Research Article |
Corresponding author: Yucheng Lin ( linyucheng@scu.edu.cn ) Academic editor: Cristina Rheims
© 2023 Jianshuang Zhang, Chengcheng Feng, Hao Yu, Yucheng Lin.
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:
Zhang J, Feng C, Yu H, Lin Y (2023) A review of the spider genus Sinoalaria (Araneae, Theridiosomatidae), with the descriptions of four new species and two new combinations. ZooKeys 1173: 307-338. https://doi.org/10.3897/zookeys.1173.105123
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The spider genus Sinoalaria Zhao & Li, 2014 is redefined and reviewed. A total of ten species are studied, including four new species: S. chi Yu & Lin, sp. nov. (♂♀), S. shenhei Yu & Lin, sp. nov. (♀), S. shuidi Yu & Lin, sp. nov. (♀), S. xiaotu Yu & Lin, sp. nov. (♂♀). Two new combinations are proposed: Sinoalaria nitida (Zhao & Li, 2012), comb. nov. and S. prolata (Zhao & Li, 2012), comb. nov., both transferred from Karstia Chen, 2010. The material of six known species were re-examined and photographed, including the type species, S. chengguanensis (Zhao & Li, 2012). A key is provided for all species of the genus, as well as diagnoses, illustrations, and a distribution map.
Araneoidea, identification key, Southeast Asia, southern China, morphology
Sinoalaria Zhao & Li, 2014 is a replacement name for Alaria Zhao & Li, 2012, which was originally erected as a monotypic genus based on A. chengguanensis Zhao & Li, 2012 from China. It is a relatively small genus, distributed exclusively in South East Asia, with only four known species described so far (
This paper reports our findings on the study of recently available samples from southwest China, Vietnam, Laos, and Thailand, which revealed four new species of Sinoalaria. Furthermore, two combinations are proposed based on morphological evidence and a preliminary molecular phylogeny (unpublished): S. nitida (Zhao & Li, 2012), comb. nov. and S. prolata (Zhao & Li, 2012), comb. nov., ex Karstia Chen, 2010. With that, the total number of Sinoalaria species reaches ten species. The goals of this paper are: 1) to conduct a comprehensive review of the genus Sinoalaria, provide an identification key and a distribution map for all species; 2) to describe the four new species under the name of S. chi Yu & Lin, sp. nov., S. shenhei Yu & Lin, sp. nov., S. shuidi Yu & Lin, sp. nov. and S. xiaotu Yu & Lin, sp. nov.; 3) to re-illustrate all six known species, including the type species and the two combinations.
All specimens were preserved in 95% ethanol. Specimens were examined and measured with a Leica M205 C stereomicroscope. Further details were studied with an Olympus BX43 compound microscope. Male and female genitalia were examined and illustrated after dissection. Epigynes were removed and cleared in lactic acid before illustration. All vulvae were imaged after being embedded in Arabic gum. Photos were taken with a Canon EOS 60D wide zoom digital camera (8.5 megapixels) mounted on an Olympus BX43 stereomicroscope. The images were montaged using Helicon Focus 3.10 (
The distribution map was generated with ArcGIS 10.5 (Environmental Systems Research Institute, Inc.). Locality coordinates for all species are copied from the original publications (see
All examined materials are deposited in the Natural History Museum of Sichuan University in Chengdu (
Family Theridiosomatidae Simon, 1881
Alaria Zhao & Li, 2012: 7 (type species Alaria chengguanensis Zhao & Li, 2012, by original designation).
Sinoalaria
Zhao & Li, 2014: 41 (replacement name for Alaria Zhao & Li, 2012, preoccupied in Platyhelminthes by
Sinoalaria species can be recognised by the following combination of genitalic characters: In males, palpal tibia retrolaterally bears an apophysis (Figs
See
Ten species from southwestern China to Laos, Vietnam and to Thailand: Sinoalaria bicornis (♂♀) and S. navicularis (♂♀) from Laos, S. chengguanensis (♂♀), S. nitida comb. nov. (♂♀), S. prolata comb. nov. (♂♀), S. shenhei sp. nov. (♀) and S. shuidi sp. nov. (♀) endemic to China, S. xiaotu sp. nov. (♂♀) endemic to Vietnam, S. cavernicola from Laos and Thailand, S. chi sp. nov. (♂♀) from Vietnam and Thailand.
