Research Article |
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Corresponding author: Jibao Jiang ( jibao@sjtu.edu.cn ) Academic editor: John Reynolds
© 2024 Qing Jin, Jiali Li, Jibao Jiang, Jiangping Qiu.
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:
Jin Q, Li J, Jiang J, Qiu J (2024) Four new earthworm species of the genera Amynthas and Metaphire (Oligochaeta, Megascolecidae) from Hunan and Anhui provinces, China. ZooKeys 1210: 247-271. https://doi.org/10.3897/zookeys.1210.125963
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This paper describes four new species earthworms from Hunan and Anhui provinces, China, Amynthas xiangtanensis Qiu & Jin, sp. nov., Amynthas taoyuanensis Qiu & Jin, sp. nov., Amynthas xuanchengensis Jin & Li, sp. nov. and Metaphire donganensis Jin & Jiang, sp. nov. Amynthas xiangtanensis sp. nov., and A. taoyuanensis sp. nov. belong to the Amynthas corticis group. Both have four pairs of intersegmental spermathecal pores in 5/6–8/9; male pores in segment XVIII, separated by 1/3 of body circumference, each on top of a slightly raised porophore, surrounded by several tiny genital papillae. Amynthas taoyuanensis sp. nov. prostate glands are degenerated. Amynthas xuanchengensis sp. nov. belongs to the Amynthas morrisi group, it has two pairs of spermathecal pores in 5/6 and 6/7; male pores in XVIII, separated by 1/3 of body circumference, each on top of a slightly raised, circular porophore. Metaphire donganensis sp. nov. belongs to the Metaphire houlleti group. It has three pairs of spermathecal pores in 6/7–8/9; male pores in XVIII, separated by 1/3 of body circumference, each on the bottom center of the longitudinal copulatory chamber.
Barcode, COI gene, Megascolecidae, new species, Oligochaeta, taxonomy
Earthworms belonging to the family Megascolecidae are the most important and widely distributed in China. Hunan Province (24°38'–30°08'N, 108°47'–114°15'E) is located in the south of China and has a subtropical monsoon climate. Anhui Province (29°41'–34°38'N, 114°54'–119°37'E) is located in the east of China, a transitional region between the warm temperate zone and the subtropical zone. Both provinces are among the most abundant biological diversity areas in China, yet only a few earthworm species have been reported there. Previously, only five (Amynthas moniliatus moniliatus (Chen, 1946), A. triastriatus triastriatus (Chen, 1946), Metaphire biforatum Tan & Zhong, 1987, M. bifoliolare Tan & Zhong, 1987 and M. hunanensis Tan & Zhong, 1986) and eight (A. carnosus carnosus (Goto & Hatai, 1899), A. corticis (Kinberg, 1867), A. hupeiensis (Michaelsen, 1895), A. loti (Chen & Hsu, 1975), A. pectieniferus (Michaelsen, 1931), A. robustus (Perrier, 1872), M. guillelmi (Michaelsen, 1895) and M. tschiliensis tschiliensis (Michaelsen, 1928)) Megascolecidae species have been recorded from Hunan and Anhui, respectively (
In 2015, 2016, and 2019, we investigated earthworm diversity in the two provinces and more than 31 and 24 Megascolecidae species were recorded from Hunan and Anhui, respectively. In this paper, we describe three new species of the genus Amynthas and one new species of the genus Metaphire found in those surveys (Fig.
The earthworms were collected in 2015, 2016, and 2019. Specimens were anaesthetized in a 10% ethanol solution and preserved in a 95% ethanol solution. Holotypes and paratypes are deposited in the Shanghai Natural History Museum.
DNA was extracted from several specimens of A. xiangtanensis sp. nov., A. taoyuanensis sp. nov., A. xuanchengensis sp. nov., M. donganensis sp. nov. by using the E.Z.N.A. Mollusc DNA Kit (Omega Bio-tek, Norcross, GA, USA). The gene cytochrome c oxidase subunit I (COI) was amplified by polymerase chain reaction (PCR). The PCR amplification mixture (50 μL total) consisted of 1 μL of DNA template, 2 μL of each primer, 35.4 μL double-distilled H2O, and 9.6 μL Trans TaqTM Polymerase High Fidelity containing 0.6 μL TransTaqTM HiFi DNA polymerase, 4 μL 2.5 mM dNTPs and 5 μL 10 × TransTaqTM HiFi Buffer I. Primers used in the research were COI,5’-GGTCAACAAATCATAAAGATATTGG-3’ and 5’-TAAACTTCAGGGTGACCAAAAAATCA-3’ (
Specimens with molecular data used in this study. Some species have no molecular data in GenBank. Abbreviations, HT holotype, PT paratype.
