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Research Article
Review of the genus Coccus Linnaeus from Korea, with description of a new species (Hemiptera, Coccomorpha, Coccidae)
expand article infoJinyeong Choi, Seunghwan Lee
‡ Seoul National University, Seoul, Korea, South
† Deceased author
Open Access

Abstract

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.

Keywords

Coccinae , Coccini , soft scale insect, taxonomy

Introduction

The genus Coccus Linnaeus, 1758, which is a species-rich group in the family Coccidae, comprises approximately 111 species worldwide (Hodgson 1994; García-Morales et al. 2016). This group is defined morphologically by the distribution of ventral tubular ducts, the shape of dorsal and marginal setae, and the presence of a tibio-tarsal articulatory sclerosis on each leg (Hodgson 1994); however, the molecular phylogeny using some of the taxa included in Coccidae revealed that it is not a monophyletic group and needs taxonomic revision (Lin et al. 2013). Among the genus Coccus, C. hesperidum Linnaeus, C. viridis Green, and C. celatus De Lotto are known as economic pests of ornamental and agricultural products (Kapranas et al. 2007; Waller et al. 2007). Two species of the genus, C. hesperidum and C. pseudomagnoliarum (Kuwana) have been previously recorded from Korea. Here, a new species is described, Coccus ficicola sp. n. and its morphology is compared with congeners. A hitherto unrecorded species is also redescribed, C. malloti (Takahashi) and recorded for the first time from Korea.

Materials and methods

The specimens were mounted on microscope slides using the method of Hodgson and Henderson (2000) and Danzig and Gavrilov-Zimin (2014). The micrographs of slide-mounted materials were taken and measured using analysis software (Active Measure ver. 3.0.3, Mitani Co. Ltd, Japan). The terminology follows Hodgson (1994) and Hodgson and Henderson (2000), except that the term “pregenital disc-pores” is replaced with “multilocular pores” suggested by Kondo and Hardy (2008). The type specimens are deposited in the Insect Biosystematics Laboratory, Research Institute for Agriculture and Life Science, Seoul National University, Korea (SNU).

Taxonomy

Coccus Linnaeus, 1758: 455

Type species

Coccus hesperidum Linnaeus, 1758, designated by Opinion 1303 (1985).

Diagnosis

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 Hodgson (1994) and Wang and Feng (2012).

Key to species of genus Coccus in Korea

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.

Coccus ficicola sp. n.

Figs 1A–D, 2A–Q

Material examined

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♀♀.

Diagnosis

Adult females in life (Fig. 1A–D) with a reticulated pattern of brown stripes and a longitudinal ridge medially on dorsum; dermal areolations present but small; dorsal tubercles present; dorsal tubular ducts sparse on submarginal area; dorsal setae with bluntly rounded apices; marginal setae usually with simple pointed apices; multilocular disc-pores usually with ten loculi; ventral tubular ducts with a narrow inner ductule, frequent on posterior region of the head, medial area of thorax, and inner submarginal area of abdomen; antennae each with eight segments; legs each with a tibio-tarsal sclerosis on the articulation.

Figure 1. 

Coccus ficicola Choi & Lee sp. n. A instar female B immature adult female C mature adult female D mature (upper) and immature (under) adult females.

Description

Adult female. Living appearance (Fig. 1A–D). Body oval, flattened, or moderately convex. Young adult females yellowish to brownish, with a reticulated pattern of brown stripes except for a longitudinal ridge on mid dorsum. Older adult females becoming more convex and darker. Eggs not seen.

Slide-mounted material (Fig. 2A–Q). Body oval, 2.6–3.5 mm long, 2.0–3.6 mm wide, with distinct stigmatic cleft; anal clefts approximately 1/6 of body length.

Figure 2. 

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.

Etymology

Named after its host plant, Ficus benghalensis L.

Host plant

Moraceae: Ficus benghalensis L.

Comments

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 (Starr et al. 2003). In order to know the exact origin of the new species, further investigations are needed.