1 | Males | 2 |
– | Females | 7 |
2 | Cymbium dorsal-basally bears a cluster of several long setae (Figs |
3 |
– | Cymbium dorsal-basally without a cluster of several long setae (Figs |
5 |
3 | Embolic bases prominently visible, median apophysis large, > 1/2 of tegulum length (Fig. |
S. chengguanensis |
– | Embolic base indistinct, median apophysis relatively small, ≤ 1/2 of tegulum length (Figs |
4 |
4 | Cymbium basally with a cluster of 6 setae; median apophysis extremely small, ≤ 1/3 of tegulum length, both proximal process and distal process are indistinct; the apex of conductor needle-shaped, sharp (Fig. |
S. xiaotu sp. nov. |
– | Cymbium basally with a cluster of 8 setae; median apophysis comparably larger, ca 1/2 of tegulum length, both proximal process and distal process are distinct; the apex of conductor nearly triangular, relatively blunt (Fig. |
S. bicornis |
5 | Median apophysis navicular, proximal process sharp and not serrated, distal process not furcated (Fig. |
S. navicularis |
– | Median apophysis not navicular, proximal process with a blunt and serrated tip, apex of distal process slightly furcated (Figs |
6 |
6 | Distal process of median apophysis short and wide, slightly shorter and narrower than base of median apophysis; the lower ramus on distal process of median apophysis tooth-shaped, apex sharp, distinctly longer than the indistinct upper ramus; embolus distinctly long, terminating at ca 8 o’clock position in retrolateral view, terminating at ca 4 o’clock position in ventral view; embolic base relatively smaller, its width ca 1/2 of tegulum length (Fig. |
S. chi sp. nov. |
– | Distal process of median apophysis long and narrow, slightly longer and distinctly narrower than base of median apophysis; both rami on distal process of median apophysis distinct, are of equal length, the lower one with a relatively blunt tip; embolus relatively short, terminating at ca 10 o’clock position both in retrolateral and ventral view; embolic base relatively larger, wider than 2/3 of tegulum length (Fig. |
S. cavernicola |
7 | Scape shorter than length of epigynal plate (from anterior level of vulva to posterior margin of epigynal plate) (Figs |
8 |
– | Scape longer than length of epigynal plate (Figs |
9 |
8 | Scape shaped like a water drop, apex distinctly wider than its stem; spermathecae globular (Fig. |
S. shuidi sp. nov. |
– | Scape shaped like a nose, apex nearly as wide as stem; spermathecae peanut-shaped (Fig. |
S. nitida |
9 | Scape rugose, membranous except slightly sclerotized apex (Figs |
10 |
– | Scape not rugose, heavily sclerotized (Figs |
13 |
10 | Scape wide, ca 1/2 width of epigynal plate, apex nearly triangular (Fig. |
S. navicularis |
– | Scape narrow, ca 1/4–1/3 width of epigynal plate, apex nearly digitiform (Figs |
11 |
11 | Scape apically with 2 notches, ventral and anterior folds of copulatory ducts slightly sclerotized (Fig. |
S. bicornis |
– | Scape apically with a hood, ventral and anterior folds of copulatory ducts completely membranous (Figs |
12 |
12 | The ventral and anterior folds of copulatory ducts represented by 2 oblong bursae, the dorsal and posterior folds represented by a longitudinal loop and a horizontal loop (Fig. |
S. chi sp. nov. |
– | The ventral and anterior folds represented by 2 globular bursae, the dorsal and posterior folds running horizontally, forming only 1 loop (Fig. |
S. cavernicola |
13 | Scape wide, triangular, protrudes vertically from the posterior epigynal margin; copulatory ducts located laterally to spermathecae (Fig. |
S. prolata |
– | Scape not as above; copulatory ducts located anterolaterally to spermathecae (Figs |
14 |
14 | Scape shaped like a dumbbell, apically swollen (Fig. |
S. xiaotu sp. nov. |
– | Scape tongue-shaped or like an inverted bowling pin, apically narrowed (Figs |
15 |
15 | Scape shaped like an inverted bowling pin, slightly narrowed proximally; the dorsal and posterior folds of copulatory ducts trapeziform, heavily sclerotized; dorsum of abdomen basically black, with three bands which consisting of small white spots, forming a trident-shaped pattern (Fig. |
S. shenhei sp. nov. |
– | Scape tongue-shaped, proximally part distinctly narrowed; the dorsal and posterior folds of copulatory ducts nearly circular, slightly sclerotized; abdomen dorsally white with numerous small black spots (Fig. |
S. chengguanensis |
Alaria bicornis Lin, Li & Jäger, 2014: 90, figs 11A–D, 12A–F, 13A–E, 14A–C, 15A–F, 16A–C (♂♀).
