperidum Linnaeus, 1758, designated by Opinion 1303 (1985).
Dorsal setae pointed or blunt; dorsal tubular ducts and dorsal tubercles present or absent; marginal setae with pointed or frayed apices; ventral tubular ducts present or absent, if present, mainly distributed on medial area of thorax or submarginal area; a tibio-tarsal articulatory sclerosis present or absent on each leg. For further diagnostic characteristics, see
1 | Dorsal tubercles absent; legs without tibio-tarsal articulatory scleroses; ventral tubular ducts present on abdomen only | C. pseudomagnoliarum (Kuwana) |
– | Dorsal tubercles present; legs with tibio-tarsal articulatory scleroses; ventral tubular ducts present on thorax and abdomen | 2 |
2 | Ventral tubular ducts of three types (Type I: each with a broad inner ductule; Type II: each with a narrow inner ductule; Type III: each with a filamentous inner ductule) present | C. malloti (Takahashi) |
– | Ventral tubular ducts of type I, each with a narrow inner ductule | 3 |
3 | Antenna 7-segmented; ventral tubular ducts scarce: a small group of 0–3 ducts present between mouthparts and each procoxa; a thin transverse band containing one or two ducts vertically present between mesocoxae; absent on inner submarginal area of abdomen | C. hesperidum (Linnaeus) |
– | Antenna 8-segmented; ventral tubular ducts abundant: a large group of 16–20 ducts present between mouthparts and each procoxa; a broad transverse band containing 4–7 ducts present vertically between mesocoxae; present on inner submarginal area of abdomen | C. ficicola sp. n. |
Holotype: adult female: Korea, Gangnam-gu, Yeoksam-dong, 18.iv.2015, coll. J.Y. Choi, on Ficus benghalensis L. (Moraceae). Paratypes: same data as holotype, 9♀♀.
Adult females in life (Fig.
Adult female. Living appearance (Fig.
Slide-mounted material (Fig.
Coccus ficicola sp. n., adult female. A dorsal microduct B dorsal tubercle C dermal areolations D dorsal seta E stigmatic spines F marginal setae G dorsal tubular duct H anal plates I preopercular pore J multilocular disc-pores K ventral tubular duct on abdomen L leg M ventral seta N ventral tubular duct on head and thorax. O spiracular pore P ventral microduct Q antenna. Scale bars: 200 μm (L, Q); 100 μm (H); 50μm (C, E); others = 10μm.
Dorsum. Derm membranous. Dermal areolations oval and small, each with a microduct. Dorsal tubercles normally convex, present on submarginal area, 4–6 in total on each side: two between apex of head and anterior stigmatic cleft, one or two between anterior and posterior stigmatic clefts, and one or two between posterior stigmatic cleft and anal cleft. Dorsal setae cylindrical, short, stout, blunt apically, each 6–9 μm long, moderately distributed on dorsum. Dorsal tubular ducts each with a developed outer ductule and a slender inner ductule with a developed terminal gland, sparsely present on submargin. Dorsal microducts evenly scattered over entire dorsum. Preopercular pores round and small, rather inconspicuous, 6–7 μm wide, set in a small group of approximately 6–15 in front of anal plates. Anal plates quadrate, 190–223 μm long, 160–203 μm wide, usually posterolateral margin slightly longer than anterolateral margin; anterolateral margin 119–144 μm long, posterolateral margin 130–154 μm long; each plate with four apical setae. Ano-genital fold with two pairs of anterior and three pairs of lateral margin setae. Anal ring with six long setae.
Margin. Marginal setae spinose, slender, slightly bent, each 16–32 μm long, mostly with simple pointed apices, but sometimes with bifid tips; with 52–59 present between anterior stigmatic clefts; 12–17 laterally present between anterior and posterior stigmatic clefts, 45–53 present between posterior stigmatic cleft and anal clefts. Stigmatic clefts deep, each with three stigmatic spines, median spine 2–3 times as long as lateral spine: medians 48–77 μm long, laterals 14–30 μm long. Eyespots located near margin.