| Species | Species number | GenBank acc. no. |
|---|---|---|
| Amynthas xiangtanensis sp. nov. (HT) | P1CJHUSH190510083 N1-05A | PP497097 |
| Amynthas xiangtanensis sp. nov. (PT) | P1CJHUSH190519808 N5-02 | PP497098 |
| Amynthas xiangtanensis sp. nov. (PT) | P1CJHUSH190526781 N5-01 | PP497099 |
| Amynthas taoyuanensis sp. nov. (HT) | P1CJHUSH190517069 N8-01A | PP497095 |
| Amynthas taoyuanensis sp. nov. (PT) | P1CJHUSH190526781 R8-04 | PP497096 |
| Amynthas xuanchengensis sp. nov. (PT) | P1CJHUSH190521800 Q6-03 | PP497100 |
| Metaphire donganensis sp. nov. (PT) | P1CJHUSH190511779 N9-03 | PP497092 |
| Metaphire donganensis sp. nov. (PT) | P1CJHUSH190512778 N11-01 | PP497094 |
| Metaphire donganensis sp. nov. (PT) | P1CJHUSH190512096 Q3-04 | PP497093 |
| Amynthas corticis (Kingberg, 1867) | HN201035-02 | KF205966 |
| Amynthas maximus Qiu & Dong, 2019 | GX201304-01 | MG450707 |
| Amynthas tortuosus Qiu & Dong, 2019 | GX201306-06 | MG450708 |
| Amynthas stricosus Qiu & Sun, 2012 | HN201104-04 | JX315345 |
| Amynthas homosetus (Chen, 1938) | - | No data in GenBank |
| Amynthas genitalis Qiu & Sun, 2012 | - | No data in GenBank |
| Amynthas recavus Yuan & Jiang, 2019 | YN201109-09 | KF205473 |
| Amynthas endophilus Zhao & Qiu, 2013 | HN201011-03 | KF240560 |
| Amynthas fucatus Zhao & Qiu, 2013 | HN201114-01 | KF151185 |
| Amynthas infuscuatus Jiang & Sun, 2015 | - | No data in GenBank |
| Amynthas zonarius Sun & Qiu, 2015 | HN201114-06 | JQ982486 |
| Amynthas baikmudongensis Hong, 2017 | - | No data in GenBank |
| Metaphire vulgaris agricola (Chen,1930) | - | No data in GenBank |
| Metaphire tschiliensis lanzhouensis (Feng, 1984) | - | No data in GenBank |
| Metaphire viridis Feng & Ma, 1987 | - | No data in GenBank |
| Metaphire ptychosiphona Qiu & Zhong, 1993 | - | No data in GenBank |
| Metaphire sanmingensis Sun & Jiang, 2018 | FJ201008-02 | KY774380 |
Genus Amynthas Kinberg, 1867
Holotype. • 1 clitellate (P1CJHUSH190510083 N1-05A), China, Hunan Province, Xiangtan City (27.98312°N, 112.81616°E), 47 m elevation, brownish yellow soil under shrub in farmland, 10 May 2019, JB Jiang, JL Li and BY Yin. Paratypes. 10 clitellates in total • 2 clitellates (P1CJHUSH190510083 N1-05B), China, Hunan Province, Xiangtan City (27.98312°N, 112.81616°E), 47 m elevation, brownish yellow soil under shrub in farmland, 10 May 2019, JB Jiang, JL Li and BY Yin • 2 clitellates (P1CJHUSH190519808 N5-02), China, Hunan Province, Yongzhou City (25.83566°N, 112.27331°E), 163 m elevation, brown soil under weeds in field, 19 May 2019, JB Jiang, JL Li and Y Wang • 2 clitellates (P1CJHUSH190526781 N5-01), China, Hunan Province, Liling City (27.54671°N, 113.54837°E), 74 m elevation, brown soil under weeds in farmland, 26 May 2019, JB Jiang, JL Li and Y Wang • 2 clitellates (P1CJHUSH190514791 N13-01), China, Hunan Province, Yueyang City (29.07035°N, 113.23038°E), 57 m elevation, yellow soil under vegetable field in farmland, 14 May 2019, Y Dong, YF Qin and YZ Wu • 2 clitellates (AH201612-02), China, Anhui Province, Huangshan City (30.58531°N, 117.87033°E), 506 m elevation, brown soil under vegetable field in farmland, 8 May 2016, JB Jiang, J Sun, Y Dong and Y Zheng.
Size medium to large. Spermathecal pores in 5/6–8/9, separated by 1/3 of body circumference. Male pores in XVIII, separated by 1/3 of body circumference, each on the top of a slightly raised, circular porophore. Spermathecae four pairs in VI–IX, ampulla heart-shaped, duct thick and ~ 1/4 of ampulla. Diverticulum is ~ 2/3 of main pouch (duct and ampulla together), terminal 1/2 dilated into rod-shaped seminal chamber. Intestinal caeca are simple. Prostate glands are well developed.
Yellowish brown dorsal pigmentation, pale yellowish brown ventral pigmentation. Dimensions 110–184 mm by 5.0–7.0 mm at clitellum, segments 111–133. Annulus present on VIII–XIII. The dorsal midline is clearly visible and purplish brown. First dorsal pore of all examined individuals in 9/10. Prostomium 1/2 epilobous. Clitellum annular, taupe, in XIV– XVI, swollen, setae invisible externally, but dorsal pores visible on clitellum. Setae numbering 16–28 at III, 26–32 at V, 40–46 at VIII, 46–54 at XX, 54–58 at XXV; 14–16 between male pores; 8–12 (V), 10–12 (VI), 12–14 (VII) and 14–17 (VIII) between spermathecal pores; setal formula, aa = 1.0–1.2ab, zz = 1.4–2.0zy. Male pores one pair in XVIII, separated by 1/3 of body circumference, each on the top of a slightly raised, circular porophore. A pair of oval medium-sized flat-topped papillae on XVIII, after the setae ring near male pores, the interval is ~ 1/4 of body circumference (Fig.
A. xiangtanensis sp. nov. A ventral view showing spermathecal pores, and male pores B prostate glands C intestinal caeca D spermathecae E male pores of paratypes (P1CJHUSH190519808 N5-02) F male pores of paratypes (P1CJHUSH190514791 N13-01). Abbreviations: sp, spermathecal pores; mp, male pores.
Septa 5/6–7/8 thick and muscular, 10/11–12/13 slightly thickened, 8/9 and 9/10 absent. Gizzard long bucket-shaped, in IX–X. Intestine enlarged distinctly from XV. Intestinal caeca paired in XXVII, extending anteriorly to XXIII, simple, smooth on both sides or sometimes weakly constricted on ventral margin (Fig.
The species is named after its type locality.
Amynthas xiangtanensis sp. nov., with four pairs of spermathecal pores in 5/6–8/9, keys to the Amynthas corticis group (
A comparison of characters of A. xiangtanensis sp. nov., A. taoyuanensis sp. nov., and similar species of the Amynthas corticis group. Abbreviations: sp, spermathecal pores, mp, male pores.
