Research Article |
Corresponding author: Fenglong Jia ( fenglongjia@aliyun.com ) Academic editor: Mariano Michat
© 2022 Zuqi Mai, Jian Hu, Fenglong Jia.
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:
Mai Z, Hu J, Jia F (2022) Additional fauna of Coelostoma Brullé, 1835 from China, with re-establishment of Coelostoma sulcatum Pu, 1963 as a valid species (Coleoptera, Hydrophilidae, Sphaeridiinae). ZooKeys 1091: 15-56. https://doi.org/10.3897/zookeys.1091.79564
|
Six new species of Coelostoma Brullé, 1835 are described from China: Coelostoma bannanicum Mai & Jia, sp. nov., C. dactylopunctum Mai & Jia, sp. nov., C. fortunum Mai & Jia, sp. nov., and C. pseudomartensi Mai & Jia, sp. nov. from Yunnan; C. mixtum Mai & Jia, sp. nov. from Fujian; and C. nankunshanense Mai & Jia, sp. nov. from Guangdong. Coelostoma surkhetensis Hebauer, 2002 is a new record from China (Xizang). Coelostoma huangi Jia, Aston & Fikáček, 2014 is reported from Yunnan, C. hajeki Jia, Aston & Fikáček, 2014 from Hunan, C. jaechi Jia, Lin, Chan, Skale & Fikáček, 2017 from Guangdong, C. turnai Hebauer, 2006 from Fujian, Guizhou and Chongqing, and C. wui Orchymont, 1940 from Shanxi and Zhejiang, all for the first time. Coelostoma transcaspicum Reitter, 1906 is excluded from Chinese fauna. Coelostoma sulcatum Pu, 1963 is confirmed as a valid species and its variations of aedeagus are illustrated. The specimens treated as C. wui Orchymont, 1940 by previous authors possibly contain two species. The diversity and habitats of Chinese Coelostoma are discussed. A modified key to the species of Chinese Coelostoma is provided.
habitat, modified key, new records, new species, valid name, variations
Coelostoma Brullé, 1835 is a typical Old World group, with most of the species distributed in the Oriental, Afrotropical, and Palearctic regions. Only a few species occur in the Australian region (
Coelostoma can be separated from other genera of Coelostomatini by following characters combined: broadly oval and convex body shape; loosely segmented antennal club; prosternum convex medially; mesoventrite with an elevated arrowhead-shaped process in the middle; metaventral process projecting anteriorly between mesocoxae, abutting mesoventral elevation; the first metatarsomere distinctly longer than the second one; elytra with dense punctures and sutural stria, without serial punctures in most species; the first abdominal ventrite with or without carina medially (e.g.,
A total of 24 species has been recorded from China since 1874 (
The aim of this study is to describe the new species, update the species of the Chinese fauna, and verify the status of C. sulcatum Pu, 1963 as well as promoting the knowledge of Chinese Coelostoma. Until now, a total of 30 species has been recorded in China including the six new species in this paper.
Representative specimens were dissected. After 8 min in 10% KOH at 100 °C, dissected male genitalia were transferred to a drop of distilled water, remaining membranes were removed under a compound microscope, and the cleaned genitalia were subsequently mounted in a drop of soluble resin on a piece of paper card attached below the respective specimen after photography. Habitus photographs were taken using a Nikon DS-Ri2 mounted on a Nikon SMZ25; layers were captured and stacked in the NIS-Elements software. Photographs of genitalia were taken using a Zeiss AxioCam HRc mounted on a Zeiss AX10 microscope with the Axio Vision SE64 software. These images were then stacked in Helicon focus (v7.0.2). Habitat images were taken using Canon or Nikon digital camera. SEM photographs were taken with a Phenom Pro scanning electronic microscope. All images were digitally enhanced using Adobe Photoshop CS6. Label data of the type specimens are cited verbatim and enclosed in double quotes; a slash divides separate rows on the same label, a double slash divides separate labels. Morphological terminology used in the description largely follows
The following additional specimen was examined for comparative purposes:
Dactylosternum latum (Sharp, 1873): 1 male (
Holotype
: male (
Length 5.4–5.7 mm. Head, pronotum and elytra with similar punctation. Prosternal carina with a prominent tooth anteromedially. Elytra slightly parallel-sided in the middle, without serial punctures laterally. Mesofemora densely pubescent, except on extreme apex. First abdominal ventrite with distinct median carina on basal two-thirds. Fifth ventrite slightly emarginate and with a row of stout setae apically. Aedeagus (Fig.
Form and colour
(Fig.
