Abstract
Background
China is one of the countries with the greatest species diversity of Chilocorini (Coleoptera: Coccinellidae), including nearly forty-five percent of the known genera and fourteen percent of all described species in this tribe. Recently, we discovered three species previously not recorded in China.
New information
In this study, three species Priscibrumus uropygialis (Mulsant, 1853), Priscibrumus disjunctus Canepari, 1997 and Brumus octosignatus (Gebler, 1830) are documented for the first time in China. Brumus octosignatus is the first member of the genus Brumus Mulsant, 1850 recorded in China. Detailed descriptions, illustrations and distributions of these three species are provided. A checklist of Chinese Chilocorini is also given.
Keywords: Coleoptera , Coccinellidae , Chilocorini , new record, checklist, China
Introduction
The members of family Coccinellidae, commonly known as colourful and shiny beetles, predators of plant pests, contain 6000 species distributed worldwide (Vandenberg 2002). The tribe Chilocorini is well-known as a primary predator of coccids. Many species of this tribe are widely used as biological control agents. In recent years, phylogenetic and evolutionary studies of Chilocorini indicated that this tribe is a monophyletic group, closely related to the tribe Coccinellini (Magro et al. 2010, Seago et al. 2011, Escalona et al. 2017, Che et al. 2017) or Plotinini (Li et al. 2020). At present, Chilocorini contains 22 genera and more than 280 species distributed worldwide (Łączyński and Tomaszewska 2012, Li et al. 2020).
China is one of the countries with the greatest species of Chilocorini (Coleoptera: Coccinellidae), including 10 genera and 39 species (Li et al. 2017a). However, in Li et al. (2017a), the genus Brumus Mulsant was incorrectly recorded in the Chinese checklist of Chilocorini (no literature existed to indicate that the members of this genus were distributed in China). After a phylogenetic study of the Chilocorini, the genus Phaenochilus Weise was synonymised with Chilocorus Leach (Li et al. 2020).
In this study, we report for the first time in China the genus Brumus with the species Brumus octosignatus (Gebler, 1830), as well as the species Priscibrumus uropygialis (Mulsant, 1853) and Priscibrumus disjunctus Canepari, 1997. A revised checklist of Chinese Chilocorini is also provided, containing all nine genera and 42 species.
Materials and methods
Specimens, examined in this study, were collected in China (Tibet and Xinjiang) and deposited at the Department of Entomology, South China Agriculture University (SCAU), Guangzhou.
The newly-collected specimens of Priscibrumus disjunctus were identified based on the original species description (Canepari 1997). The specimens of the other two new Chinese records i.e. Priscibrumus uropygialis and Brumus octosignatus, were identified from the secondary descriptions and illustrations by Miyatake (1985) and Kovář (1997), respectively.
External morphology was observed with a dissecting stereomicroscope (SteREO Discovery V20, Zeiss). Male and female genitalia were dissected, cleared in 10% solution of sodium hydroxide (NaOH) by boiling for several minutes and examined with an Olympus BX51 microscope. Photographs of the genitalia and other morphological characters were taken with digital cameras (AxioCam HRc and Coolsnap-Procf & CRI Micro*Color), attached to microscopes using AxioVision Rel. ver. 4.8 and Image-Pro Plus ver. 6.0. Images were cleaned up and laid out in plates with Adobe Photoshop CS ver. 8.0. Terminology follows Ślipiński (2007).
Abbreviations
TL = total length: length from apical margin of clypeus to apex of elytra
TW = total width: width across both elytra at widest part
TH = body height measured across the highest point of the elytra
HW = head width in a frontal view
PL = pronotal length: from middle of anterior margin to base of pronotum
PW = pronotal width at widest part
EL = elytral length: from apex to base including scutellum
EW = elytral width, equal to TW
Taxon treatments
Priscibrumus
Kovář, 1997
878E2E1D-DEEB-5C59-8608-647643362168
Priscibrumus Kovář, 1997 in Kovář 1997: 114
Priscibrumus Exochomus puniceipennis Semenov, 1900
Diagnosis
Priscibrumus can be distinguished from other genera of the tribe Chilocorini by the following combination of characters: body densely covered with short, greyish pubescence; antenna composed of 10 antennomeres, with terminal antennomere very small and embedded in the penultimate antennomere; pronotal basal margin entirely bordered with submarginal line; base of pronotum and elytra contiguous all along their length; elytral epipleura narrow, more or less horizontal and without foveae; abdominal postcoxal lines almost complete; mid and hind tibiae with two apical spurs.
Priscibrumus disjunctus
Canepari, 1997
DBFEBD22-DF86-5F7D-9C90-20724315E3A3
Priscibrumus disjunctus Canepari, 1997 in Canepari 1997: 45.
Materials
Type status: Holotype. Occurrence: recordedBy: Wenjing Li; individualCount: 2; sex: 1 male, 1 female; lifeStage: adult; Taxon: scientificName: Priscibrumus disjunctus; Location: country: China; stateProvince: Tibet; locality: Jilong County; verbatimElevation: 2900 m; verbatimCoordinates: 28°18.05'N; 85°23.12'E; georeferenceProtocol: label; Identification: identifiedBy: Wenjing Li; dateIdentified: 2017; Event: samplingProtocol: sweeping; eventDate: 28/08/1984; Record Level: basisOfRecord: PreservedSpecimen
Description
TL: 3.75–4.10 mm, TW: 2.80–2.91 mm, TH: 1.83–2.12 mm, TL/TW: 1.31–1.34, PL/PW: 0.45–0.47, EL/EW: 1.07–1.10.