Sinoalaria bicornis:
2♂ 6♀, Laos: Vien Tiane Province, Vang Vieng District: 13.2 km north of Vieng keo Village, Tham Hoy, 19°02.352'N, 102°25.422'E, 256 m, 3.XII.2012, Z. Yao and S. Li leg.; 4♂ 12♀, 11.95 km north of Vieng keo Village, Pha Thao Cave, 19°01.752'N, 102°25.956'E, 290 m, 3.XII.2012, Z. Yao and S. Li leg.; 3♂ 8♀, 10.37 km north of Vieng keo Village, Kieo Cave, 19°00.882'N, 102°25.902'E, 286 m, 2.XII.2012, Z. Yao and S. Li leg.; 1♂ 7♀, 4.01 km north of Vieng keo Village, Lom Cave, 18°57.456'N, 102°26.244'E, 314 m, 2.XII.2012, Z. Yao and S. Li leg.
Males of S. bicornis resemble those of S. xiaotu sp. nov. in the general shape of the male palp. The palps of the two species share the similarly short median apophysis which is ≤ 1/2 of tegulum length, and the indistinct embolic base (Figs
Male palp of Sinoalaria bicornis (Lin, Li & Jäger, 2014) A prolateral view B retrolateral view C dorsal view D ventral view. Abbreviations: Co = conductor; Cy = cymbium; EB = embolic base; Em = embolus; MA = median apophysis; Pc = paracymbium; RTA = retrolateral tibial apophysis; SCy = setae on cymbium; Te = tegulum. Scale bars: 0.20 mm.
Sinoalaria bicornis (Lin, Li & Jäger, 2014), male habitus (A, B), female habitus (C, D) and epigyne (E–G) A dorsal view B ventral view C ventral view D dorsal view E intact, ventral view F cleared, ventral view G cleared, dorsal view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A–D); 0.20 mm (E–G).
See
Laos (Fig.
Alaria cavernicola Lin, Li & Jäger, 2014: 77, figs 1A–H, 2A–E, 3A–G, 4A, B (♂♀).
Sinoalaria cavernicola:
1♂ 10♀, Laos: Ban Kouanphavang Khammouane Province: 30.02 km northeast of Thakhek Town, Tham Kamouk Cave, 17°37.914'N, 104°07.458'E, 193 m, 24.XI.2012, Z. Yao and S. Li leg.; 2♂ 5♀, Thailand: Satun Province: Thung wa District, Cave without name-cave A & B, 07°03.966'N, 99°50.478'E, 12 m, 1–3.XII.2013, F. Ballarin leg.; 1♂ 8♀, Thung wa District, Cave without name,07°06.276'N, 99°47.502'E, 25 m, 29.XI.2013, F. Ballarin leg.
Males of S. cavernicola and S. chi sp. nov. share the following features: the proximal process of median apophysis with a blunt and serrated tip, and the distal process of median apophysis with a slightly furcated apex (Figs
Male palp of Sinoalaria cavernicola (Lin, Li & Jäger, 2014) A prolateral view B retrolateral view C ventral view D dorsal view. Abbreviations: Co = conductor; Cy = cymbium; EB = embolic base; Em = embolus; MA = median apophysis; Pc = paracymbium; RTA = retrolateral tibial apophysis; Te = tegulum. Scale bars: 0.20 mm.