| Character | A. xiangtanensis | A. taoyuanensis | A. corticis | A. maximus | A. tortuosus | A. stricosus | A. homosetus | A. genitalis | A. recavus |
|---|---|---|---|---|---|---|---|---|---|
| Body size (mm) | 110–184*5.0–7.0 | 41–120*3.5–4.5 | 45–170*3.0–6.0 | 145–170*5.8–6.2 | 55–86*2.5–2.8 | 72–97*2–2.8 | 116*5.2 | 83–97*2.3–2.5 | 58-64*2.1–2.3 |
| Pigment dorsum | Yellowish brown | None | Greenish brown | Pale purple-brown to brown | Purple-brown to pale purple-brown | None | Dark chocolate to grey | None | Pink to pale brown |
| Pigment ventrum | Pale yellowish brown | None | None | None to yellowish | Pale purple-brown to None | None | Grey | None | None |
| First dorsal pore | 9/10 | 10/11 | 11/12 | 13/14 | 13/14 | 11/12 or 12/13 | 12/13 | 12/13 | 12/13 |
| Clitellum | XIV–XVI | XIV– 2/3XVI | XIV–XVI | 2/5XIV–XVI | XIV–XVI | XIV–XVI | XIV–XVI | XIV–XVI | XIV–XVI |
| Setae numbering | 16–28/III, 40–46/VIII, 46–54/XX | 32–42/III, 44–50/VIII, 50–58/XX | 36–40/VII, 40–46/XXV | 33–38/III, 29–33/VIII, 18–22/XX | 24–26/III, 34–36/VIII, 32–36/XX | 30–54/III, 62–72/VIII, 40–70/XX | 44/VIII | 30–36/III, 32–36/VIII, 38–46/XX | 21–22/III, 34–36/VIII, 34–36/XX |
| Setae number between sp | 14–17 (VIII) | 19–22 (VIII) | 12 (VIII) | 18–22 (VIII) | 12–13 (VIII) | 23–29 (VIII) | - | 8–11 (VIII) | 12 (VIII) |
| Setae number between mp | 14–16 (XVIII) | 13–14 (XVIII) | 10–14 (XVIII) | 9–13 (XVIII) | 8–9 (XVIII) | 10–12 (XVIII) | 9(XVIII) | 11–12 (XVIII) | 9–10 (XVIII) |
| Ventral distance of sp | 1/3C | 1/3C | 1/3C | 1/3C | 1/4C | 2/5C | 1/4C | 1/3C | 2/5C |
| Papillae within sp region | Paired before setae in VIII or IX | Paired after setae in VI, VII or VIII | Paired before or after setae near sp | Two pairs after setae in VII and VIII | Four pairs after setae in VI–IX | None | Invisible | None | None |
| Papillae within mp region | Two or more near male pore | Two paired or more in XVI, XVII, XVIII, or XIX | One or more papillae near male pore | Paired before setae medial of male pore | Paired medial of male pore | Single or paired in XVII, XIX, and XX | Invisible | 4 paired in XVII, XVIII and XIX | Paired in XVII, 3 papillae in XIX |
| Prostate glands | Well developed with accessory gland | Degenerated with accessory glands | Developed | Underdeveloped | Developed | Developed, | Developed | Developed | Developed |
| Diverticulum | 2/3 of main pouch, terminal 1/2 dilated into rod-shaped seminal chamber | 2/3 of main pouch, terminal 2/3 dilated into bag-shaped seminal chamber | Shorter than main pouch and terminal dilated into round or elongate oval seminal chamber | Shorter, lightly twist in middle, terminal 2/5, dilated into rod-shaped seminal chamber | Shorter, terminal 4/5 dilated into S-shaped twisted seminal chamber | As long as main pouch, slender, terminal 2/5 dilated into a band shaped chamber | Shorter, terminal dilated into rod-shaped seminal chamber | Longer than main pouch, terminal 0.29 dilated into rod-shaped seminal chamber | ~ 3/5 of main pouch, terminal 1/4 dilated into ovoid-shaped seminal chamber |
| Accessory glands | Paired on VIII or IX | Two paired on VII or VIII | Bound down to parietes or retained within body wall | One or two on VI, VII, VIII, and IX | One near the each spermatheca | None | Invisible | None | None |
The new species is also fairly close to Amynthas corticis (Kinberg, 1867) by having medium to large size, spermathecal pores location and simple intestinal caeca (Table
Another similar species with four pairs of spermathecal pores in 5/6–8/9 is Amynthas maximus Qiu & Dong, 2019. The two species share some similarities, such as body size, pigmentation, spermathecal pores, simple intestinal caeca (Table
Regarding the pigmentation, clitellum, the position of spermathecal and male pores, simple intestinal caeca, and characteristics of the spermathecae, the new species is similar to Amynthas tortuosus Qiu & Dong, 2019. However, the two species are distinguished by body size, the position of the first dorsal pore, setae number, the number and position of papillae within the spermathecal pore and male pore region, the characteristics and ventral distance of male pores, the position of prostate glands and the existence or nonexistence of accessory glands, and the characteristics of the diverticulum and accessory glands (Table
In addition, the new species is somewhat similar to Amynthas stricosus Qiu & Sun, 2012 in the clitellum, the position of spermathecal and male pores, the ventral distance of male pores, and the simple intestinal caeca. Nevertheless, the two species are different in body size, pigmentation, the position of the first dorsal pore, setae number, the ventral distance of spermathecal pores and the existence or nonexistence of papillae, the characteristics of male pores, the number and position of papillae within the male pore region, the position of prostate glands and the existence or nonexistence of accessory glands, the characteristics of spermathecae, diverticulum and existence or nonexistence of accessory glands (Table
Holotype. • 1 clitellate (P1CJHUSH190517069 N8-01A), China, Hunan Province, Yongzhou City (25.30170°N, 111.63432°E), 201 m elevation, brown soil under weeds in dry farmland, 17 May 2019, JB Jiang, JL Li and BY Yin. Paratypes. 7 clitellates in total • 3 clitellates (P1CJHUSH190517069 N8-01B), China, Hunan Province, Yongzhou City (25.30170°N, 111.63432°E), 201 m elevation, brown soil under weeds in dry farmland, 17 May 2019, JB Jiang, JL Li and BY Yin • 1 clitellates (P1CJHUSH190526781 R8-04), China, Hunan Province, Liling City (27.63491°N, 113.54768°E), 55 m elevation, yellow soil under weeds in woodland, 26 May 2019, JB Jiang, JL Li and Y Wang • 2 clitellates (HU201601-03), China, Hunan Province, Changde City (28.70779°N, 111.38225°E), 66 m elevation, brown soil under moss and fern in farmland, 2 May 2016, JB Jiang, J Sun, Y Dong and Y Zheng • 1 clitellates (HU201602-04), China, Hunan Province, Changde City (28.65267°N, 111.34672°E), 69 m elevation, reddish brown soil under moss and fern in farmland, 2 May 2016, JB Jiang, J Sun, Y Dong and Y Zheng.
Size small to medium. Spermathecal pores in 5/6–8/9, separated by 1/3 of body circumference. Male pores in XVIII, separated by 1/3 of body circumference, each on the top of a slightly raised, circular porophore. Spermathecae four pairs in VI–IX, ampulla heart- or rod-shaped, duct is thick and ~ 1/3 of ampulla. Diverticulum is ~ 3/4 of main pouch (duct and ampulla together), terminal 3/4 dilated into bag-shaped chamber intestinal caeca are simple. Prostate glands are degenerated.