Body oval, moderately convex. Head black, densely covered with short, greyish pubescence. Mouthparts and antenna black. Pronotum black, densely covered with short, greyish pubescence. Scutellum black. Elytra reddish-brown, with two pairs of broadly black stripes: outer stripes approximately 2/5 width of elytra, 3/4 length of elytra; inner stripes situated on suture, almost as long as elytra, distinctly broadening at base and weakly broadening at apex, densely covered with extremely short, greyish pubescence (Fig. 1a–c). Underside black except elytral epipleura brownish-yellow, densely covered with short, greyish pubescence. Abdominal postcoxal lines incomplete, arcuate. Posterior margin of male abdominal ventrite 5 slightly emarginate at middle and of ventrite 6 distinctly emarginate (Fig. 1d).
Figure 1.
Priscibrumus disjunctus Canepari, 1997. a. dorsal view; b. lateral view; c. frontal view; d. abdomen; e. penis; f. apex of penis; g. tegmen, lateral view; h. tegmen, ventral view. Scale bars: 0.1 mm.
Male genitalia: penis slender, penis capsule with short outer and inner arm, apex of penis truncate with membranous appendage (Fig. 1e–f). Tegmen stout, penis guide narrow at base, parallel along basal 1/3, after that, gradually broadening to basal 2/3, then narrowing to apex in ventral view; penis guide in lateral view, widest near base, gradually narrowing to apex; parameres distinctly longer than penis guide with dense short setae on the inner sides and distal end with a group of short setae in lateral view (Fig. 1g–h).
Female genitalia: coxites distinctly elongated. Spermatheca approximately C-shaped, cornu without appendage.
Diagnosis
This species can be distinguished from other species of Priscibrumus by the following combination of characters: elytra reddish-brown, with two pair of broadly black stripes, inner stripes situated on suture, almost as long as elytra; parameres distinctly longer than penis guide.
Distribution
Nepal (Canepari 1997) and Tibet, China (present study) (Fig. 4).
Figure 4.
Distribution map. (▲) Priscibrumus disjunctus Canepari, 1997; (●) Priscibrumus uropygialis (Mulsant, 1853); (○) Brumus octosignatus (Gebler, 1830).
Priscibrumus uropygialis
(Mulsant, 1853)
6F63801E-90E2-5C22-AD8C-2E493A7F22A7
Exochomus uropygialis Mulsant, 1853 in Mulsant 1853: 196.
Brumus uropygialis (Mulsant, 1853): Crotch, 1874 (see Crotch 1874: 196, Gordon 1985: 25).
Exochomus (Exochomus) uropygialis (Mulsant, 1853) Barovskij, 1922 (see Barovskij 1922: 297-8; Mader 1955: 789, 797-8; Bielawski 1979: 114; Miyatake 1985: 11).
Priscibrumus uropygialis (Mulsant, 1853): Kovář, 1997 (see Kovář 1997: 117; Canepari 1997: 45; Poorani 2002: 314).
Materials
Type status: Other material. Occurrence: recordedBy: Wenjing Li; individualCount: 2; sex: 1 male, 1 female; lifeStage: adult; Taxon: scientificName: Priscibrumus uropygialis; Location: country: China; stateProvince: Tibet; locality: Mama village, Cuona County; verbatimElevation: 2800 m; verbatimCoordinates: 27°53.03'N; 91°47.34'E; georeferenceProtocol: label; Identification: identifiedBy: Wenjing Li; dateIdentified: 2017; Event: samplingProtocol: sweeping; eventDate: 25/05/2011; Record Level: basisOfRecord: PreservedSpecimen
Type status: Other material. Occurrence: recordedBy: Wenjing Li; individualCount: 1; sex: 1 male; lifeStage: adult; Taxon: scientificName: Priscibrumus uropygialis; Location: country: China; stateProvince: Tibet; locality: Jilong County; verbatimElevation: 2800 m; verbatimCoordinates: 28°18.05'N; 85°23.12'E; georeferenceProtocol: label; Identification: identifiedBy: Wenjing Li; dateIdentified: 2017; Event: samplingProtocol: sweeping; eventDate: 29/05/2011; Record Level: basisOfRecord: PreservedSpecimen
Description
TL: 3.65–4.24 mm, TW: 2.72–3.31 mm, TH: 1.46–1.82 mm, TL/TW: 1.28–1.34, PL/PW: 0.46–0.51, EL/EW: 1.07–1.16.
Body oval, moderately convex. Head black, densely covered with short, greyish pubescence. Mouthparts and antenna black. Pronotum black, densely covered with short, greyish pubescence. Scutellum black. Elytra reddish-brown, with a pair of black spots at elytral apex, densely covered with short, greyish pubescence (Fig. 2a–c). Underside black, except elytral epipleura brownish-yellow, densely covered with short, greyish pubescence. Abdominal postcoxal lines incomplete, arcuate. Posterior margin of male abdominal ventrite 5 slightly emarginate at middle and of ventrite 6 distinctly emarginate (Fig. 2d).
Figure 2.