Sinoalaria cavernicola (Lin, Li & Jäger, 2014), male habitus (A, B), female habitus (C, D) and epigyne (E–G) A dorsal view B ventral view C dorsal view D ventral view E intact, ventral view F cleared, ventral view G cleared, dorsal view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A–D); 0.20 mm (E–G).
See
Laos and Thailand (Fig.
Alaria chengguanensis Zhao & Li, 2012: 8, figs 1A–D, 2A–D, 3A–D, 4A–F, 5A–D (♂♀).
Sinoalaria chengguanensis:
Holotype
♂ and paratypes 13♂ 8♀ (
Males of S. chengguanensis resembles those of S. bicornis and S. xiaotu sp. nov. by their cymbium dorsal-basally bears a cluster of several long setae (Figs
Male palp of Sinoalaria chengguanensis (Zhao & Li, 2012) A prolateral view B retrolateral view C embolic division, dorsal view D ventral view. Abbreviations: Co = conductor; Cy = cymbium; EB = embolic base; Em = embolus; MA = median apophysis; Pc = paracymbium; RTA = retrolateral tibial apophysis; SCy = setae on cymbium; Te = tegulum. Scale bars: 0.20 mm.
Sinoalaria chengguanensis (Zhao & Li, 2012), male habitus (A), female habitus (B, C) and epigyne (D–F) A dorsal view B dorsal view C lateral view D intact, ventral view E cleared, ventral view F cleared, dorsal view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A–C); 0.20 mm (D–F).
See
China (Guizhou) (Fig.
Holotype ♂ and Paratype 1♀, Vietnam: Hang Dai Ca, 20°33.520'N, 105°53.287'E, 30.XII.2013, H. Sterner leg.
Vietnam. 1♂ 2♀, Hang Boi, 20°15.533'N, 105°53.278'E, 24–25.XII.2013, H. Steiner leg.; 1♂ 5♀, Hang Tra Tu, 20°06.492'N, 105°55.040'E, 28.XII.2013, H. Steiner leg.
The specific name is derived from the Chinese pinyin ‘chǐ’, which means ‘tooth’, referring to the tooth-shaped lower ramus on distal process of median apophysis; noun in apposition.
Males of S. chi sp. nov. resemble those of S. cavernicola. The two species share a similarly slender and filiform embolus, sheet-shaped paracymbium with a spine-like tip, and the median apophysis with serrated proximal process and slightly bifurcated distal process but differ by: (1) distal process of median apophysis short and wide, slightly shorter and narrower than the base of median apophysis in S. chi sp. nov. (long and narrow, slightly longer and distinctly narrower than base in S. cavernicola) (cf. Fig.
Male palp of the holotype of Sinoalaria chi sp. nov. A prolateral view B retrolateral view C ventral view D dorsal view. Abbreviations: Co = conductor; Cy = cymbium; EB = embolic base; Em = embolus; MA = median apophysis; Pc = paracymbium; RTA = retrolateral tibial apophysis; Te = tegulum. Scale bars: 0.20 mm.
Sinoalaria chi sp. nov., female paratype and male holotype, male habitus (A, B), female habitus (C, D) and epigyne (E–G) A dorsal view B ventral view C dorsal view D ventral view E intact, ventral view F cleared, ventral view G cleared, dorsal view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A–D); 0.20 mm (E–G).
Male (holotype) (Fig.
Palp
(Fig.
Female (paratype). Somatic features as in Fig.
Epigyne
(Fig.
Northern Vietnam (Fig.
Alaria navicularis Lin, Li & Jäger, 2014: 83, figs 5A–F, 6A–D, 7A–D, 8A–E, 9A–D, 10A–C (♂♀).