Head. Dorsal surface with dense fine punctures. Interstices between punctures smooth. Clypeus subtruncate anteriorly. Eyes of moderate size, distinctly emarginated anteriorly in lateral view, separated by ca. 3.6 × the width of one eye. Mentum strongly emarginate anteriorly and depressed in anterior half, with sparse punctures except on the depressed portion. Antennae with 9 antennomeres, antennal club (antennomeres 7–9) densely pubescent. Maxillary palpomere 2 strongly swollen, palpomere 4 truncate apically, slightly longer than palpomere 3. Gula narrow and glabrous.
Thorax. Pronotum widest posteriorly, gradually narrowed anteriad, with punctures as on head, anterolateral angles obtusely rounded, posterolateral angles blunt, anterior and lateral margins with narrow marginal bead. Prosternum with a carina medially and a prominent tooth anteromedially. Scutellum almost in shape of equilateral triangle, with punctures finer than those on pronotum. Elytra with more or less coarser punctures than those on pronotum, punctures on lateral and posterior portions somewhat coarser than those on disc; elytra without serial punctures; sutural stria reaching anterior third of elytra; lateral margin of elytra with bead but not explanate.
Legs. Pro- and mesofemora bearing dense pubescence, except on extreme apex. Metafemora not pubescent, with dense microsculptures and spares fine punctures. Meso- and Metatibiae slightly flattened, with strong apical spurs and series of sparse stout spines laterally. Tarsi with long dorsal setae and gold ventral setae; metatarsi with fifth tarsomere almost as long as third and fourth combined. Claws curved, with a pair of long setae beneath.
Abdomen. Abdominal ventrites densely pubescent. First ventrite with distinct median carina on basal two-thirds. Fifth ventrite with fine marginal bead and slightly emarginated apically, with a row of stout setae apically.
Male genitalia
(Fig.
This species is named after the type locality, Xishuangbanna Dai Autonomous Prefecture.
On the basis of private communication to the collector, Dr. Ri-Xin Jiang (Guizhou University), the specimens were collected under stones at the edges of a mountain stream.
This species is very similar to C. coomani Orchymont, 1932 and C. jaechi Jia, Lin, Chan, Skale & Fikáček, 2017 in the weakly emarginate apex of the median lobe, and to C. surkhetensis Hebauer, 2002 in the shape of the median lobe. It can be distinguished from C. coomani (
Only known from type locality. China (Yunnan).
Holotype
: male (
Length 4.3–4.5 mm. Head and pronotum with similar punctation. Prosternum carinate medially, with a fine tooth anteromedially. Elytra not parallel-sided in the middle, each elytron with ten distinct rows of serial punctures; intervals between series with two sizes of punctures especially in posterior half of elytron, the finer punctures as on pronotum and much finer and shallower than the coarser punctures, coarser punctures almost as coarse as those of the series (Fig.
Form and colour
(Fig.
Head. Dorsal surface with dense fine punctures. Interstices between punctures smooth. Clypeus subtruncate anteriorly. Eyes of moderate size, slightly emarginated anteriorly in lateral view, separated by ca. 5.3 × the width of one eye. Mentum emarginate anteriorly and depressed in anterior half, with sparse punctures except on the depressed portion. Antennae with nine antennomeres, antennal club (antennomeres 7–9) densely pubescent. Maxillary palpomere 2 strongly swollen, palpomere 4 truncate apically, slightly longer than palpomere 3. Gula narrow and glabrous.
Thorax. Pronotum widest posteriorly, gradually narrowed anteriad, with punctures as on head, anterolateral angles obtusely rounded, posterolateral angles blunt, anterior and lateral margins with narrow marginal bead. Prosternum with a carina medially and slightly projecting anteromedially. Scutellum in shape of equilateral triangle, with punctures as on pronotum. Each elytron with ten distinct rows of serial punctures; intervals between series with two sizes of punctures especially in posterior half of elytron, the finer punctures as on pronotum and much finer and shallower than the coarser punctures, coarser punctures almost as coarse as those of the series (Fig.
Legs. Pro- and mesofemora bearing dense pubescence, except on extreme apex. Metafemora not pubescent, with dense microsculptures and spares fine punctures. Meso- and Metatibia slightly flattened, with strong apical spurs and series of sparse stout spines laterally. Tarsi with long dorsal setae and gold ventral setae; metatarsi with fifth tarsomere almost as long as third and fourth combined. Claws curved, with a pair of long setae beneath.