Priscibrumus uropygialis (Mulsant, 1853). a. dorsal view; b. lateral view; c. frontal view; d. abdomen; e. penis; f. apex of penis; g. tegmen, lateral view; h. tegmen, ventral view; i. ovipositor. Scale bars: 0.1 mm.
Male genitalia: penis slender, penis capsule with short outer arm and long inner arm, apex of penis truncate with membranous appendage (Fig. 2e–f). Tegmen stout, penis guide parallel along 2/3 length, then gradually narrowing to apex in ventral view; penis guide in lateral view, widest at base, parallel along 2/5 length, then gradually narrowing to apex; parameres as long as penis guide with dense long setae on the inner sides and distal end with a group of long setae in lateral view (Fig. 2g–h).
Female genitalia: coxites distinctly elongate (Fig. 2i). Spermatheca approximately C-shaped, cornu without appendage.
Diagnosis
This species can be easily distinguished from other species of Priscibrumus by the following combination of characters: elytra reddish-brown, with a pair of black spots at elytral apex; parameres as long as penis guide.
Distribution
Kashmir, Nepal, India, Bhutan, Pakistan (Mulsant 1850, Crotch 1874, Gordon 1987, Barovskij 1922, Bielawski 1979, Miyatake 1985, Kovář 1997, Canepari 1997, Poorani 2002, Mader 1955) and Tibet, China (present study) (Fig. 4).
Brumus
Mulsant, 1850
49BD6CF2-085C-5C68-A9A6-62CC9C552965
Brumus Mulsant, 1850 in Mulsant 1850: 492.
Brumus Coccinella octosignata Gebler, 1830
Diagnosis
Brumus can be distinguished from other genera of the tribe Chilocorini by the following combination of characters: antenna 10-segmented, terminal antennomere very small and embedded in penultimate segment; pronotal basal margin entirely bordered with submarginal line; elytral epipleura narrow, more or less horizontal and without foveae; abdominal postcoxal lines complete; mid and hind tibiae with two apical spurs; tarsal claws without basal tooth, only slightly swollen at base.
Brumus octosignatus
(Gebler, 1830)
E53379AA-1E43-5A00-BFDA-D1E4683C8933
Coccinella octosignata Gebler, 1830 in Gebler 1830: 225.
Coccinella deserta Motschulsky, 1840 in Motschulsky 1840: 175.
Brumus desertorum Mulsant, 1850 in Mulsant 1850: 493.
Brumus 8-signata (Gebler, 1830) Crotch, 1874 see Crotch 1874: 38.
Brumus octosignatus (Gebler, 1830) Crotch, 1874, see Crotch 1874: 195, Barovskij 1927: 196, 199; Korschefsky 1932: 199; Mader 1955: 803, 805; Bielawski 1975: 255; Bielawski 1984: 315; Kovář 1997: 36.
Materials
Type status: Other material. Occurrence: recordedBy: Wenjing Li; individualCount: 2; sex: 1 male, 1 female; lifeStage: adult; Taxon: scientificName: Brumus octosignatus (Gebler, 1830); Location: country: China; stateProvince: Xinjiang; locality: Kezier village, Baicheng County; verbatimElevation: 850 m; verbatimCoordinates: 43°27.23'N; 82°27.40'E; georeferenceProtocol: label; Identification: identifiedBy: Wenjing Li; dateIdentified: 2017; Event: samplingProtocol: sweeping; eventDate: 31/07/1995; Record Level: basisOfRecord: PreservedSpecimen
Type status: Other material. Occurrence: recordedBy: Wenjing Li; individualCount: 4; sex: 2 male, 2 female; lifeStage: adult; Taxon: scientificName: Brumus octosignatus (Gebler, 1830); Location: country: unknown; stateProvince: unknown; locality: unknown; verbatimElevation: unknown; verbatimCoordinates: unknown; Identification: identifiedBy: Wenjing Li; dateIdentified: 2017; Record Level: basisOfRecord: PreservedSpecimen
Description
TL: 3.80–4.00 mm, TW: 2.80–3.20 mm, TH: 1.87–2.07 mm, TL/TW: 1.32–1.35, PL/PW: 0.47–0.50, EL/EW: 1.05–1.10.
Body oval, moderately convex. Head, mouthparts and antenna brownish-yellow. Pronotum orange-yellow, with a black spot at centre of basal margin. Scutellum black. Elytra orange-yellow, with four pairs of black spots, the first one situated at the humeral angle; the second one situated at basal 2/5, near suture; the third one situated at basal 3/5, near outer margin; the fourth one situated basal 4/5, near suture (Fig. 3a–c). Underside orange-yellow, densely covered with short, greyish pubescence. Abdominal postcoxal lines complete and semicircular. Posterior margin of male abdominal ventrite 5 and of ventrite 6 distinctly emarginate (Fig. 3d).
Figure 3.
Brumus octosignatus (Gebler, 1830). a. dorsal view; b. lateral view; c. frontal view; d. abdomen; e. penis; f. apex of penis; g. tegmen, lateral view; h. tegmen, ventral view; i. ovipositor. Scale bars: 0.1 mm.