Sinoalaria navicularis:
1♂ 4♀, Laos: Khammouan Province, 2.5 km WNW, Ban Tathot, Eingang, 17°37.897'N, 103°07.502'E, 200 m, 20.II.2003, P. Jäger leg.; 4♀, Tham Koun Dôn, 17°333.82'N, 104°52.132'E, 24.II.2006, H. Steiner leg.
Males of S. navicularis can be distinguished from all other congeners by the navicular median apophysis (Fig.
Male palp of Sinoalaria navicularis (Lin, Li & Jäger, 2014) A prolateral view B retrolateral view C ventral view D dorsal view. Abbreviations: Co = conductor; Cy = cymbium; EB = embolic base; Em = embolus; MA = median apophysis; Pc = paracymbium; RTA = retrolateral tibial apophysis; Te = tegulum. Scale bars: 0.20 mm.
Sinoalaria navicularis (Lin, Li & Jäger, 2014), male habitus (A, B), female habitus (C, D) and epigyne (E–G) A dorsal view B ventral view C dorsal view D ventral view E intact, ventral view F cleared, ventral view G cleared, dorsal view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A–D); 0.20 mm (E–G).
See
Laos (Fig.
Karstia nitida Zhao & Li, 2012: 20, figs 11A–E, 12A, B (♀).
Paratype
1♀ (
Sinoalaria nitida morphologically is similar to S. shuidi sp. nov. by the scape shorter than length of epigynal plate, with a swollen apex (Figs
Sinoalaria nitida (Zhao & Li, 2012), comb. nov., female habitus (A, B) and epigyne (C–G) A dorsal view B ventral view C intact, ventral view D cleared, ventral view E cleared, dorsal view F anterior view G posterior view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; N = notch; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A, B); 0.20 mm (C–G).
See
China (Guangxi) (Fig.
Karstia prolata Zhao & Li, 2012: 23, figs 13A–F, 14A, B (♀).
Holotype
♀ (
This species can be distinguished from all other species of the genus Sinoalaria by the wide, triangular, scape that protrudes vertically from the posterior epigynal margin (Fig.
Sinoalaria prolata (Zhao & Li, 2012), comb. nov., female habitus (A–C) and epigyne (D–F) A dorsal view B ventral view C lateral view D intact, ventral view E cleared, posterior view F cleared, anterior view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A–C); 0.20 mm (D–F).
See
China (Guangxi) (Fig.
Holotype ♀, China: Yunnan, Baoshan City, Longling County, Longjiang Town, Xiaoheishan Provincal Nature Reserve, 24°49.733'N, 98°45.601'E, 2020 m, 22.VIII.2018, Y. Lin et al. leg.
11 juv., same data as holotype.
The specific name is an adjective and derived from the Chinese pinyin ‘shēn hēi’, which means ‘pitch-black’, referring to the basic color of body.
The new species is similar to S. chengguanensis (Fig.
Holotype female of Sinoalaria shenhei sp. nov., habitus (A, B) and epigyne (C–F) A dorsal view B ventral view C intact, lateral view D intact, ventral view E cleared, ventral view F cleared, dorsal view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A, B); 0.20 mm (C–F).
Female (holotype) (Fig.
Epigyne
(Fig.
Male. Unknown.
Known only from the type locality (Fig.
Holotype ♀ and Paratypes 3♀, China: Guangxi, Liuzhou City, Luzhai County, Zhongdu Town, Xiamotun Village, Jiulong Cave, 24°44.917'N, 109°39.844'E, 257 m, 15.VII.2013, H. Zhao leg.
The specific name is derived from the Chinese pinyin ‘shuǐ dī’, which means water drop, referring to the scape that is shaped like a water drop; noun in apposition.
This new species is similar to S. nitida in having the scape shorter than length of epigynal plate, with a swollen apex (Figs
Holotype female of Sinoalaria shuidi sp. nov., habitus (A, B) and epigyne (C–E) A dorsal view B ventral view C intact, ventral view D cleared, ventral view E cleared, dorsal view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; N = notch; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A, B); 0.20 mm (C–E).
Female (holotype) (Fig.
Epigyne
(Fig.