Abdomen. Abdominal ventrites densely pubescent. First ventrite with complete median carina. Fifth ventrite slightly emarginate and with fine marginal bead, with a row of stout setae apically.
Male genitalia
(Fig.
The species name is a combination of Dactylosternum, a genus in the same tribe Coelostomatini, and the Latin punctum. The name refers to the fact that this species with distinct serial punctures on elytra which is similar to many species of Dactylosternum Wollaston, 1854.
Aquatic. Adults were found on the edges of a forest stream at night.
This species is easily recognized as a member of Dactylosternum by the ten rows of serial punctures on elytra, which is different to any known Coelostoma species. Coelostoma martensi Hebauer, 2002 and C. gentilii Jia, Aston & Fikáček, 2014 (Fig.
Only known from type locality. China (Yunnan).
Holotype
: male (
Length 4.4–4.6 mm. Head and pronotum with similar punctation. Prosternum carinate medially, with a prominent tooth anteromedially. Elytra not parallel-sided in the middle, each elytron with 10 distinct rows of serial punctures; intervals between series with two sizes of punctures especially in posterior half of elytron, all finer than those of the series, the finer punctures finer and shallower than the coarser punctures but not extremely so (Fig.
Form and colour
(Fig.
Head. Dorsal surface with dense fine punctures. Interstices between punctures smooth. Clypeus subtruncate anteriorly. Eyes of moderate size, slightly emarginated anteriorly in lateral view, separated by ca. 4.2 × the width of one eye. Mentum emarginated anteriorly and depressed in anterior half, with sparse punctures and transverse microsculpture. Antennae with 9 antennomeres, antennal club (antennomeres 7–9) densely pubescent. Maxillary palpomere 2 strongly swollen, palpomere 4 truncate apically, slightly longer than palpomere 3. Gula narrow and glabrous.
Thorax. Pronotum widest posteriorly, gradually narrowed anteriad, with punctures slightly sparser than those on head, anterolateral angles obtusely rounded, posterolateral angles blunt, anterior and lateral margins with narrow marginal bead. Prosternum with a carina medially and a prominent tooth anteromedially. Scutellum in shape of equilateral triangle, with punctures as on pronotum. Each elytron with ten distinct rows of serial punctures; intervals between series with two sizes of punctures especially in posterior half of elytron, all finer than those of the series, the finer punctures finer and shallower than the coarser punctures but not extremely so (Fig.
Legs. Pro- and mesofemora bearing dense pubescence, except on extreme apex. Metafemora not pubescent, with dense microsculptures and spares fine punctures. Meso- and Metatibia slightly flattened, with strong apical spurs and series of sparse stout spines laterally. Tarsi with long dorsal setae and gold ventral setae; metatarsi with fifth tarsomere almost as long as third and fourth combined. Claws curved, with a pair of long setae beneath.
Abdomen. Abdominal ventrites densely pubescent. First ventrite with distinct median carina on basal two-thirds. Fifth ventrite slightly emarginate and with fine marginal bead, with a row of stout setae apically.
Male genitalia
(Fig.
This new species is derived from the Latin adjective fortuna, fortunate, meaning the senior author was lucky to collect the new species.
All specimens were collected in a light trap
This species also with ten rows of serial punctures on elytra as C. dactylopunctum sp. nov. It can be distinguished from the latter by apex of median lobe truncate and not emarginate (Fig.
Only known from type locality. China (Yunnan).
Holotype
: male (
Length 6.13 mm. Head, pronotum and elytra with similar punctation. Prosternum carinate medially, with a prominent tooth anteromedially. Elytra parallel-sided in the middle, without serial punctures laterally. Mesofemora densely pubescent, except on extreme apex. First abdominal ventrite with median carina on basal one-thirds. Fifth ventrite slightly emarginate and with a row of stout setae apically. Aedeagus (Fig.
Form and colour
(Fig.
Head. Dorsal surface with dense fine punctures. Interstices between punctures smooth. Clypeus subtruncate anteriorly. Eyes of moderate size, distinctly emarginate anteriorly in lateral view, separated by ca. 4 × the width of one eye. Mentum strongly emarginated anteriorly and depressed in anterior half, with sparse fine punctures. Antennae with nine antennomeres, antennal club (antennomeres 7–9) densely pubescent. Maxillary palpomere 2 strongly swollen, palpomere 4 truncate apically, slightly longer than palpomere 3. Gula narrow and glabrous.