No pigmentation on dorsal and ventral. Dimensions 41–120 mm by 3.5–4.5 mm at clitellum, segments 49–112. Annulus present on X–XIII. The dorsal midline is clearly visible and green-brown. First dorsal pore of all examined individuals in 10/11. Prostomium 1/2 epilobous. Clitellum annular, grey, in XIV–2/3XVI, smooth, setae numbering 8 at XIV (P1CJHUSH190517069 N8-01 and HU201602-04), 11 at XV (HU201602-04), 13 at XVI (P1CJHUSH190517069 N8-01) or 14 at XVI (P1CJHUSH190526781 R8-04), dorsal pores visible on clitellum. Setae numbering 32–42 at III, 40–48 at V, 44–50 at VIII, 50–58 at XX, 56–62 at XXV; 13–14 between male pores; 18–19 (V), 18–21 (VI), 19–21 (VII) and 19–22 (VIII) between spermathecal pores; setal formula, aa = 1.0–1.4ab, zz = 1.0–1.4zy. Male pores one pair in XVIII, separated by 1/3 body circumference, each on the top of a slightly raised, circular porophore. Two pairs of circular medium-sized flat-topped papillae present after the setae ring in XVI and XVII, the interval is ~ 1/3 of body circumference. Two pairs of circular medium-sized flat-topped papillae present before the setae ring on XVII and XVIII, the interval is ~ 1/9 of body circumference (Fig.
Septa 5/6–7/8 thick and muscular, 10/11–12/13 slightly thickened, 8/9 and 9/10 absent. Gizzard spherical, in IX–X. Intestine enlarged distinctly from XV. Intestinal caeca paired in XXVII, extending anteriorly to XXIV, simple, smooth on both sides (Fig.
The species is named after its type locality.
Amynthas taoyuanensis sp. nov., with four pairs of spermathecal pores in 5/6–8/9, also belongs to the Amynthas corticis group. Amynthas taoyuanensis sp. nov. is close to Amynthas xiangtanensis sp. nov. in setae number, the position and characteristics of spermathecal pores and male pores, and the simple intestinal caeca. However, the differences between the two new species are body size, pigmentation, the position of the first dorsal pore, the position of clitellum, the position of papillae within spermathecal pores and male pore region, the existence or nonexistence of prostate glands, and the characteristics of the spermathecae and diverticulum (Table
The new species is similar to Amynthas homosetus (Chen, 1938) in terms of body size, position and characteristics of spermathecal pores and male pores, and simple intestinal caeca. Whereas, the new species differs from A. homosetus in being unpigmented, the first dorsal pore in 10/11, clitellum in XIV–2/3XVI, 13 or 14 setae between male pores, the ventral distance of spermathecal pores is 1/3C, papillae paired after setae in VI, VII, or VIII, two paired or more papillae in XVI, XVII, XVIII, or XIX, prostate glands degenerated, spermathecae ~ 0.6–1.8 mm long, ampulla oval-shaped, duct is thick and ~ 1/2 of ampulla, diverticulum terminal 3/4 dilated into bag-shaped seminal chamber, two paired large lumpy accessory glands on VII or VIII. Amynthas homosetus is dark chocolate-colored anteriorly and grey on other parts of dorsum, grey on the ventrum, the first dorsal pore in 12/13, clitellum without setae in XIV–XVI, nine setae between male pores, the ventral distance of spermathecal pores is 1/4C, papillae invisible within spermathecal pores and male pore regions, prostate glands developed in XVI–XXI, spermathecae heart-shaped, diverticulum seminal chamber ovoid (Table
Amynthas taoyuanensis sp. nov. is also close to Amynthas stricosus Qiu & Sun, 2012 by body size, pigmentation, setae number, the characteristics of spermathecal pores and male pores, the ventral distance of male pores, simple intestinal caeca. However, the new species is distinguished from A. stricosus by the first dorsal pore, the position and existence or nonexistence setae of clitellum, and the ventral distance of spermathecal pores; A. taoyuanensis sp. nov. has paired papillae after setae in VI, VII, or VIII, but A. stricosus has no papillae; in addition, A. taoyuanensis sp. nov. has two paired papillae or more in XVI, XVII, XVIII, or XIX, but A. stricosus has a single or paired after setae in XVII, XIX, and XX; prostate glands of A. taoyuanensis sp. nov. are degenerated with several lumpy accessory glands in XVI, XVII, and XVIII, while developed in XVI–XX with accessory glands invisible in A. stricosus; furthermore, spermathecae of A. taoyuanensis sp. nov. are ~ 0.6–1.8 mm long, ampulla oval-shaped, duct is thick and ~ 1/2 of ampulla, whereas ~ 1.6 mm long, ampulla heart-shaped, gradually slender duct as long as ampulla in A. stricosus; diverticulum of A. taoyuanensis sp. nov. is ~ 3/4 of main pouch, terminal 3/4 dilated into bag-shaped seminal chamber, but as long as main pouch, slender, terminal 2/5 dilated into band-shaped seminal chamber in A. stricosus; two paired large lumpy accessory glands on VII or VIII in A. taoyuanensis sp. nov., but no accessory glands in A. stricosus (Table
According to body size, pigmentation, setae ventrally in clitellum, the position and characteristics of spermathecal pores and male pores, simple intestinal caeca, and the characteristics of spermathecae, A. taoyuanensis sp. nov. is somewhat similar to Amynthas genitalis Qiu & Sun, 2012. However, the new species is characterized by the first dorsal in 10/11, clitellum in XIV–2/3 XVI, 19–22 setae between spermathecal pores (VIII), papillae paired within spermathecal pore region, two paired or more papillae in XVI, XVII, XVIII, or XIX, prostate glands degenerated with several lumpy accessory glands in XVI, XVII, and XVIII, diverticulum ~ 3/4 of main pouch, terminal 3/4 dilated into bag-shaped seminal chamber, two paired large lumpy accessory glands on VII or VIII; whereas A. genitalis has 8–11 setae between spermathecal pores (VIII), no papillae within spermathecal pore region, paired papillae before setae annulet in XVIII and XIX, after setae annulet in XVII and XVII, prostate glands developed with accessory glands invisible in XVII–XX, diverticulum longer than main pouch, slender, terminal 0.29 dilated into rod-shaped seminal chamber, no accessory glands (Table
Another similar species with four pairs of spermathecal pores in 5/6–8/9 is Amynthas recavus Yuan & Jiang, 2019. The two species share some similarities, such as the position of male pores and simple intestinal caeca. However, the two species are distinguished by body size, pigmentation, the first dorsal pore position, setae number, the ventral distance of spermathecal pores, existence or nonexistence of papillae within spermathecal pore region, the characteristics of male pores and papillae within male pore region, prostate glands, the characteristics of spermathecae and diverticulum, and the existence or nonexistence of accessory glands (Table
Holotype. • 1 clitellate (AH201517-06), China, Anhui Province, Xuancheng City (30.89694°N, 118.48889°E), 860 m elevation, black sandy soil under shrubbery in front of the house, 11 October 2015, Y Dong, Z Yuan, MS Chen and YL Wang. Paratype. • 1 clitellate (P1CJHUSH190521800 Q6-03), China, Hunan Province, Chenzhou City (26.16809°N, 113.05648°E), 79 m elevation, sandy soil under weeds beside roadways, 21 May 2019, JB Jiang, JL Li and Y Wang.