Venter. Derm membranous. Multilocular disc-pores 7–8 μm wide, each with 10–12 loculi, mostly with ten loculi, abundant around vulvar area, but less frequent on anterior segments of abdomen. Spiracular pores 4–5 μm wide, each with five loculi, in a narrow band 1–2 pores wide between each spiracle and stigmatic cleft. Ventral tubular ducts of a single type, each 21–28 μm long, with a developed outer ductule and a narrow inner ductule with a flower-shaped terminal gland, approximately 16–20 ducts densely present between mouthparts and procoxa on each side; a broad transverse band containing around 4–7 ducts vertically present between mesocoxae; abundant between each meso- and metacoxa, extending around spiracles; and also sparsely scattered on inner submarginal area between anal plates and each metacoxa. Ventral microducts present on entire venter, especially frequent on submargin. Ventral setae with three pairs of long pregenital setae; two pairs of long setae between antennae; other setae sharply spinose, each 7–14 μm long, moderately distributed over entire venter. Legs well developed, each with a tibio-tarsal articulation and an articulatory sclerosis; total length of each metathoracic leg 560–638 μm long: each coxa 138–163 μm long, trochanter+femur 181–213 μm long, tibia+tarsus 218–237 μm long, claw 17–26 μm long. Tarsal digitules thinner and longer than claw digitules. Spiracles normal, mostly posterior peritreme broader than anterior: anterior peritremes each 38–49 μm wide, posterior peritremes each 45–57 μm wide. Antenna 8-segmented, each 279–339 μm long. Clypeolabral shield 127–138 μm wide.
Named after its host plant, Ficus benghalensis L.
Moraceae: Ficus benghalensis L.
Coccus ficicola sp. n. is probably a non-endemic species because it occurs on an imported ornamental plant, Ficus benghalensis, which is widely cultivated in tropical areas (
Based on taxonomic articles, such as
Comparison of morphological characters of adult females of Coccus ficicola sp. n. and its related taxa.
Species | Dorsal tubercles | Dorsal tubular ducts | Dorsal setae | Preopercular pores | Marginal setae | Antenna | Pregenital setae | Tibio-tarsal sclerosis | Ventral tubular ducts | Reference |
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C. ficicola sp. n. | Present | Present | Bluntly rounded | Present | Pointed or frayed | Eight segments | Three pairs | Present | Present on medial area of head, pro-, and mesothorax; submarginal area of abdomen | This study |
C. capparidis | Present | Absent | Bluntly rounded | Present | Pointed or frayed | Six or seven segments | One or two pairs | Absent | Present on submarginal area of abdomen |
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C. discrepans | Present | Absent | Sharply pointed or bluntly rounded | absent | Pointed or frayed | Seven segments | Three pairs | Present | Present on medial area of mesothorax |
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C. elatensis | Present | Present | Bluntly rounded | Present | Pointed or frayed | Eight segments | Four pairs | Present | Present on medial area of mesothorax |
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C. formicarii | Absent | Absent | Sharply pointed (setose) | Present | Pointed (setose) | Seven or eight segments | Three pairs | Absent | Present on medial area of head, pro-, and mesothorax |
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C. gymnospori | Present | Present | Bluntly rounded | Present | Pointed or frayed | Eight segments | Three pairs | Present | Present on medial area of head and mesothorax |
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C. hesperidum | Present | Present or absent |
Sharply pointed | Present | Pointed or frayed | Seven segments | Three pairs | Present | Present on medial area of pro- and mesothorax; laterad to genital opening |
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C. latioperculatum | Absent | Absent | Bluntly rounded | Present | Frayed | Seven segments | Two pairs | Present | Present on medial area of pro- and mesothorax |
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C. moestus | Present | Present | Bluntly rounded | Present | Frayed | Seven or eight segments | Three pairs | Present | Present on medial area of mesothorax |
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C. praetermissus | Present | Present | Bluntly rounded | Present | Pointed or frayed | Seven segments | Three pairs | Present | Present on medial area of mesothorax |
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C. sulawesicus | Absent | Absent | Sharply pointed | Present | Pointed or frayed | Seven or eight segments | Three pairs | Present | Present on medial area of meso- and metathorax |
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In the morphological comparison, Coccus ficicola shows a new combination of morphological characters; in particular, the distributional pattern of ventral tubular ducts of the species reveals uniqueness among the nine morphological characters. Coccus ficicola is most closely related to C. gymnospori (Green), in having (i) dorsal tubercles, (ii) dorsal tubular ducts on submarginal area, (iii) dorsal setae with bluntly rounded apices, (iv) preopercular pores, (v) marginal setae with pointed or frayed apices, (vi) antenna with eight segments, (vii) three pairs of pregenital setae, and (viii) tibio-tarsal sclerosis. However, C. ficicola differs from C. gymnospori in having the following combination of character states (character states of C. gymnospori in parenthesis): (i) ventral tubular ducts abundant, 16 to 20 ducts present between mouthparts and each procoxa (few, only 3 or 4 ducts); a broad transverse band containing 4–7 ducts vertically between metacoxae (thin, containing one or two ducts); and present on inner submarginal area of abdomen (entirely absent), and (ii) multilocular disc-pores extending further anteriorly (restricted to preceding two abdominal segments) (
Although the African species, C. africanus (Newstead) and C. alpinus De Lotto, are not included in the list of related taxa for morphological comparison, C. ficicola is similar to both species in having abundant ventral tubular ducts. However, C. ficicola does not have continuous ventral tubular ducts between the metacoxae, whereas both African species have this character state (
Coccus hesperidum Linnaeus, 1758: 455.