Male genitalia: penis slender, penis capsule with short outer and long inner arm, apex of penis acute with membranous appendage (Fig. 3e–f). Tegmen stout, penis guide narrow at base, widest at basal 1/2, then gradually converging to blunt tip, symmetrical in ventral view; penis guide in lateral view, widest at base, parallel along basal 1/3, after that gradually converging to apex; parameres nearly as long as the penis guide with dense short setae on the inner sides and distal end with a group of short setae in lateral view (Fig. 3g–h).
Female genitalia: coxites distinctly elongate (Fig. 3i). Spermatheca approximately C-shaped, cornu without appendage.
Diagnosis
This species can be easily distinguished from other species of Brumus by the following combination of characters: pronotum with a black spot at central of basal margin; elytra orange-yellow, with four pairs of black spots; parameres nearly as long as the penis guide.
Distribution
Azerbaijan, Armenia, France, Italy, Russia: south European Territory, Afghanistan, Iran, Iraq, Kyrgyzstan, Mongolia, Tajikistan, Turkmenistan, Turkey, Uzbekistan (Gebler 1830, Motschulsky 1840, Mulsant 1850, Crotch 1874, Barovskij 1927, Mader 1955, Bielawski 1975, Bielawski 1984, Kovář 1997) and Xinjiang, China (present study) (Fig. 4).
Checklists
Checklist of Chinese Chilocorini sensu Li et al. 2020
Brumoides
Chapin, 1965
C1D7BFF4-D441-5864-BA87-EA89FF4A197B
Brumoides hainanensis
Miyatake, 1970
B90B50F7-8D94-52DA-91BD-3D699476C1B6
Distribution
China (Miyatake 1970).
Brumoides lineatus
(Weise, 1885)
791DCD0B-A327-58B2-B5AF-F7FEB09F146A
Distribution
China, Myanmar, Thailand, Bangladesh, India, Sri Lanka, Nepal, Pakistan and Ceylon (Weise 1885, Miyatake 1970, Poorani 2002).
Brumoides maai
Miyatake, 1970
55161FAB-69B9-5C69-92E6-2875FA2880C5
Distribution
China (Miyatake 1970).
Brumoides ohtai
Miyatake, 1970
931B69C1-43A9-58BF-9170-6E2540BB448C
Distribution
China (Miyatake 1970).
Brumus
Mulsant, 1850
6D97AC34-4DB2-5549-A8AC-DEDA064BA76C
Brumus octosignatus
(Gebler, 1830)
8DCC3591-1A72-593D-BFDC-2E14429E1874
Distribution
China, Azerbaijan, Armenia, France, Italy, Russia: south European Territory, Afghanistan, Iran, Iraq, Kyrgyzstan, Mongolia, Tajikistan, Turkmenistan, Turkey and Uzbekistan (Gebler 1830, Motschulsky 1840, Mulsant 1850, Crotch 1874, Barovskij 1927, Mader 1955, Bielawski 1975, Bielawski 1984, Kovář 1997).
Chilocorus
Leach, 1815
45531EBC-C889-5E16-B124-7A819DAE4F79
Chilocorus albusolomus
(Li &Wang, 2017)
3AF2B3AD-6D3A-5C11-8136-600B09A769FF
Distribution
China (Li et al. 2017b).
Chilocorus alishanus
Sasaji, 1968
4905DAF8-8834-5104-B150-15FDEA4A7387
Distribution
China (Sasaji 1968).
Chilocorus bijugus
Mulsant, 1853
F09738D9-694E-5A6D-BCB3-73A84F30427D
Distribution
China, Japan, India, Nepal, Pakistan and Palaearctic (Mulsant 1853, Kovář 2007).
Chilocorus bipustulatus
(Linnaeus, 1758)
85C81AFD-2392-558A-A259-1840F0846712
Distribution
China, Europe, Middle East, Central Asia and North Africa (Linnaeus 1758, Li et al. 2018).
Chilocorus chalybeatus
Gorham, 1892
D151B203-56C4-5B92-9D2D-85DECE6E49B3
Distribution
China (Gorham 1892, Kovář 2007, Li et al. 2018).
Chilocorus chinensis
Miyatake, 1970
E87AB265-8D3D-5A20-8BB1-95BEDD3F9FF0
Distribution
China (Miyatake 1970, Li et al. 2018).
Chilocorus circumdatus
(Gyllenhal, 1808)
B65D3A0E-FA95-532C-A5B9-96252D2C718F
Distribution
China, Indonesia, India, Sri Lanka and introduced to Australia and America (Gyllenhal 1808, Ślipiński 2007, Kovář 2007, Li et al. 2018).
Chilocorus esakii
Kamiya, 1959
991E8AB0-2D2F-57CA-B2F5-A335CFB43D73
Distribution
China, Japan (Kamiya 1959, Li et al. 2018).
Chilocorus geminus
Zaslavskij, 1962
A4209690-8C5D-5A64-A0BD-E812386D1605
Distribution
China, Mongolia, Turkey and Central Asia (Zaslavskij 1962, Bielawski 1975, Kovář 2007, Li et al. 2018).
Chilocorus hauseri
Weise, 1895
5F196210-6323-5E08-AA54-1D2B3EC8D25C
Distribution
China, India and Myanmar (Weise 1895, Korschefsky 1932, Kovář 2007, Li et al. 2018).
Chilocorus hupehanus
Miyatake, 1970
68CC0E29-A578-593E-AF05-79F1D07345BB
Distribution
China (Miyatake 1970).