Size small. Spermathecal pores in 5/6 and 6/7, separated by 1/3 of body circumference. Male pores in XVIII, separated by 1/4 of body circumference, each on the top of a slightly raised, circular porophore. Spermathecae two pairs in VI and VII, ampulla heart-shaped, duct is thick and ~ 1/2 of ampulla. Diverticulum as long as main pouch (duct and ampulla together), terminal 1/2 dilated into ovoid-shaped seminal chamber. Intestinal caeca are simple. Prostate glands are developed.
No dorsal or ventral pigmentation. Dimensions 26–32 mm by 1.5–2.0 mm at clitellum, segments 60–65. The dorsal midline is not clearly visible. First dorsal pore of all examined individuals in 12/13. Prostomium 1/2 epilobous. Clitellum annular, taupe, in XIV– XVI, smooth, setae invisible externally. Setae numbering 32–40 at III, 36–44 at V, 40–48 at VIII, 50–54 at XX, 56–58 at XXV; 4 between male pores; setal formula, aa = 1.0–1.4ab, zz = 1.4–2.0zy. Male pores one pair in XVIII, separated by 1/4 body circumference, each on the top of a slightly raised, circular porophore (Fig.
Septa 5/6–7/8 thick and muscular, 10/11–12/13 slightly thickened, 8/9 and 9/10 absent. Gizzard spherical in IX–X. Intestine enlarged distinctly from XV. Intestinal caeca paired in XXVII, extending anteriorly to XXVI, simple, smooth on both sides (Fig.
The species is named after its type locality.
Amynthas xuanchengensis sp. nov., with two pairs of spermathecal pores in 5/6 and 6/7, can be assigned to Amynthas morrisi group (
A comparison of characters of A. xuanchengensis sp. nov. and similar species of the Amynthas morrisi group. Abbreviations: sp, spermathecal pores, mp, male pores.
| Character | A. xuanchengensis | A. endophilus | A. fucatus | A. infuscuatus | A. zonarius | A. baikmudongensis |
|---|---|---|---|---|---|---|
| Body size (mm) | 26–32*1.5–2.0 | 96*3 | 137*4.0 | 60–78*1.4–1.6 | 52–103*1.6–3.1 | 75*4.3 |
| Pigment dorsum | None | None | Dark red brown to pale | Purple to pale brown | None | None |
| Pigment ventrum | None | None | Pale red brown to pale | None | None | - |
| Setae numbering | 32–40/III, 40–48/VIII, 50–54/XX | 56/III, 72/V, 84/VIII | 44/III, 60/VIII, 40/XX | 46–48/III, 49–52/VIII, 42–46/XX | 38–56/III, 46–52/VIII, 34–54/XX | 24/VII, 53/XX |
| Setae number between mp | 4 (XVIII) | 2 (XVIII) | 8 (XVIII) | 8–10 (XVIII) | 6–12 (XVIII) | 7 (XVIII) |
| Spermathecal pores | Inconspicuous, 1/3C | Obvious, 2/5C | Obvious, 2/5C | Obvious, 1/3C | Inconspicuous, 2/5C | Inconspicuous |
| Male pores | 1/4C | Surrounded by folds, 1/3C | Surrounded by folds, 2/5C | Surrounded by folds, 1/3C | Surrounded by folds, 1/3C | Large circular raised pads in XVIII |
| Papillae within mp region | None | None | Two in the inner side of male pore., another paired in XVIII | Two in the inner side of male pore | One or two in the inner side of male pore, sometimes another two on XVIII | None |
| Prostate glands | Developed | Developed | Well developed | Developed | Developed | Underdeveloped |
| Diverticulum | As long as main pouch, terminal 1/2 dilated into ovoid-shaped seminal chamber | 4/5 of main pouch, terminal 3/5 dilated into zonal seminal chamber | 1/3 of main pouch, terminal 2/5 dilated into virgulate seminal chamber | Slightly longer than main pouch, terminal 1/3 dilated into swollen seminal chamber | Long as main pouch, terminal 1/2 dilated into band-shaped seminal chamber | 1/2 of main pouch, terminal 1/2 dilated into club-shaped seminal chamber |
Another similar species with two pairs of spermathecal pores in 5/6 and 6/7 is Amynthas fucatus Zhao & Qiu, 2013. The two species share some similarities, such as clitellum in XIV–XVI, the position of spermathecal pores and male pores, invisible papillae within spermathecal pore region, and simple intestinal caeca. By contrast, the new species and A. fucatus can be separated on the basis of smaller body size and fewer setae at VIII. In addition, A. xuanchengensis sp. nov. is unpigmented, but dark red brown before clitellum and pale after clitellum in dorsum, pale red brown before clitellum and pale after clitellum in ventrum in A. fucatus; the first dorsal pore of A. xuanchengensis sp. nov. in 12/13, but 11/12 in A. fucatus; the ventral distance of spermathecal pores and male pores are 1/3C and 1/4C in A. xuanchengensis sp. nov., but 2/5C and 2/5C in A. fucatus; spermathecal pores of A. xuanchengensis sp. nov. are inconspicuous, but obvious in A. fucatus; male pores of A. xuanchengensis sp. nov. not surrounded by folds, but A. fucatus surrounded by three to five folds; invisible papillae within male pore region of A. xuanchengensis sp. nov., whereas two papillae on the inner side of male pore, another paired in XVIII in A. fucatus; prostate glands in XVI–XIX of A. xuanchengensis sp. nov., while XV–XXI with accessory glands invisible in A. fucatus; spermathecae ~ 0.7 mm long, ampulla heart-shaped in A. xuanchengensis sp. nov., nevertheless spermathecae longer, ampulla elongated ovoid in A. fucatus; diverticulum is as long as main pouch, terminal 1/2 dilated into ovoid-shaped seminal chamber in A. xuanchengensis sp. nov., but diverticulum ~ 1/3 of main pouch, straight, terminal 3/4 dilated into virgulate seminal chamber in A. fucatus (Table