Adult female: Daehak-dong, Gwanak-gu, Seoul, 09.iv.2014, coll. J.Y. Choi, on Orchidaceae sp., 5♀♀; Sinhyo-dong, Seogwipo-si, Jeju-do, 14.ix.2014, coll. J.Y. Choi, on Asplenium antiquum Makino (Aspleniaceae), 5♀♀; Sinbuk-eup, Chuncheon-si, Gangwon-do, 31.v.2015, coll. J.Y. Choi, on Heteropanax fragrans (Roxb.) (Araliaceae), 5♀♀; Geumam-dong, Deokjin-gu, Jeonju-si, Jeollabuk-do, 06.vi.2015, on same host, 5♀♀; Songhyeon-dong, Andong-si, Gyeongsangbuk-do, 07.vi.2015, on same host, 5♀♀; Guseo-dong, Geumjeong-gu, Busan, 07.vi.2015, on Ficus benghalensis L., 5♀♀.
Adult females in life (Fig.
Host plant. Recorded from 346 genera in 121 families (
Known from all zoogeographical regions (
Pulvinaria malloti Takahashi, 1956: 25.
Adult female: Jeollanam-do, Gwangyang-si, Ongnyong-myeon, Chusan-ri, 28.v.2015, coll. J.Y. Choi, on Ilex cornuta Lindl. (Aquifoliaceae), 9♀♀; Jeju-do, Seogwipo-si, Andeok-myeon, Gamsan-ri, 27.iv.2016, coll. J.Y. Choi, on Aphananthe aspera (Thunb.) (Cannabaceae), 1♀.
Adult females in life (Fig.
Adult female. Living appearance (Figs
Slide-mounted material (Fig.
Coccus malloti (Takahashi, 1956), adult female. A dorsal microduct B dorsal tubercle C dermal areolations D dorsal seta E stigmatic spines F marginal seta G anal plates H preopercular pore I multilocular disc-pores J ventral tubular duct (Type II) K leg L ventral tubular duct (Type III) M ventral tubular ducts (Type I) N ventral seta O spiracular pore P ventral microduct Q antenna. Scale bars: 200 μm (K, Q); 100μm (G); 50 μm (C, E); others = 10 μm.
Dorsum. Derm membranous. Dermal areolations oval and small, each with a microduct. Dorsal tubercles normally convex, present on submarginal area, 1–5 in total on each side: one or two between apex of head and anterior stigmatic cleft, zero to two between anterior and posterior stigmatic clefts, and zero or one between posterior stigmatic cleft and anal cleft. Dorsal setae sharply spinose, short, stout, each 6–9 μm long, moderately distributed on dorsum. Dorsal tubular ducts absent. Dorsal microducts evenly scattered over entire dorsum. Preopercular pores round and small, 3–5 μm wide, set in a small group of approximately 8 in front of anal plates. Anal plates quadrate, 217–249 μm long, 169–198 μm wide, each with slightly concaved posterolateral margin, usually posterolateral margin quite longer than anterolateral margin; anterolateral margin 123–143 μm long, posterolateral margin 150–169 μm long; each plate with four apical setae. Ano-genital fold with two pairs of anterior and three pairs of lateral margin setae. Anal ring with six long setae.