Chilocorus kuwanae
Silvestri, 1909
65527081-C9DB-5CD4-AF88-182BE71C2F95
Distribution
China, Japan, North Korea and Italy and introduced to America (Silvestri 1909, Gordon 1985, Kovář 2007, Li et al. 2018).
Chilocorus melas
Weise, 1898
B62642EB-2D3E-5753-8D18-49B58C1997D6
Distribution
China, Myanmar, Thailand, Laos, India, Nepal, Bhutan and Indonesia (Weise 1898, Miyatake 1970, Poorani 2002, Li et al. 2018).
Chilocorus metasternalis
(Miyatake, 1970)
894CF8A1-FF01-501F-8A29-AEF030F1DCA3
Distribution
China, Laos, Vietnam, Singapore and Indonesia (Miyatake 1970).
Chilocorus nigricaeruleus
Li & Wang, 2018
3E8BEDE9-074A-5832-B1B6-DEBE7B310BE8
Distribution
China (Li et al. 2018).
Chilocorus nigrita
(Fabricius, 1798)
EE478024-15C8-53B6-8682-8891F1CF7D91
Distribution
China, Myanmar, Indonesia, India, Pakistan, Sri Lanka, Bangladesh, Australian, USA, Brazil and South Africa (Fabricius 1798, Crotch 1874, Poorani 2002, Kovář 2007, Li et al. 2018).
Chilocorus rubidus
Hope, 1831
3D230278-A1BF-53AB-9677-1AE2D30D99D0
Distribution
China, Mongolia, Korea, Japan, India, Nepal, Indonesia and Siberia (Hope 1831, Kovář 2007, Li et al. 2018).
Chilocorus rufitarsis
Motschulsky, 1853
C587DF55-9353-54EF-8AD1-B076B8F1DCB7
Distribution
China and Vietnam (Motschulsky 1853, Korschefsky 1932, Kovář 2007, Li et al. 2018).
Chilocorus shirozui
Sasaji, 1968
F7567A00-A9F8-5784-8B73-8605A13C98AF
Distribution
China (Sasaji 1968, Li et al. 2018).
Chilocorus strenotubus
Li & Wang, 2018
FE188300-49C7-5748-918D-AABA36906CCB
Distribution
China (Li et al. 2018).
Chilocorus yunlongensis
Cao & Xiao, 1984
38FF0EAF-9949-5E22-82FB-74F3B93443BE
Distribution
China (Cao and Xiao 1984, Li et al. 2018).
Chilocorus politus
Mulsant, 1850
777DC001-0AD5-570E-88FE-5A74AA32C452
Distribution
China, Thailand, Laos, India, Nepal, Bhutan and Indonesia (Mulsant 1850, Poorani 2002, Kovář 2007, Li et al. 2018).
Chujochilus
Sasaji, 2005
83C54B32-E64B-569B-9C1A-E100C4B54F81
Chujochilus parisensis
Wang & Ren, 2010
D753815F-B596-59EF-BA18-D3F3EFDE8F88
Distribution
China (Wang and Ren 2010).
Chujochilus sagittatus
Wang & Ren, 2010
826D936D-5A08-5EA4-BE96-7839C7E22531
Distribution
China (Wang and Ren 2010).
Exochomus
Redtenbacher, 1843
DA672CD7-7EDD-5525-8B2A-9A972FE9FF97
Exochomus mongol
Barovsky, 1922
A7CDD3E8-7CF5-58CD-A735-EF070910BC7F
Distribution
China, Mongolia, Korea and Far East (Barovskij 1922, Kovář 1997, Kovář 2007, Li et al. 2015b).
Exochomus quadripustulatus
(Linnaeus, 1758)
8DA47D42-8923-5406-B997-A6986E2C5969
Distribution
China, Europe, Russia, Middle East and Mongolia. Introduced to USA and Australia (Linnaeus 1758, Kovář 1997, Kovář 2007, Li et al. 2015b).
Exochomus rubistictus
Li & Ren 2015
E206704F-7AEE-50A2-B7DE-F5A62C1DDFFC
Distribution
China (Li et al. 2015b).
Parexochomus
Barovsky, 1922
84D7CE08-742E-592B-AE56-56034C5C3ECD
Parexochomus nigromaculatus
(Goeze, 1777)
585E64CC-8D15-5A37-86A1-85ABC02CFE60
Distribution
China, Mongolia, Iran, Siberia, Arabia, Europe and Africa (Chapin 1965, Goeze 1777, Ślipiński 2007, Li et al. 2015b).
Parexochomus oligotrichus
Li & Ren 2015
631C7451-643A-5CBE-8045-46CB7046CBC5
Distribution
China (Li et al. 2015b).
Parexochomus semenowi
Weise, 1887
D0A9F7DF-D0B7-5D97-8F99-24CAC26050BD
Distribution
China and Mongolia (Weise 1887, Barovskij 1922, Kovář 2007).
Priscibrumus
Kovář, 1997
5018E44F-F924-5821-8237-9ACAD06A9891
Priscibrumus disjunctus
Canepari, 1997
B9367917-52D5-5022-A85E-AA5395813E9A
Distribution
China and Nepal (Canepari 1997, Poorani 2002).
Priscibrumus himalayensis
(Kapur, 1958)
36D37861-2E37-515F-81EB-064A0E14D041
Distribution
China and Nepal (Kapur 1958, Poorani 2002, Hu et al. 2013).