Morphological comparison of adult females of Coccus ficicola sp. n. and its related taxa

Based on taxonomic articles, such as Gill et al. (1977), Ben-Dov (1981), Avasthi and Shafee (1991), and Lin et al. (2017), we selected ten species morphologically similar to C. ficicola sp. n.: C. capparidis (Green, 1904), C. discrepans (Green, 1904), C. elatensis (Ben-Dov, 1981), C. formicarii (Green, 1896), C. gymnospori (Green, 1908), C. hesperidum (Linnaeus, 1758), C. latioperculatum (Green, 1922), C. moestus (De Lotto, 1959), C. praetermissus Lin & Tanaka, 2017, and C. sulawesicus Gavrilov, 2013. The morphological characters of adult females of Coccus ficicola and the ten species are summarized in Table 1.

Table 1.

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
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 Williams and Watson 1990
C. discrepans Present Absent Sharply pointed or bluntly rounded absent Pointed or frayed Seven segments Three pairs Present Present on medial area of mesothorax Avasthi and Shafee 1991; Tao et al. 1983
C. elatensis Present Present Bluntly rounded Present Pointed or frayed Eight segments Four pairs Present Present on medial area of mesothorax Ben-Dov 1981
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 Hodgson 1994
C. gymnospori Present Present Bluntly rounded Present Pointed or frayed Eight segments Three pairs Present Present on medial area of head and mesothorax Ben-Dov 1981
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 Hodgson 1994; Lin et al. 2017
C. latioperculatum Absent Absent Bluntly rounded Present Frayed Seven segments Two pairs Present Present on medial area of pro- and mesothorax Avasthi and Shafee 1991
C. moestus Present Present Bluntly rounded Present Frayed Seven or eight segments Three pairs Present Present on medial area of mesothorax Gill et al. 1977
C. praetermissus Present Present Bluntly rounded Present Pointed or frayed Seven segments Three pairs Present Present on medial area of mesothorax Lin et al. 2017
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 Gavrilov-Zimin 2013

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) (Ben-Dov 1981; Avasthi and Shafee 1989).

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 (De Lotto 1957; De Lotto 1960; Granara de Willink et al. 2010).

Coccus hesperidum (Linnaeus, 1758)

Fig. 3A–F

Coccus hesperidum Linnaeus, 1758: 455.

Material examined

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♀♀.

Diagnosis

Adult females in life (Fig. 3A–F) highly variable in body color and pigment pattern, but usually dorsum pale yellowish to brownish, with black or brown spots; dermal areolations present; dorsal tubercles present; dorsal tubular ducts present or absent; dorsal setae with sharply pointed apices; marginal setae usually with pointed, bifid or fimbriate apices; multilocular disc-pores usually with ten loculi; ventral tubular ducts with a narrow inner ductule, few present around meso- and procoxa, and anal plates; antennae each 7-segmented; legs each with a tibio-tarsal articulatory sclerosis.

Figure 3. 

Coccus hesperidum (Linnaeus, 1758). A population on Asplenium antiquum B adult female on Ficus benghalensis C adult female on Orchidaceae sp. D, E, F populations on Heteropanax fragrans.

Host plant. Recorded from 346 genera in 121 families (García-Morales et al. 2016). For Korean records, see Paik (1978).

Distribution

Known from all zoogeographical regions (García-Morales et al. 2016).

Coccus malloti (Takahashi, 1956)

Figs 4A–B, 5A–Q

Pulvinaria malloti Takahashi, 1956: 25.

Material examined

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♀.

Diagnosis

Adult females in life (Fig. 4A–B) with a reticulated pattern of black stripes and a longitudinal band medially on dorsum; dermal areolations present but small; dorsal tubercles present; dorsal tubular ducts absent; dorsal setae sharply spinose; marginal setae mostly with simple pointed apices; multilocular disc-pores usually with ten loculi; ventral tubular ducts of three types: Type I with a broad inner ductule, 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 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 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-, metathorax and anterior abdomen, and submarginal area of head; antennae each eight segments; legs each with a tibio-tarsal sclerosis on the articulation.

Figure 4. 