Amynthas xuanchengensis sp. nov. appears to be closely related to A. infuscuatus Jiang & Sun, 2015 in the combined characters of ventrum pigmentation, first dorsal pore in 12/13, clitellum in XIV–XVI, the position and characteristics of spermathecal pores, invisible papillae within spermathecal pore region, the position of male pores, simple intestinal caeca, and ampulla heart-shaped. Conversely, they still have some differences, such as body size, dorsum pigmentation, and setae number. Amynthas xuanchengensis sp. nov. smaller, without pigment on dorsum, but purple before VIII; pale brown after VIII in A. infuscuatus; four setae between male pores of A. xuanchengensis sp. nov., but 8–10 setae in A. infuscuatus; the ventral distance of male pores is 1/4C in A. xuanchengensis sp. nov., but 1/3C in A. infuscuatus; spermathecal pores of A. xuanchengensis sp. nov. are inconspicuous, but obvious in A. infuscuatus; male pores of A. xuanchengensis sp. nov. surrounded by no folds, but A. infuscuatus surrounded by three or four folds; invisible papillae within male pore region of A. xuanchengensis sp. nov., whereas two papillae on the inner side of male pore in A. infuscuatus; prostate glands in XVI–XIX of A. xuanchengensis sp. nov., while XVI–1/2XX with accessory glands invisible in A. infuscuatus; spermathecae ~ 0.7 mm long, duct is thick and ~ 1/2 of ampulla in A. xuanchengensis sp. nov., but spermathecae longer, duct slender, twice as long as ampulla in A. infuscuatus; diverticulum is as long as main pouch, terminal 1/2 dilated into ovoid-shaped seminal chamber in A. xuanchengensis sp. nov., but diverticulum longer than main pouch, slender, terminal 1/3 dilated into swollen seminal chamber in A. infuscuatus (Table
Amynthas xuanchengensis sp. nov. and Amynthas zonarius Sun & Qiu, 2015 share some common characters in pigmentation, first dorsal pore in 12/13, clitellum in XIV–XVI, the characteristics of spermathecal pores, invisible papillae within spermathecal pore region, the position of male pores, simple intestinal caeca. In contrast, A. xuanchengensis sp. nov. differs from A. zonarius in smaller body size and fewer setae. Additionally, the ventral distance of spermathecal pore and male pores are 1/3C and 1/4C in A. xuanchengensis sp. nov., but 2/5C and 1/3C in A. zonarius; male pores of A. xuanchengensis sp. nov. surrounded by no folds, but A. zonarius surrounded by five folds; invisible papillae within male pore region of A. xuanchengensis sp. nov. whereas one or two papillae on the inner side of male pore, sometimes another two on XVIII in A. zonarius; prostate glands in XVI–XIX in A. xuanchengensis sp. nov., while XVI–1/2XX with accessory glands invisible in A. zonarius; spermathecae ~ 0.7 mm long, ampulla heart-shaped, duct is thick and ~ 1/2 of ampulla in A. xuanchengensis sp. nov., whereas spermathecae longer, ampulla ovoid, duct as long as ampulla in A. zonarius; ovoid-shaped seminal chamber in A. xuanchengensis sp. nov., but band-shaped seminal chamber in A. zonarius (Table
In terms of pigmentation, clitellum, setae number, the positions of spermathecal pores and male pores, inconspicuous spermathecal pores, invisible papillae within spermathecal pore and male pore regions, simple intestinal caeca. Amynthas xuanchengensis sp. nov. is somewhat similar to Amynthas baikmudongensis Hong, 2017. On the contrary, the new species is easily distinguished from A. baikmudongensis by body size, prostate glands, the characteristics of male pores, spermathecae, and diverticulum (Table
Holotype. • 1 clitellate (HU201613-01A), China, Hunan Province, Dongan City (26.35499°N, 111.19531°E), 172 m elevation, brown soil under vegetable field in farmland, 4 May 2016, JB Jiang, J Sun, Y Dong, and Y Zheng. Paratypes. 13 clitellates in total • 1 clitellate (HU201613-01B), China, Hunan Province, Dongan City (26.35499°N, 111.19531°E), 172 m elevation, brown soil under vegetable field in farmland, 4 May 2016, JB Jiang, J Sun, Y Dong and Y Zheng • 2 clitellates (P1CJHUSH190511779 N9-03), China, Hunan Province, Liuyang City (28.32795°N, 113.52008°E), 112 m elevation, red soil under vegetable in vegetable garden, 11 May 2019, Y Dong, YF Qin and YZ Wu • 7 clitellates (P1CJHUSH190512778 N11-01), China, Hunan Province, Yueyang City (28.91995°N, 113.70132°E), 179 m elevation, brown soil under rape in rape field, 12 May 2019, Y Dong, YF Qin and YZ Wu • 1 clitellate (P1CJHUSH190512096 Q3-04), China, Hunan Province, Linxiang City (29.33550°N, 113.40176°E), 51 m elevation, brown sandy soil under shrub in grove, 12 May 2019, Y Dong, YF Qin and YZ Wu • 2 clitellates (P1CJHUSH190518092 Q6-01), China, Hunan Province, Yuanjiang City (29.04369°N, 112.29798°E), 48 m elevation, brown soil under litter next to the house, 18 May 2019, Y Dong, YF Qin and YK Li.
Size medium to large. Spermathecal pores in 6/7–8/9, separated by 1/3 of body circumference. Male pores in XVIII, separated by 1/3 of body circumference, each on the bottom center of the longitudinal copulatory chamber. Sometimes the copulatory chamber eversion is ridged. Spermathecae three pairs in VII–IX, ampulla heart- or rod-shaped, duct thick and as long as ampulla. Diverticulum as long as main pouch (duct and ampulla together), slender and straight at proximal part, terminal 1/2 dilated into twisted in zigzag fashion. Intestinal caeca are simple. Prostate glands are well developed.