Thorax. Pronotum widest posteriorly, gradually narrowed anteriad, with punctures as on head, anterolateral angles obtusely rounded, posterolateral angles blunt, anterior and lateral margins with narrow marginal bead. Prosternum with a carina medially and a prominent tooth anteromedially. Scutellum in shape of equilateral triangle, with punctures finer and denser than those on pronotum. Elytra with punctures as on pronotum; elytra without serial punctures; sutural stria reaching anterior half of elytra; lateral margin of elytra with bead but not explanate.
Legs. Pro- and mesofemora bearing dense pubescence, except on extreme apex. Metafemora not pubescent, with dense microsculptures and spares fine punctures. Meso- and Metatibia slightly flattened, with strong apical spurs and series of sparse stout spines laterally. Tarsi with long dorsal setae and gold ventral setae; metatarsi with fifth tarsomere almost as long as third and fourth combined. Claws curved, with a pair of long setae beneath.
Abdomen. Abdominal ventrites densely pubescent. First ventrite with distinct median carina on basal one-thirds. Fifth ventrite with fine marginal bead and slightly emarginate apically.
Male genitalia
(Fig.
This species is derived from the Latin verb mixtus, mix, refers to the fact that this species is similar to Coelostoma vagum Orchymont, 1940 in shape of the median lobe and similar to C. wui Orchymont, 1940 in shape of parameres.
Unknown, this species was collected with C. wui Orchymont, 1940 in the same place.
The holotype of this species was identified as C. vagum Orchymont, 1940 in
Only known from type locality. China (Fujian).
Holotype
: male (
Length 4.7–5.0 mm. Head, pronotum and elytra with similar punctation. Prosternum carinate medially, with a prominent tooth anteromedially. Elytra slightly parallel-sided in the middle, without serial punctures laterally. Mesofemora densely pubescent, except on extreme apex. First abdominal ventrite with median carina on basal two-thirds. Fifth ventrite slightly emarginate and with a row of stout setae apically. Aedeagus (Fig.
Form and colour
(Fig.
Head. Dorsal surface with dense fine punctures. Interstices between punctures smooth. Clypeus subtruncate anteriorly. Eyes of moderate size, distinctly emarginate anteriorly in lateral view, separated by ca. 3.5 × the width of one eye. Mentum strongly emarginated anteriorly and depressed in anterior half, with sparse punctures and dense transverse microsculpture. Antennae with nine antennomeres, antennal club (antennomeres 7–9) densely pubescent. Maxillary palpomere 2 strongly swollen, palpomere 4 truncate apically, slightly longer than palpomere 3. Gula narrow and glabrous.
Thorax. Pronotum widest posteriorly, gradually narrowed anteriad, with punctures as on head, anterolateral angles obtusely rounded, posterolateral angles blunt, anterior and lateral margins with narrow marginal bead. Prosternum with a carina medially and a prominent tooth anteromedially. Scutellum in shape of equilateral triangle, with punctures finer and denser than those on pronotum. Elytra with punctures as on pronotum, punctures on lateral and posterior portions somewhat coarser than those on disc; elytra without serial punctures; sutural stria reaching anterior third of elytra; lateral margin of elytra with bead but not explanate.
Legs. Pro- and mesofemora bearing dense pubescence, except on extreme apex. Metafemora not pubescent, with dense microsculptures and spares fine punctures. Meso- and Metatibia slightly flattened, with strong apical spurs and series of sparse stout spines laterally. Tarsi with long dorsal setae and gold ventral setae; metatarsi with fifth tarsomere almost as long as third and fourth combined. Claws curved, with a pair of long setae beneath.
Abdomen. Abdominal ventrites densely pubescent. First ventrite with distinct median carina on basal two-thirds. Fifth ventrite with fine marginal bead and slightly emarginate apically.
Male genitalia
(Fig.
This species is named after the type locality, Nankunshan, a nature reserve in Guangdong Province.
(Figs
Only known from type locality. China (Guangdong).
Holotype
: male (
Length 5.4 mm. Head and pronotum with similar punctation. Prosternum carinate medially, with a prominent tooth anteromedially. Elytra slightly parallel-sided in the middle, each elytron with ten serial punctures, somewhat difficult to separate them from the ground punctures in anterior half of elytron; intervals between series with two sizes of punctures especially in posterior half of elytron, coarser punctures slightly finer than those of the series (Fig.
Form and color
(Fig.
Head. Dorsal surface with dense fine punctures. Interstices between punctures smooth. Clypeus subtruncate anteriorly. Eyes of moderate size, slightly emarginate anteriorly in lateral view, separated by ca. 4.5 × the width of one eye. Mentum strongly emarginate anteriorly and depressed in anterior half, with sparse punctures and transverse microsculpture. Antennae with 9 antennomeres, antennal club (antennomeres 7–9) densely pubescent. Maxillary palpomere 2 strongly swollen, palpomere 4 truncate apically, slightly longer than palpomere 3. Gula narrow and glabrous.