Priscibrumus uropygialis
(Mulsant, 1853)
ACCCE50E-5357-56D1-8DB5-A52FCE4B014E
Distribution
China; Kashmir, Nepal, India, Bhutan and Pakistan (Mulsant 1850, Crotch 1874, Gordon 1987, Barovskij 1922, Bielawski 1979, Miyatake 1985, Kovář 1997, Canepari 1997, Poorani 2002, Mader 1955).
Renius
Li et Wang, 2017
CA8FFA57-6831-5125-AEBD-23750A0B99B8
Renius cornutus
Li & Wang, 2017
7AB4E103-5BFF-5BC7-9907-56A3916AB330
Distribution
China and India (Li et al. 2017a, Poorani and Rojeet 2019).
Xanthocorus
Miyatake, 1970
EC055B36-58E0-5035-90FB-39EBB9B695A8
Xanthocorus mucronatus
Li & Ren, 2015
B9ED6E68-4E04-5658-A710-78CA04501A1A
Distribution
China (Li et al. 2015a).
Xanthocorus nigromarginatus
Miyatake, 1970
FE2B6E4C-4732-565A-9942-28C8359E9994
Distribution
China (Miyatake 1970, Li et al. 2015a).
Xanthocorus nigrosuturarius
Li & Ren, 2015
02083B29-0124-5BB0-905B-7DDA2BAFF015
Distribution
China (Li et al. 2015a).
Discussion
The genus Priscibrumus was erected by Kovář (1997), when he revised Exochomus and Brumus Mulsant, 1850 from the Palearctic Region. This revision was a huge contribution to our understanding of the relationships between Exochomus and its closely-related genera. Until now, Priscibrumus only contained seven species which mainly occur in the Pamir Mountains, especially in the western part of the Himalaya Mountains (Kovář 1997, Poorani 2002). Priscibrumus himalayensis (Kapur) was the first member of this genus recorded in China (Tibet) by Hu et al. (2013).
Brumus was considered a junior synonym of Exochomus by Ślipiński and Giorgi (2006). Subsequently, Kovář (2007) accepted this point of view and transferred all species of Brumus from the Palearctic Region to Exochomus. The recent phylogenetic studies of Chilocorini indicated that the relationship between Exochomus and Brumus is not so close. Exochomus is the sister group of a large clade containing various genera, for example, Xanthocorus, Priscibrumus, Parexochomus, Brumus and Brumoides, while Brumus is closely related to Brumoides (Li et al. 2020). Actually, Brumus can be easily distinguished from Exochomus by the following characters: elytral epipleuron more or less horizontal; tarsal claws without basal tooth, only slightly swollen at base. In Exochomus, the outer part of elytral epipleuron is distinctly descending; and the tarsal claws have a large triangular tooth at the base.
Supplementary Material
Acknowledgements
The research was supported by the National Natural Science Foundation of China (31802003, 31970441), Science and Technology Planning Project of Guangdong Province (2017A020208060) and the Science and Technology Program of Guangzhou (201804020070, 151800033).
References
- Barovskij V. Revisio specierum palaearticarum Coccinellidarum generis Exochomus Redtb. Annuaire du Musée Zoologique de l’Académie des Sciences de Russie. 1922;23:289–303. [Google Scholar]
- Barovskij V. Revision specierum palaearticarum Coccinellidarum generis Brumus Muls. (Coleoptera). Annuaire du Musée Zoologique de I’ Académie de Sciences de I’ URSS. 1927;28:194–200. [Google Scholar]
- Bielawski R. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei, Nr. 352. Coccinellidae V and VI (Coleoptera) Fragmenta Faunistica (Warszawa) 1975;20:247–271. doi: 10.3161/00159301FF1975.20.16.247. [DOI] [Google Scholar]
- Bielawski R. Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen Museums in Basel. Coleoptera: Faro. Coccinellidae. Entomologica Basiliensia. 1979;4:83–125. [Google Scholar]
- Bielawski R. Coccinellidae (Coleoptera) of Mongolia. Annales Zoologici. 1984;38:281–460. [Google Scholar]
- Canepari C. Coccinellidae (Coleoptera) from the Nepal Himalayas. Stuttgarter Beiträge zur Naturkunde (A) (Biologie) 1997;565:1–65. [Google Scholar]
- Cao C. Y., Xiao N. N. New species of Coccinellidae from Yunnan, China. Entomotaxonomia. 1984;6:109–132. Chinese. [Google Scholar]
- Chapin E. A. The genera of the Chilocorini (Coleoptera, Coccinellidae). Bulletin of the Museum of Comparative Zoology, Harvard University. 1965;133:227–27. [Google Scholar]
- Che L. H., Zhang S. Q., Li Y., Liang D., Pang H., Ślipiński A., Zhang P. Genome-wide survey of nuclear protein-coding markers for beetle phylogenetics and their application in resolving both deep and shallow-level divergences. Molecular Ecology Resources. 2017;17(6):1342–1358. doi: 10.1111/1755-0998.12664. [DOI] [PubMed] [Google Scholar]
- Crotch G. R. A Revision of the Coleopterous Family Coccinellidae. E. W. Janson; London: 1874. 311 [Google Scholar]