Male. Unknown.
Known only from the type locality (Fig.
Holotype ♂ and Paratypes 6♀, Vietnam: Bac Can, Barbie National Park, Beilan cave, 22°22.766'N, 105°36.790'E, 280 m, 2.IV.2012, Z. Yao leg.; 1♂ 6♀, same data as holotype, 18.X.2012, H. Zhao and Z. Chen leg.
1♂ 2♀, Vietnam: Phu Tho, Xuan Son National Park, Lun Cave, 21°07.022'N, 104°57.443'E, 398 m, 26.X.2012, H. Zhao and Z. Chen leg.
The specific name is derived from the Chinese pinyin ‘xiǎo tū’, which means ‘small apophysis’, referring to the small median apophysis which is no more than 1/3 of tegulum length; noun in apposition.
Males of S. xiaotu sp. nov. can be distinguished from congeners except S. bicornis by the similar indistinct embolic base and the small median apophysis which is no more than 1/2 of tegulum length (Figs
Male palp of the holotype of Sinoalaria xiaotu sp. nov. A prolateral view B retrolateral view C ventral view D dorsal view. Abbreviations: Co = conductor; Cy = cymbium; EB = embolic base; Em = embolus; MA = median apophysis; Pc = paracymbium; RTA = retrolateral tibial apophysis; SCy = setae on cymbium; Te = tegulum. Scale bars: 0.20 mm.
Male (holotype) (Fig.
Sinoalaria xiaotu sp. nov., female paratype and male holotype, male habitus (A, B), female habitus (C, D) and epigyne (E–G) A dorsal view B ventral view C dorsal view D ventral view E intact, ventral view F cleared, ventral view G cleared, dorsal view. Abbreviations: CD = copulatory duct; DPF = dorsal and posterior fold of copulatory duct; FD = fertilization duct; Sc = scape; Sp = spermatheca; VAF = ventral and anterior fold of copulatory duct. Scale bars: 0.50 mm (A–D); 0.20 mm (E–G).
Palp
(Fig.
Female (one paratype). Somatic features as in Fig.
Epigyne
(Fig.
The genus Sinoalaria shows a distinct set of genital characters, such as palpal tibia retrolaterally bearing an apophysis in male, and copulatory ducts rising and curling up to form two folds (or chambers, or bursae) at each side in females. In contrast, the RTA is lacking and such conformation of the copulatory ducts has never been seen in any other theridiosomatid genus; therefore, Sinoalaria can be separated from all other genera of Theridiosomatidae. Furthermore, a preliminary molecular analysis of Theridiosomatidae from Southeast Asia was carried out, based on five targeted genes (two mitochondrial genes 16S and COI; three nuclear genes 18S, 28S, and H3). According to the unpublished results, the monophyly of Sinoalaria is strongly supported.
Sinoalaria nitida and S. prolata were assigned to the genus Karstia in the original publication (
In view of the above-mentioned facts, it is currently impossible to discern any obvious derived features that could indicate a close relationship between the two species and the genus Karstia, leaving no doubts that our transfer is correct.
We are grateful to Dr. Cristina A. Rheims (São Paulo, Brazil) for editing the manuscript. Special thanks to Abel A. Bustamante (Belém, Brazil) and an anonymous referee for providing constructive comments on the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was supported by the National Natural Science Foundation of China to Yucheng Lin (NSFC-31972870, 31750002) and Hao Yu (NSFC-32060113, 31702006), the Natural Science Foundation of Guizhou Province to Hao Yu ([2020]1Y081).
Y.C.L. designed the study; Y.C.L. and C.C.F. performed the field work; J.S.Z. and C.C.F. conducted the lab work; Y.C.L. and C.C.F. analysed the data; J.S.Z., Y.C.L. and H.Y. interpreted the results and wrote the paper.
Jianshuang Zhang https://orcid.org/0000-0003-4010-3082
Hao Yu https://orcid.org/0000-0002-9113-2425
Yucheng Lin https://orcid.org/0000-0002-5054-0633
All of the data that support the findings of this study are available in the main text.