Thorax. Pronotum widest posteriorly, gradually narrowed anteriad, with punctures as on head, anterolateral angles obtusely rounded, posterolateral angles blunt, anterior and lateral margins with narrow marginal bead. Prosternum with a carina medially and a prominent tooth anteromedially. Scutellum slightly longer than wide, in shape of equilateral triangle, with punctures as on pronotum. Elytra with ground punctures as on pronotum, becoming coarser posteriorly. Each elytron with ten rows of serial punctures, somewhat difficult to separate them from the ground punctures in anterior half of elytron; intervals between series with two sizes of punctures especially in posterior half of elytron, coarser punctures slightly finer than those of the series; series 1 overlap with sutural stria; series 8 and 9 slightly sulcate posteriorly; series 10 short and becoming indistinct posteriorly (Fig.
Legs. Pro- and mesofemora bearing dense pubescence, except on extreme apex. Metafemora not pubescent, with dense microsculptures and spares fine punctures. Meso- and Metatibia slightly flattened, with strong apical spurs and series of sparse stout spines laterally. Tarsi with long dorsal setae and gold ventral setae; metatarsi with fifth tarsomere almost as long as third and fourth combined. Claws curved, with a pair of long setae beneath.
Abdomen. Abdominal ventrites densely pubescent. First ventrite with distinct median carina on basal half. Fifth ventrite slightly emarginate and with fine marginal bead, with a row of stout setae apically.
Male genitalia
(Fig.
This species name is a combination of the Latin pseudo-, false, and another species of the genus, Coelostoma martensi. The name refers to the fact that this species is similar to C. martensi in some morphology characters.
This species lives mixed with Coelostoma dactylopunctum sp. nov.
This species is similar to Coelostoma martensi Hebauer, 2002 (
Only known from type locality. China (Yunnan).
China: Xizang: 1 male (
Length 5.4–5.8 mm. Head, pronotum and elytra with similar punctation. Prosternum carinate medially, with a prominent tooth anteromedially. Elytra not parallel-sided in the middle, without serial punctures laterally. Mesofemora densely pubescent, except on extreme apex. First abdominal ventrite with distinct median carina on basal two-thirds. Fifth ventrite slightly emarginate and with a row of stout setae apically. Aedeagus (Fig.
(Fig.
Previously known only from Nepal (
China: Yunnan: 32 spec. (
In Yunnan, specimens were collected along sides of a rocky stream in a valley at night. Coelostoma coomani Orchymont, 1932 and C. phallicum Orchymont, 1940 were also collected in the same habitat. Adult individuals attracted by light (
China (Guangxi, Jiangxi, Yunnan), Thailand (
China: Hunan: 30 spec. (
China (Guangdong, Guangxi, Hunan). New record for Hunan.
China: Guangdong: 1 male (
China (Hong Kong, Guangdong). New record for Guangdong.
China: Chongqing: 4 spec. (
(Fig.
Only known from China (Fujian, Guizhou, Hubei, Hunan, Chongqing) (
China: Hunan: 4 spec. (
(possibly another undescribed species). China: Guangdong: 5 spec. (
(Figs
The specimens we studied contained two different forms of male genitalia (Fig.
China (Hubei, Hunan, Jiangxi, Shaanxi, Shandong, Shanxi, Taiwan, Xinjiang, Zhejiang), Korea. (
Coelostoma sulcatum Pu, 1963: 77. Type locality: Xishuangbanna Dai Autonomous Prefecture, Yunnan, China.
Coelostoma (Holocoelostoma) stultum
(Walker, 1858):
Coelostoma (Holocoelostoma) bhutanicum
Jayaswal, 1972:
Coelostoma sulcatum: Holotype (Fig.
Aedeagus of Coelostoma sulcatum Pu, 1963 (dorsal view) A, B holotype of C. sulcatum A labels B aedeagus C from Jing’an County (Jiangxi) D from Shenzhen City (Guangdong) E from Longlin County (Guangxi) F from Xishuangbanna (Yunnan) G from Xima (昔马) Town (Yunnan) H from Tongbiguan Town (Yunnan) I from Muotuo County (Xizang) J from Macao. Scale bar: 0.5 mm (A–J).