- Escalona H. E., Zwick A., Li H. S., Li H. S., Wang X. M., Pang H., Hartley D., Jermiin L. S., Nedvěd O., Misof B., Niehuis O., Ślipiński A., Tomaszewska W. Molecular phylogeny reveals food plasticity in the evolution of true ladybird beetles (Coleoptera: Coccinellidae: Coccinellini) BMC Evolutionary Biology. 2017;17(1):151. doi: 10.1186/s12862-017-1002-3. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Fabricius J. C. Supplementum entomologiae systematicae. Proft and Storch; Copenhagen: 1798. 572 [Google Scholar]
- Ledebour C. F. Gebler A. Reise durch das Altai-Gebirge und die songorische Kir gisen-Steppe. G. Reimer; Berlin: 1830. Bemerkungen über die Insecten Sibiriens, vorzüglich des Altai. (Part 3)427 [Google Scholar]
- Goeze A. E. Entomologische Beitrage I. Weidmanns Erben und Reich; Leipzig: 1777. 736 [Google Scholar]
- Gordon R. D. The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society. 1985;93:1–912. [Google Scholar]
- Gordon R. D. A catalogue of the Crotch collection of Coccinellidae (Coleoptera). Occasional Papers on Systematic Entomology. 1987;3:1–46. [Google Scholar]
- Gorham H. S. Coleoptera from Central China and the Korea. The Entomologists. 1892;25(4):81–85. [Google Scholar]
- Schönherr C. J. Gyllenhal L. Synonyma Insectorum, oder: Versuch einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum & c. geordnet. Erster Band. Eleutherata oder Käfer. Zweiter Theil. Sprechus-Cryptocepalus; C.F Marquard, Stockholm: 1808. [New taxa.]424 [Google Scholar]
- Gray J. E. Hope F. W. Zoologica1 Miscellany. Treutel, Wurtz and Co; London: 1831. Synopsis of the new species of Nepal insects in the collection of Major General Hardwicke.40 [Google Scholar]
- Hu S. C., Lin X. W., Wang B. H. Coccinellidae of the Qinghai-Xizang. Henan Science and Technology Press; 2013. 213 [Google Scholar]
- Kamiya H. A revision of the tribe Chilocorini of Japan and the Loochoos (Coleoptera: Coccinellidae) Kontyu. 1959;27(2):99–105. [Google Scholar]
- Kapur A. P. Coccinellidae of Nepal. Records of the Indian Museum. 1958;53:309–338. [Google Scholar]
- Junk W., Schenkling S. Korschefsky R. Coleopterorum Catalogus. W. Junk; Berlin: 1932. Pars 120: Coccinellidae. II. pp. 225–659. [DOI] [Google Scholar]
- Kovář I. Revision of the genera Brumus Muls. and Exochomus Redtb. (Coleoptera: Coccinellidae) of the Palaearctic region. Part I. Acta Entomologica Musei Nationalis Pragae. 1997;44:5–124. [Google Scholar]
- Löbl I., Smetana A. Kovář I. Catalogue of Palaearctic Coleoptera. Apollo Books; Stenstrup: 2007. New nomenclatorial and taxonomic acts and comments Coccinellidae, pp. 568–631.905 [Google Scholar]
- Łączyński P., Tomaszewska W. Chapinaria, new genus of Chilocorini for Endochilus meridionalis Sicard from Africa (Coleoptera: Coccinellidae) Annales Zoologici (Warszawa) 2012;62(1):1–9. doi: 10.3161/000345412X633658. [DOI] [Google Scholar]
- Linnaeus C. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species cum chatacteribus, differentiis, synonymis, locis. Holmiae: Laurentii Salvii; 1758. 823 [Google Scholar]
- Li W. J., Chen X. S., Wang X. M., Ren S. X. Contribution to the genus Xanthocorus Miyatake (Coleoptera, Coccinellidae, Chilocorini) ZooKeys. 2015;511:89–98. doi: 10.3897/zookeys.511.9584. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Li W. J., Huo L. Z., Wang X. M., Chen X. S., Ren S. X. The genera Exochomus Redtenbacher, 1843 and Parexochomus Barovsky, 1922 (Coleoptera: Coccinellidae: Chilocorini) from China, with descriptions of two new species. The Pan-pacific Entomologist. 2015;91(4):291–304. doi: 10.3956/2015-91.4.291. [DOI] [Google Scholar]
- Li W. J., Huo L. Z., Ahrens D., Ren S. X., Wang X. M. Renius cornutus, a new genus and species of Chilocorini from Tibet, China (Coleoptera, Coccinellidae) ZooKeys. 2017;678:121–128. doi: 10.3897/zookeys.678.11862. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Li W. J., Huo L. Z., Chen X. S., Ren S. X., Wang X. M. A new species of the genus Phaenochilus Weise from China (Coleoptera, Coccinellidae, Chilocorini) ZooKeys. 2017;644:33–41. doi: 10.3897/zookeys.644.9825. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Li W. J., Huo L. Z., Wang D., Ahrens D., Wang X. M. Contribution to the genus Chilocorus Leach, 1815 (Coleoptera: Coccinellidae: Chilocorini), with descriptions of two new species from China. European Journal of Taxonomy. 2018;469:1–34. doi: 10.5852/ejt.2018.469. [DOI] [Google Scholar]