Pale brown dorsal and ventral pigmentation. Dimensions 72–159 mm by 4.6–7.0 mm at clitellum, segments 78–111. Annulus present on IX–XVIII. The dorsal midline is clearly visible and purplish brown. First dorsal pore of all examined individuals in 12/13. Prostomium 1/2 epilobous. Clitellum annular, pale taupe, in XIV– XVI, smooth, setae invisible externally. Setae numbering 20–34 at III, 20–46 at V, 28–58 at VIII, 52–66 at XX, 58–70 at XXV; 8–13 between male pores; 14–17 (VI), 16–20 (VII) and 17–21 (VIII) between spermathecal pores; setal formula, aa = 1.0–1.6ab, zz = 1.2–1.8zy. Male pores one pair in XVIII, separated by 1/3 body circumference, each on the bottom center of the longitudinal copulatory chamber, multiple radioactive folds on the outer edge (Fig.
Septa 5/6–7/8 thick and muscular, 10/11–12/13 slightly thickened, 8/9 and 9/10 absent. Gizzard spherical, in IX–X. Intestine enlarged distinctly from XV. Intestinal caeca paired in XXVII, extending anteriorly to XXIII, simple, smooth dorsal margin, weakly constricted on ventral margin (Fig.
The species is named after its type locality.
Metaphire donganensis sp. nov. with three pairs spermathecal pores in 6/7–8/9, keys to the Metaphire houlleti group, which includes 44 species (
A comparison of characters of M. donganensis sp. nov. and similar species of the Metaphire houlleti group. Abbreviations: sp, spermathecal pores, mp, male pores.
| Character | M. donganensis | M. vulgaris agricola | M. tschiliensis lanzhouensis | M. viridis | M. ptychosiphona | M. sanmingensis |
|---|---|---|---|---|---|---|
| Body size (mm) | 72–159*4.6–7.0 | 118*6.0 | 245–310*6–7 | 192–230*9.5–10 | 196–295*6.0–9.0 | 55–113* 4–5.5 |
| Pigment dorsum | Pale brown | Earthy yellow | Earthy yellow | Dark green | Grey-brown | Pale brown to brown |
| Pigment ventrum | Pale brown | Earthy yellow | Earthy yellow | Pale green | Grey-brown | None |
| First dorsal pore | 12/13 | 12/13 | 12/13 | 12/13 | 11/12 | 11/12, 12/13 or 13/14 |
| Setae numbering | 20–34/III, 28–58/VIII, 52–66/XX | 29/III, 54/VIII, 62/XXV | 32–40/III, 46–55/VIII | 47–50/III, 64–67/VIII, 92–95/XXV | 57–61III, 64–79/VIII, 87–117/XX | 16–24/III, 33–40/VIII, 44–48/XX |
| Setae number between sp | 17–21 (VIII) | - | 16–24 (VIII) | - | 28–34 (VIII) | 12 (VIII) |
| Setae number between mp | 8–13 (XVIII) | - | 8–14 (XVIII) | - | 15–27 (XVIII) | 8–9 (XVIII) |
| Ventral distance of sp | 1/3C | 1/4C | 1/3C | 1/2C | 2/5C | 1/3C |
| Papillae within sp region | None | Paired in VII | None | Paired in VII, VIII, and IX | None | Two in VII and VIII, or extra two paired in VIII |
| Ventral distance of mp | 1/3C | 1/4C | 1/3C | 1/2C | 1/3C | 1/3C |
| Papillae within mp region | None | Paired in the inner side of male pore | One in pouches | Four in pouches | None | Three in the inner side of male pores, extra three in XVIII |
| Prostate glands | Well developed | Developed | Underdeveloped with accessory gland | Developed with accessory gland | Well developed | Developed |
| Diverticulum | As long as main pouch, terminal 1/2 dilated into zigzag fashion | Shorter, terminal 2/3 dilated into twisted zigzag fashion | As long as main pouch, terminal 2/3 dilated into zigzag fashion | Longer, terminal 1/2 dilated into zigzag fashion | Shorter, terminal 0.6 dilated into zigzag fashion | Shorter, terminal dilated into rod-shaped seminal chamber |
| Accessory glands | None | Paired in VIII | None | Paired in VII, VIII, and IX | None | Invisible |
We compare the new species to Metaphire tschiliensis lanzhouensis (Feng, 1984), which has three pairs of spermathecal pores in 6/7–8/9. They share several common characters in the first dorsal pore position, clitellum in XIV–XVI, setae number, the position of spermathecal pores, the position and characteristics of male pores, no papillae within spermathecal pore region, simple intestinal caeca, and the characteristics of the diverticulum. Quite the contrary, the new species is smaller than M. tschiliensis lanzhouensis. Beyond that, coloration is pale brown in M. donganensis sp. nov., earthy yellow in M. tschiliensis lanzhouensis; spermathecal pores are obvious in M. donganensis sp. nov., but inconspicuous in M. tschiliensis lanzhouensis; no papillae within male pore region of M. donganensis sp. nov., but one in pouch of M. tschiliensis lanzhouensis; prostate glands well developed in XVI–XXII of M. donganensis sp. nov., while underdeveloped in XVII–XIX with a large lumpy accessory gland in M. tschiliensis lanzhouensis; spermathecae ~ 3.2–8.0 mm long, ampulla heart- or rod-shaped of M. donganensis sp. nov., but spermathecae ~ 4 mm long, ampulla spherical-shaped in M. tschiliensis lanzhouensis (Table
Considering the three pairs of spermathecal pores in 6/7–8/9, we compared the new species with Metaphire viridis Feng & Ma, 1987. Both have first dorsal pores in 12/13, clitellum in XIV–XVI, the same position and characteristics of spermathecal pores, simple intestinal caeca, and the same characteristics of diverticulum. However, the new species differs from M. viridis in many respects. Metaphire donganensis sp. nov. is smaller than M. viridis with pale brown coloration; setae number of M. donganensis sp. nov. is less than M. viridis; the ventral distance of spermathecal pore and male pores are 1/3C and 1/3 C in M. donganensis sp. nov., but 1/2C and 1/2C in M. viridis; no papillae within spermathecal pore region of M. donganensis sp. nov., while paired in VII, VIII, and IX of M. viridis; no papillae within male pore region of M. donganensis sp. nov., but four in pouches in M. viridis; prostate glands in XVI–XXII of M. donganensis sp. nov., while XV–XX or XVII–XIX with a lumpy accessory gland in M. viridis; no accessory glands in M. donganensis sp. nov., whereas paired in the VII, VIII and IX of M. viridis; spermathecae ~ 3.2–8.0 mm long, ampulla heart- or rod-shaped of M. donganensis sp. nov., nevertheless spermathecae ~ 4 mm long, ampulla spherical- or oval-shaped in M. viridis; no accessory glands of M. donganensis sp. nov., while paired in VII, VIII, and IX of M. viridis (Table