China: Fujian: 1 spec. (
Length 4.5–5.8 mm. Head, pronotum and elytra with similar punctation. Prosternum moderately convex medially, not carinate, without anteromedian process. Elytra slightly or not parallel-sided in the middle, without serial punctures. Mesofemora without dense pubescence, but with punctures bearing strong setae laterally. First abdominal ventrite not carinate, fifth ventrite emarginate and with a row of stout setae apically. Aedeagus (Fig.
(Figs
Coelostoma sulcatum Pu, 1963 is morphologically variable in shape of aedeagus, especially in parameres. Compared with the original description (
China (Fujian, Guangdong, Guangxi, Jiangxi, Macao, Taiwan, Yunnan, Xizang, Zhejiang).
For complete synonymy, see
China: Guangdong: 3 spec. (
Cambodia: 17 spec. (
Length 4.3–5.2 mm. similar to C. sulcatum in morphological characters. Aedeagus (Fig.
Lateral view of Coelostoma spp. (the red figures indicate the serial number of elytral serial punctures; the white arrows indicate the punctures of series; the circles indicate the coarser punctures; the squares indicate the finer punctures) A Coelostoma dactylopunctum sp. nov. B Coelostoma fortunum sp. nov. Scale bar: 1 mm (A, B).
This species has the similar habitations of C. sulcatum Pu, 1963.
Lateral view of Coelostoma spp. (the red figures indicate the serial number of elytral serial punctures; the white arrows indicate the punctures of series; the square indicates the elytral ground punctures) A Coelostoma pseudomartensi sp. nov. B Coelostoma gentilii Jia, Aston & Fikáček, 2014. Scale bar: 1 mm (A, B).
This species was reported widely distributed in China (Chongqing, Fujian, Guangxi, Guangdong, Jiangxi, Hainan, Hunan, Hong Kong, Shandong, Sichuan, Taiwan, Yunnan, Xizang) (
Aedeagus of Coelostoma wui Orchymont, 1940 (dorsal view) A illustration by
Habitats of Chinese Coelostoma A stone walls with running waters in forest (Yunnan) B wet stone walls with moss (Guangdong) C mountain stream in forest (Guangdong) D mountain river with rocky edges (Shanxi) E wastewater in city downtown (Guangdong) F lowland marshes with vegetation (Guangdong) G artificial lake (Yunnan) H brackish lagoon in mangrove reserve (Macao).
Habitats of Chinese Coelostoma spp. A C. turnai Hebauer, 2006 active at wet rocky ground beside a river at night (Fujian) B C. sulcatum Pu, 1963 on sandy gutterway with shallow flowing waters at night (Guangdong) C C. phallicum Orchymont, 1940 hided under a brick beside the wastewater during the day (Guangdong) D C. phallicum Orchymont, 1940 dived under water and fed on algal mats at night (Guangdong) E C. bifidum Jia, Aston & Fikáček on wet stone wall at night (Guangdong) F C. sulcatum Pu, 1963 on muddy edge of artificial lake at night (Yunnan) G C. nankunshanense sp. nov. on a stone in the middle of a forest stream at night (Guangdong) H C. surkhetensis Hebauer, 2002 in a wet stone on the edges of a forest stream at night (Xizang) I C. wui Orchymont, 1940 on the edges of a mountain river at night (Shanxi) J C. sulcatum Pu, 1963 on muddy edges of brackish lagoon at night, with a marine Amphipoda beside it (Macao) K C. nankunshanense sp. nov. mating at night (Guangdong) L C. wui Orchymont, 1940 ovipositing eggs on wet land at night (Shanxi).