- Li W. J., Łączyński P., Escalona H. E., Eberle J., Huo L. Z., Chen X. S., Huang W. D., Chen B. X., Ahrens D., Ahrens D., Tomaszewska W., Wang X. M. Combined molecular and morphological data provide insights into the evolution and classification of Chilocorini ladybirds (Coleoptera: Coccinellidae) Systematic Entomology. 2020;45:447–463. doi: 10.1111/syen.12409. [DOI] [Google Scholar]
- Mader L. Evidenz der palearktischen Coccinelliden und ihrer Aberrationen in Wort und Bild. 2. Entomologische Arbeiten aus dem Museum G. Frey Tutzing bei Muenchen. 1955;6:765–1035. [Google Scholar]
- Magro A., Lecompte E., Magne F., Hemptinne J., Crouau-Roy B. Phylogeny of ladybirds (Coleoptera: Coccinellidae): are the subfamilies monophyletic? Molecular Phylogenetics and Evolution. 2010;54:833–848. doi: 10.1016/j.ympev.2009.10.022. [DOI] [PubMed] [Google Scholar]
- Miyatake M. The East-Asian coccinellid beetles preserved in the California Academy of Sciences. Tribe Chilocorini. Memoirs of the College of Agriculture, Ehime University. 1970;14(3):19–56. [Google Scholar]
- Miyatake M. Coccinellidae collected by the Hokkaido University Expedition to Nepal Himalaya, 1968 (Coleoptera) Insecta Matsumurana (New Series) 1985;30:1–33. [Google Scholar]
- Motschulsky V. Insectes du Caucase et des provinces transcasiennes recueillis et decrits (Continuation) Bulletin de la Société Impériale des Naturalistes de Moscou. 1840;13:169–180. [Google Scholar]
- Motschulsky V. Diagnoses de Coléoptères nouveaux, trouvés par M.M. Tatarinoff et Gasckéwitsch aux environs de Pékin. Etudes Entomologiques. 1853;2:44–51. [Google Scholar]
- Mulsant M. E. Species des Coléoptères Trimères Sécuripalpes. 2. Annales des Sciencies Physiques et Naturelles, d’Agriculture et d’Industrie, publiées par la Société nationale d’Agriculture, etc.; de Lyon, Deuxième Série: 1850. 1–1104 [Google Scholar]
- Mulsant M. E. Supplément à la Monographie des Coléoptères Trimerès Sécuripapes. Annales de la Société linnéenne de Lyon (Nouvelle Série) (2) 1853;2:129–333. [Google Scholar]
- Poorani J. An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian Subregion. Oriental Insects. 2002;36:307–383. doi: 10.1080/00305316.2002.10417335. [DOI] [Google Scholar]
- Poorani J., Rojeet T. New additions to Indian fauna of Coccinellidae (Coleoptera) Oriental Insects. 2019 doi: 10.1080/00305316.2018.1550448. [DOI]
- Sasaji H. A revision of the Formosan Coccinellidae (II) tribes Stethorini, Aspidimerini and Chilocorini (Coleoptera) Etizenia. 1968;32:1–24. [Google Scholar]
- Seago A. E., Giorgi J. A., Li J. H., Ślipiński A. Phylogeny, classification and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on simultaneous analysis of molecular and morphological data. Molecular Phylogenetics and Evolution. 2011;60:137–151. doi: 10.1016/j.ympev.2011.03.015. [DOI] [PubMed] [Google Scholar]
- Silvestri F. Nuovo Coccinellide introdotto in Italia. Rivista Coleotterologica Italiana. 1909;7:126–129. [Google Scholar]
- Ślipiński A., Giorgi J. A. Revision of the Australian Coccinellidae (Coleoptera). Part 6. Tribe Chilocorini. Annales Zoologici (Warszawa) 2006;56(2):265–304. [Google Scholar]
- Ślipiński A. Australian ladybird beetles (Coleoptera: Coccinellidae), their biology and classification. ABRS; Canberra: 2007. [Google Scholar]
- Arnett R. H., Thomas M. C., Skelley P. E., Frank J. H. Vandenberg N. J. American Beetles, Vol. 2. CRC Press; Boca Raton: 2002. Coccinellidae . [DOI] [Google Scholar]
- Wang X. M., Ren S. X. Two new species of Chujochilus Sasaji, 2005 (Coleoptera: Coccinellidae) from China. Annales Zoologici. 2010;60(3):319–32. doi: 10.3161/000345410X535316. [DOI] [Google Scholar]
- Weise J. Beschreibung einiger Coccinelliden. Entomologische Zeitung (Stettin) 1885;46:227–241. [Google Scholar]
- Weise J. Neue sibirische Chrysomeliden und Coccinelliden nebst Bemerkungen über früher beschriebene Arten. Archiv für Naturgeschichte. 1887;53(1):164–238. [Google Scholar]
- Weise J. Neue Coccinelliden, sowie Bemerkungen zu bekannten Arten. Annales de la Société Entomologique de Belgique. 1895;39:120–146. [Google Scholar]
- Weise J. Üeber bekannte und neue Coccinelliden. Archiv für Naturgeschichte. 1898;64(1):225–238. [Google Scholar]
- Zaslavskij V. A. Novyy palearkticheskiy vid Chilocorus (Coleoptera, Coccinellidae) Entomologicheskoe Obozrenie. 1962;41:398–401. [Google Scholar]
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