We also compare the new species with Metaphire ptychosiphona Qiu & Zhong, 1993 on the basis of the position and characteristics of male pores, no papillae within spermathecal pore and male pore regions, simple intestinal caeca, and the same characteristics of the diverticulum. Nevertheless, the new species is smaller than M. ptychosiphona. In addition, pale brown in M. donganensis sp. nov., but grey brown in M. ptychosiphona; first dorsal pore in 12/13 of M. donganensis sp. nov., while in 11/12 of M. ptychosiphona; setae number of M. donganensis sp. nov. less than M. ptychosiphona; the ventral distance of spermathecal pore is 1/3C in M. donganensis sp. nov., but 2/5C in M. ptychosiphona; spermathecal pores obvious in M. donganensis sp. nov., but inconspicuous in M. ptychosiphona; prostate glands in XVI–XXII of M. donganensis sp. nov., while XVI–XVIII or XVII–XIX in M. ptychosiphona; and spermathecae ~ 3.2–8.0 mm long, ampulla heart- or rod-shaped, duct swollen, as long as ampulla in M. donganensis sp. nov., nevertheless spermathecae shorter, ampulla rod-shaped, duct shorter than ampulla in M. ptychosiphona (Table
The new species is also close to Metaphire sanmingensis Sun & Jiang, 2018 in body size, clitellum in XIV–XVI, the position of spermathecal pores and male pores, simple intestinal caeca, prostate glands, and the characteristics of the ampulla and diverticulum. However, the new species is easily distinguished from M. sanmingensis in pigmentation, first dorsal pore position, setae number, papillae within spermathecal pore region, and the characteristics of male pores and papillae within male pore region (Table
The COI mitochondrial DNA gene, considered as a barcode for earthworm identification (
| S1 HT | S1 PT | S1 PT | S2 HT | S2 PT | S3 PT | S4 PT | S4 PT | S4 PT | A. corticis | A. maximus | A. tortuosus | A. stricosus | A. recavus | A. endophilus | A. fucatus | A. zonarius | M. sanmingensis | |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| S1 HT | ||||||||||||||||||
| S1 PT | 0.00% | |||||||||||||||||
| S1 PT | 0.16% | 0.16% | ||||||||||||||||
| S2 HT | 21.78% | 21.78% | 21.55% | |||||||||||||||
| S2 PT | 21.78% | 21.78% | 21.55% | 0.00% | ||||||||||||||
| S3 PT | 21.32% | 21.32% | 21.10% | 17.25% | 17.25% | |||||||||||||
| S4 PT | 22.53% | 22.53% | 22.29% | 18.81% | 18.81% | 20.64% | ||||||||||||
| S4 PT | 22.29% | 22.29% | 22.06% | 18.60% | 18.60% | 20.42% | 0.16% | |||||||||||
| S4 PT | 22.27% | 22.27% | 22.04% | 18.82% | 18.82% | 19.95% | 5.62% | 5.44% | ||||||||||
| A. corticis | 23.16% | 23.16% | 22.92% | 20.83% | 20.79% | 23.25% | 20.68% | 20.45% | 19.55% | |||||||||
| A. maximus | 21.05% | 21.05% | 20.79% | 19.13% | 19.09% | 18.25% | 18.16% | 17.91% | 18.38% | 15.54% | ||||||||
| A. tortuosus | 19.78% | 19.78% | 19.55% | 18.75% | 18.71% | 18.78% | 17.05% | 17.26% | 17.25% | 20.18% | 19.18% | |||||||
| A. stricosus | 18.06% | 18.06% | 17.85% | 16.53% | 16.50% | 16.62% | 17.16% | 16.95% | 16.31% | 16.91% | 15.31% | 14.18% | ||||||
| A. recavus | 18.54% | 18.54% | 18.32% | 15.90% | 15.87% | 19.85% | 15.08% | 14.88% | 15.93% | 19.58% | 19.57% | 16.12% | 14.69% | |||||
| A. endophilus | 17.17% | 17.17% | 16.96% | 17.58% | 17.54% | 17.61% | 18.99% | 18.77% | 18.37% | 18.29% | 16.59% | 17.68% | 15.04% | 17.32% | ||||
| A. fucatus | 22.48% | 22.48% | 22.07% | 21.67% | 21.67% | 23.34% | 19.67% | 19.29% | 21.24% | 17.82% | 20.52% | 18.22% | 20.22% | 18.67% | 18.22% | |||
| A. zonarius | 18.04% | 18.04% | 17.82% | 16.00% | 15.97% | 16.79% | 19.05% | 19.27% | 21.04% | 18.43% | 16.02% | 15.61% | 14.38% | 15.72% | 13.66% | 18.73% | ||
| M. sanmingensis | 20.81% | 20.81% | 20.57% | 19.96% | 19.92% | 19.24% | 19.30% | 19.06% | 20.00% | 15.30% | 19.69% | 17.71% | 15.00% | 17.96% | 18.58% | 21.54% | 15.88% |
We are grateful to Prof. Jing Sun, Dr Yan Dong, Dr Zhu Yuan, Dr Yali Wang, Dr Yifeng Qin, Mr Mingsheng Chen, Dr Yizhao Wu, Dr Yingkai Li, Mr Yong Zheng, Mr Bangyi Yin, and Ms Yue Wang for their assistance with the fieldwork. We would like to express our great appreciation to the editors and reviewers.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was supported by National Nature Science Foundation of China Grant No. 42077028, No. 41701272, No. 41471204, and National Science & Technology Fundamental Resources In-vestigation Program of China No. 2018FY100300.
Conceptualization: QJ, JBJ and JPQ. Software: QJ. Formal analysis: QJ. Investigation: JBJ and JLL. Data Curation: QJ. Writing - Original draft: QJ. Writing - Review and Editing: QJ, JBJ and JPQ. Supervision: JBJ and JPQ. Project administration: JBJ and JPQ.
Qing Jin https://orcid.org/0000-0001-7731-3150
AuthorName https://orcid.org/0000-0000-0000-0000
AuthorName https://orcid.org/0000-0000-0000-0000
All of the data that support the findings of this study are available in the main text.