This key is modified based on
1 | Mesofemora densely pubescent except at extreme apex ( |
2 |
– | Mesofemora not pubescent, glabrous, more or less coarsely punctate and sparsely covered by short setae ( |
24 |
2 | Elytra with distinct serial punctures laterally (Figs |
3 |
– | Elytra without serial punctures laterally | 6 |
3 | Elytra serial punctures only visible laterally, without serial punctures on disc (Fig. |
C. gentilii Jia, Aston & Fikáček, 2014 |
– | Elytra series punctures visible on disc (Figs |
4 |
4 | Elytron with 10 serial punctures, somewhat difficult to separate from the ground punctures in anterior half of elytron (Fig. |
C. pseudomartensi sp. nov. |
– | Elytron with 10 distinct serial punctures (Fig. |
5 |
5 | Intervals between series with two sizes of punctures, the small punctures much finer and shallower than the big punctures, big punctures almost as coarse as those of the series (Fig. |
C. dactylopunctum sp. nov. |
– | Intervals between series with two sizes of punctures, all finer than those of the series, the small punctures finer and shallower than the big punctures but not extremely so (Fig. |
C. fortunum sp. nov. |
6 | Body size < 4.0 mm. Pronotum with much finer and sparser punctation than on elytra ( |
C. hongkongense Jia, Aston & Fikáček, 2014 |
– | Body size > 4.0 mm. Pronotum with punctation at most slightly finer and sparser than punctation on elytra. Median lobe of the aedeagus without subapical lateral projections | 7 |
7 | Median lobe of aedeagus trilobate apically | 8 |
– | Median lobe of aedeagus not emarginate to deeply emarginate apically | 10 |
8 | Aedeagus narrowly elongate; Median lobe not wider than paramere ( |
C. phallicum Orchymont, 1940 |
– | Aedeagus relatively wider; median lobe wider than paramere | 9 |
9 | Aedeagus large (ca. 1.1 mm long), median lobe strongly sclerotized, highly modified, saddle-shaped in lateral view, rather shorter than parameres; parameres rather broadened subapically inwards ( |
C. tangliangi Jia, Lin, Chan, Skale & Fikáček, 2017 |
– | Aedeagus small (ca. 0.6 mm long), weakly sclerotized, median lobe plain, only slightly bent in lateral view, not so shorter than parameres; parameres no so broadened subapically inwards ( |
C. horni (Régimbart, 1902) |
10 | Aedeagus large (> 1.5 mm long), parameres largely overlapping apex of median lobe | 11 |
– | Aedeagus smaller (< 1.5 mm long), parameres only slightly longer than median lobe | 14 |
11 | Apex of the median lobe widely rounded or slightly emarginated, parameres broadly widened apically (Fig. |
C. wui Orchymont, 1940 |
– | Apex of the median lobe narrowly rounded or angulate | 12 |
12 | Median lobe widest in the middle, with a small rounded finger like apex; parameres broadly widened apically (Fig. |
C. mixtum sp. nov. |
– | Median lobe nearly parallel-sided in the middle, apex pointed or augulate; parameres not distinctly widened apically | 13 |
13 | Apex of median lobe widely augulate; parameres weakly narrowing in apical third, slightly bent inward ( |
C. taiwanense Liu, Hu & Fikáček, 2020 |
– | Median lobe strongly narrowing near apex, apex with a sharp prominent hook-shaped tooth ventrally; parameres relatively slender, with inner face almost straight (Fig. |
C. vagum Orchymont, 1940 |
14 | Median lobe of aedeagus distinctly emarginate apically; gonopore situated basally or slightly before the midlength of the median lobe | 15 |
– | Median lobe of aedeagus not or slightly emarginate; gonopore situated subapically to apically | 19 |
15 | Median lobe bottle-shaped, strongly broadened basally; gonopore situated basally | 16 |
– | Median lobe widest in the middle; gonopore situated slightly above the midlength of the median lobe | 18 |
16 | Outer face of parameres convex basally; median lobe strongly widened basally, gonopore extremely transverse | 17 |
– | Outer face of parameres nearly straight basally; basal portion of median lobe moderately widened, gonopore transverse round ( |
C. hajeki Jia, Aston & Fikáček, 2014 |
17 | Median strongly broadened basally. Outer face of parameres distinctly incised subapically (Fig. |
C. nankunshanense sp. nov. |
– | Median lobe not so broadened basally. Outer face of parameres slightly curved subapically ( |
C. turnai Hebauer, 2006 |
18 | Median lobe deeply emarginate apically; apex of paramere widened, sharply protruding inwards ( |
C. bifidum Jia, Aston & Fikáček, 2014 |
– | Median lobe shallowly emarginate apically; paramere not widended apically, obtusely truncate at apex ( |
C. coomani Orchymont, 1932 |
19 | Median lobe of aedeagus not emarginate apically | 20 |
– | Median lobe of aedeagus slightly emarginate or truncate apically | 21 |