Margin. Marginal setae spinose, straight or slightly bent, each 14–22 μm long, mostly with simple pointed apices; with 30–43 present between anterior stigmatic clefts; 13–17 laterally present between anterior and posterior stigmatic clefts, 22–38 present between posterior stigmatic cleft and anal clefts. Stigmatic clefts shallow to deep, each with three stigmatic spines, median spine nearly twice as long as lateral spine: medians 60–75μm long, laterals 24–38 μm long. Eyespots located near margin.
Venter. Derm membranous. Multilocular disc-pores 6–7 μm wide, each with 8–10 loculi, mostly with ten loculi, abundant around vulvar area; one or two transverse rows on each abdominal segments; and also small groups present laterad of each metacoxa and mesocoxa, but not observed around procoxa. Spiracular pores 4–5 μm wide, each with five loculi, in a narrow band 2–4 pores wide between each spiracle and stigmatic cleft. Ventral tubular ducts of three types: Type I each with 33–39 μm long, with a developed outer ductule and a moderately broad inner ductule, straight or slightly curved, with a flower-shaped terminal gland, densely present on posterior medial area of head; frequent on anterior medial area of prothorax, extending to inner submarginal area of thorax; and also sparsely scattered on inner submarginal area of abdomen; Type II each with 16–24 μm long, with a narrow inner ductule and a large flower-shaped terminal gland, rarely present on inner submarginal area and posterior medial area of abdomen; Type III each with 15–24 μm long, with a long filamentous inner ductule and a quite small terminal gland, moderately present on submarginal area between anal clefts and each posterior spiracular furrow; all types of ventral tubular ducts absent on medial area of meso- and metathorax and anterior abdomen, and submarginal area of head. Ventral microducts present on entire venter, especially frequent on submargin. Ventral setae with three pairs of long pregenital setae; approximately three or four pairs of long and short setae between antennae; other setae sharply spinose, each 5–10 μm long, sparsely distributed over entire venter. Legs well developed, each with a tibio-tarsal articulation and an articulatory sclerosis; total length of each metathoracic leg 614–769 μm long: each coxa 141–189 μm long, trochanter+femer 209–257 μm long, tibia+tarsus 243–311 μm long, claw 17–24 μm long. Tarsal digitules thinner and longer than claw digitules. Spiracles normal, mostly posterior peritreme broader than anterior: anterior peritremes each 37–52 μm wide, posterior peritremes each 45–61 μm wide. Antenna 8-segmented, each 339–399 μm long. Clypeolabral shield 138–157 μm wide.
Recorded from six genera in six families (
Only known from Japan (
The above description based on Korean specimens agrees well with that of
Lecanium (Eulecanium) pseudomagnoliarum Kuwana, 1914: 7.
Adult female: Sujeong-dong, Yeosu-si, Jeollanam-do, 27.v.2015, coll. J.Y. Choi, on Celtis sp. (Cannabaceae), 10♀♀.
Adult females in life (Fig.
Recorded from six genera in five families (
Mainly known from Palearctic Region including Australia, Europe, Iran, Israel, Japan, Russia, Korea, and USA (
Recently,
Under the morphological comparison with congeners, we conclude that Coccus ficicola sp. n. is a distinct species which is a morphologically differentiated lineage. The distinctive pattern of ventral tubular ducts seems to be an autapomorphic feature of Coccus ficicola because it shows uniqueness in the comparison of morphological characters. In order to clarify the phylogenetic relationships of a new species within the genus Coccus, molecular analysis employing mitochondrial and nuclear loci are required.
We thank Dr. Sora Kim for reviewing the manuscript for this study. We are also grateful for a colleague, Mr. Seunghyun Lee (Department of Agricultural Biotechnology, Seoul National University, Seoul, Korea), who provided specimens. We also thank Dr. Chris J. Hodgson (Department of Biodiversity and Biological Systematics, The National Museum of Wales, Cardiff, UK), Dr. T. Kondo (Corporación Colombiana de Investigación Agropecuaria, Corpoica, Colombia) and Dr. Yen P. Lin (School of Biological Sciences, The University of Queensland, Brisbane, Australia) for the useful comments. In addition, we thank Dr. H. Tanaka (Ehime University, Ehime, Japan) for comparing our samples with Japanese samples.
The research on the species newly recorded in Korea was supported by a grant from the National Institute of Biological Resources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR201701203).