Coccus malloti (Takahashi, 1956). A adult female on Ilex cornuta B adult female on Aphananthe aspera.

Redescription

Adult female. Living appearance (Figs 4A–B). Body elongate oval, flattened, or slightly convex. Young adult females yellowish to dark brownish, with a reticulated pattern of brown or black stripes, getting darker at maturity. Eggs reddish in color, stored beneath venter.

Slide-mounted material (Fig. 5A–Q). Body elongate oval, 3.6–4.8 mm long, 2.2–3.3 mm wide, with shallow to deep stigmatic cleft; anal clefts approximately 1/6 of body length.

Figure 5. 

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.

Host plant

Recorded from six genera in six families (García-Morales et al. 2016). In Korea, it was found on Aphananthe aspera (Cannabaceae) and Ilex cornuta (Aquifoliaceae).

Distribution

Only known from Japan (Takahashi 1956); first record for Korea.

Comments

The above description based on Korean specimens agrees well with that of Takahashi (1956), except that variation in the number of dorsal tubercles and marginal setae, and exact distributions of each type of ventral tubular ducts are newly provided in this study. Coccus malloti probably has intermediate morphological characters between the tribes Coccini and Pulvinariini. However, the woolly test, known as ovisac and one of the typical characters of the Pulvinariini, is not observed in the species. In addition, some slide-mounted specimens of C. malloti contain eggs and nymphs in their body, which indirectly indicates that they would not produce an ovisac for oviposition. Although Coccus malloti would be retained in the tribe Coccini, it needs to be reviewed with its type materials to clarify the exact generic position of the species.

Coccus pseudomagnoliarum (Kuwana, 1914)

Fig. 6A–B

Lecanium (Eulecanium) pseudomagnoliarum Kuwana, 1914: 7.

Material examined

Adult female: Sujeong-dong, Yeosu-si, Jeollanam-do, 27.v.2015, coll. J.Y. Choi, on Celtis sp. (Cannabaceae), 10♀♀.

Diagnosis

Adult females in life (Fig. 6A–B) greenish or greyish, with light or dark yellow mottling; dermal areolations present; dorsal tubercles absent; dorsal tubular ducts absent; dorsal setae with sharply pointed apices; marginal setae with simple pointed or spatulate apices; multilocular disc-pores usually with 6–10 loculi; ventral tubular ducts with a narrow inner ductule, few present on submaginal area of posterior abdomen; antennae each 8-segmented; legs without tibio-tarsal articulatory scleroses.

Figure 6. 

Coccus pseudomagnoliarum (Kuwana, 1914). A immature adult females B mature adult female and 1st instars.

Host plant

Recorded from six genera in five families (García-Morales et al. 2016). In Korea, it has been recorded from the following plants: Celtis willdenowiana (Cannabaceae), Citrus sp., Phellodendron amurense, Poncirus trifoliata (Rutaceae), Clerodendron trichotomum (Lamiaceae), and Zelkova serrata (Ulmaceae) (Paik 1978).

Distribution

Mainly known from Palearctic Region including Australia, Europe, Iran, Israel, Japan, Russia, Korea, and USA (García-Morales et al. 2016).

Discussion

Recently, Lin et al. (2017) described Coccus praetermissus Lin & Tanaka, which could be confused with a cosmopolitan species, Coccus hesperidum Linnaeus, 1758, based on morphological and molecular evidences. They pointed out that a morphological difference exists between the adult females of two genetically distinct species, the shape of dorsal setae, although molecular data (COI) should be used for exact identification. The adult female of Coccus praetermissus has dorsal setae with bluntly rounded apices, whereas those of C. hesperidum have sharply pointed tips. Coccus ficicola sp. n. is close to C. praetermissus in having the former type of dorsal setae, but differs in the distributional pattern of ventral tubular ducts, which is a reliable and constant character in each species within the genus Coccus. The ventral tubular ducts of Coccus ficicola are present on medial area of head, pro- and mesothorax, and submarginal area of abdomen, whereas C. praetermissus has the structures on medial area of mesothorax only.

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.

Acknowledgments

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).

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