20 | Median lobe with a distinct subapical tooth and a lateral ridge subapically ( |
C. jaculum Jia, Angus & Bian, 2019 |
– | Median lobe with a rounded apex ( |
C. phototropicum Jia, Angus & Bian, 2019 |
21 | Parameres obliquely truncate inwards apically ( |
C. huangi Jia, Aston & Fikáèek, 2014 |
– | Inner face of parameres rounded or augulate apically | 22 |
22 | Parameres strongly expanded apically, apex of paramere distinctly wider than apex of median lobe | 23 |
– | Parameres not expanded apically, apex of paramere narrower than apex of median lobe (Fig. |
C. surkhetensis Hebauer, 2002 |
23 | Size larger than 5.0 mm. Median lobe widest at midlength (Fig. |
C. bannanicum sp. nov. |
– | Size smaller than 5.0 mm. Median lobe widest at apical third ( |
C. jaechi Jia, Lin, Chan, Skale & Fikáček, 2017 |
24 | Fifth abdominal ventrite slightly emarginate posteromesally, bearing strong setae mesally ( |
25 |
– | Posterior margin of the fifth abdominal ventrite entire, not emarginate in the middle ( |
26 |
25 | Median lobe almost parallel at basal third-fourth, apical fourth distinctly narrowed subapically; outer face of parameres more or less parallel, only slightly curved medially (Fig. |
C. sulcatum Pu, 1963 |
– | Median lobe distinctly narrowed medially, then slightly widened at apical third; outer face of parameres broadened medially (Fig. |
C. stultum (Walker, 1858) |
26 | Aedeagus slender, median lobe gradually attenuate toward apex, sharpened apically. Parameres strongly narrowed from apical fifth to apex, pointed apically ( |
C. orbiculare (Fabricius, 1775) |
– | Aedeagus robust, median lobe and parameres not strongly narrowing apically | 27 |
27 | Posterior femora broad to almost oval in form. Median lobe of aedeagus broad and short, parameres slender ( |
C. vitalisi Orchymont, 1923 |
– | Posterior femora not broadened, aedeagus not as above | 28 |
28 | Body length 4.1–4.2 mm. Mesofemora finely and sparsely punctate. Median lobe of aedeagus strongly broadened at basal half, abruptly narrowed mesally, and almost parallel-sided in apical half, gonopore subtriangular, situated ca. at midlength of the median lobe ( |
C. vividum Orchymont, 1936 |
– | Body length 4.7–5.4 mm. Mesofemora with coarse and dense punctation. Median lobe of aedeagus not so broadened basally and not so extremely narrow from middle to apex, gonopore apical or subapical | 29 |
29 | Median lobe of the aedeagus bottle-shaped, with broad base and abruptly narrowed and gradually slightly narrowed toward apex, gonopore in shape of number 8 ( |
C. fallaciosum Orchymont, 1936 |
– | Median lobe of aedeagus gradually narrowed from base to apex, not abruptly narrowed, gonopore rhomboid in shape (in |
C. subditum Orchymont, 1936 |
All specimens with an emarginate apex were found from the north of Nanling Mountains (Henan, Hubei, Hunan, Shandong, Shaanxi, Shanxi and Zhejiang), one of which (Pingyi County) (Fig.
The genus Coelostoma is a typical tropical group, only several species occurring in temperate region (
Currently, 30 species of Coelostoma are known from China, of which two species are assigned to the subgenus Holocoelostoma, five species to Coelostoma (s. str.), and 23 species to Lachnocoelostoma. Compared with the fauna of other regions, we can come to following conclusions: (1) the Chinese fauna has a large species diversity of Lachnocoelostoma and nearly half of them seem to be very local and are likely endemic (
Previous studies on aquatic beetles indicate that species occurring in standing water tend to have larger ranges than species in running waters (e.g.,
We thank Dr. Robert B. Angus, a specialist on Hydrophiloidea (Natural History Museum, London, UK) for helping us in reviewing the manuscript. We are grateful to Dr. Martin Fikáček (Prague, Czech Republic) and Dr. Yûsuke N Minishima (Kitakyushu Museum of Natural History and Human History, Fukuoka, Japan) for their valuable suggestions and comments on the manuscript. We are grateful to Mr. Yu-Chen Zheng (China Agricultural University), Mr. Hao-Han Mao (Beijing Forestry University), Mr. Zhao-Yang Tang, Dr. Ri-Xin Jiang (Guizhou University), Dr. Jian-Yue Qiu, Dr. Hao Xu (both Mianyang Normal University), Dr. Hai-Tian Song (Fujian Academy of Forestry), Mr. Zhi-Hao Qi (Fujian Agriculture and Forestry University), Dr. Liang Tang (Shanghai Normal University), Mr. Hao-Yi Liu, Dr. Wen-Yi Zhou, and Mr. Zi-Hao Shen for donation of valuable specimens. We thank Mr. Bao-Ping Huang and Mr. Qian-Le Lu for providing habitat images of some species and donating some specimens to us as well. We are grateful to Professor Guo-dong Ren (College of Life Sciences, Hebei University, China) for his loan of specimens. We are also very grateful to Dr. Jun Chen and Miss Kui-yan Zhang (