Research Article |
Corresponding author: Yun Bu ( buy@sstm.org.cn ) Academic editor: Bruce A. Snyder
© 2023 Ya-Li Jin, Yun Bu.
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 Y-L, Bu Y (2023) Two new species of the genus Symphylella (Symphyla, Scolopendrellidae) from China and the significance of the frons chaetotaxy. ZooKeys 1138: 143-160. https://doi.org/10.3897/zookeys.1138.96424
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Symphylella macrochaeta sp. nov. and Symphylella longispina sp. nov. from China are described and illustrated. Symphylella macrochaeta sp. nov. is characterized by 10 extremely long macrosetae arranged as 4/4/2 on the frons, tergites with broad triangular processes, and 4+4 setae on the first tergite. Symphylella longispina sp. nov. is characterized by a thick and prominent labrum, distinctly long proximal spines on the mandible, eight macrosetae arranged as 4/2/2 on frons, 3+3 setae on first tergite, and narrow triangular processes on the tergites. Detailed comparisons of the new species with similar species are presented. In addition, the frons chaetotaxy of Symphylella is illustrated and discussed for the first time and proposed as a significant diagnostic character for the taxonomic study of the genus.
Chaetotaxy, frons, mandible, Myriapoda, taxonomy
Symphylans are minute soil arthropods present in various habitats, and some species are important crop pests (
Specimens were obtained by extraction of soil and litter samples from broad-leaf and bamboo forests using Berlese-Tullgren funnels. Specimens were preserved in 80% ethanol. They were mounted on slides using Hoyer’s solution and dried in an oven at 50 °C. Observations were performed under a phase contrast microscope (Leica DM 2500). Photographs were taken with a digital camera (Leica DMC 4500) mounted on the microscope. Line drawings were made using a drawing tube. All specimens are deposited in the collections of Shanghai Natural History Museum (SNHM), Shanghai, China.
Family Scolopendrellidae Bagnall, 1913
Symphylella isabellae (Grassi, 1886), described from Italy.
Symphylella macrochaeta sp. nov. is characterized by 10 extremely long macrosetae arranged as 4/4/2 on the frons, 4+4 setae on the first tergite and broad triangular processes on tergites.
Holotype : female (slide no. ZJ-ZS-SY2020029) (SNHM), China, Zhejiang Province, Zhoushan City, Changgang Mountain Forest Park, extracted from soil samples of broad-leaf forest, alt. 250 m, 30°2'N, 121°7'E, 17-XI-2020, coll. Y. L. Jin et al.
Paratypes : 10 females (slides no. ZJ-ZS-SY2020006, ZJ-ZS-SY2020008, ZJ-ZS-SY2020014–ZJ-ZS-SY2020016, ZJ-ZS-SY2020024–ZJ-ZS-SY2020028) (SNHM), same data as holotype. 2 females (slides no. SH-JZGY-SY2017032, SH-JZGY-SY2017034), China, Shanghai, Jiuzi Park, extracted from soil and litter samples of bamboo forest, alt. 14 m, 31°15'N, 121°28'E, 25-V-2017, coll. Y. L. Jin.
Non-type specimens : 18 juveniles with 7–10 pairs of legs, same data as holotype; 5 juveniles with 9 or 10 pairs of legs, China, Shanghai, Jiuzi Park, extracted from soil and litter samples of bamboo forest, alt. 14 m, 31°15'N, 121°28'E, 25-V-2017, coll. Y. L. Jin; 1 juvenile with 10 pairs of legs, China, Shanghai, Tianma Mountain, extracted from soil samples of bamboo forest, alt. 98 m, 31°5'N, 121°9'E, 10-V-2017, coll. Y. Bu.
Adult body 2.1 mm long in average (1.9–2.2 mm, n = 11), holotype 2.1 mm (Fig.
Head
length 250–280 μm, width 223–265 μm, with widest part on equal level of points of articulation of mandibles. Central rod distinct in both anterior (65–70 μm) and posterior (75–85 μm) parts, with an obvious middle node-like interruption. Head dorsally covered with setae of different lengths (Fig.
Symphylella macrochaeta sp. nov. A habitus, dorsal view B head, anterior part, dorsal view C tergites 1–3 D tergites 13–14 E Tömösváry organ F first pair of legs (arrows indicate reduced legs) G tergite 1 H styli and coxal sacs on base of leg 3 (arrows indicate styli). I cerci, dorsal view J left cercus, ventral view (arrows indicate long and erect outer setae). Scale bars: 100 μm (A); 20 μm (B–J).
Tömösváry organ
globular, diameter 15–20 μm, about half of greatest diameter of third antennomere (35–40 μm), opening round (9–12 μm), inner margins of opening covered with regular vertical striae (Fig.
Mouthparts. Mandible composed by pars incisivus (pi) and pars molaris (pm), with movable appendage lacinia mobilis (lm) inserted between them. Pars incisivus with 4 distinct thick teeth, pars molaris with 4 smaller teeth and 2 proximal spines, and lacinia mobilis with only 1 blunt process observed from lateral view (Fig.
Antennae
with 16–20 antennomeres (18 in holotype), about 0.2 of body length. First antennomere cylindrical, length about 0.5–0.8 of greatest diameter (width 33–40 μm, length 18–25 μm), with 6 or 7 setae in 1 whorl, longest inner seta 16–18 μm (Fig.
Numbers of setae and sensory organs on antennae of Symphylella macrochaeta sp. nov. (holotype).
Antennomere | Primary whorl setae | Secondary whorl setae | Rudimentary spined sensory organs | Cavity-shaped organs on dorsal side | Bladder-shaped organs |
---|---|---|---|---|---|
1 | 6 | 1 | |||
2 | 8 | 1 | |||
3 | 8 | 1 | |||
4 | 9 | 1 | |||
5 | 9 | 1 | |||
6 | 11 | 1 | 1 | ||
7 | 11 | 2 | 1 | 1 | |
8 | 11 | 2 | 0 | 1 | |
9 | 10 | 3 | 0 | 1 | |
10 | 11 | 3 | 1 | 1 | 1 |
11 | 11 | 4 | 1 | 1 | 1 |
12 | 11 | 5 | 1 | 1 | 1 |
13 | 11 | 6 | 1 | 1 | 2 |
14 | 11 | 6 | 0 | 1 | 3 |
15 | 10 | 7 | 0 | 1 | 3 |
16 | 10 | 6 | 3 | 5 | |
17 | 10 | 6 | 3 | 8 |
Trunk
with 17 tergites. Tergites 2–13, and 15 each with 1 pair of triangular processes. Length from base to tip of processes somewhat shorter than or same as its basal width; basal distance between processes longer than their length from base to tip except on tergites 2 and 3 (Table
Symphylella macrochaeta sp. nov. A mandible, lateral view (pi – pars incisivus, pm – pars molaris, lm – lacinia mobilis) B first maxilla C left 1–3 antennomere, dorsal view D terminal three antennomeres, dorsal view (bo – bladder-shaped organ, co – cavity-shaped organ, rso – rudimentary spined sensory organ, so – spined sensory organ) E terminal three antennomeres, ventral view F left cercus, dorsal view (arrows indicate long and erect outer setae) G left cercus, ventral view H frons (L1–L5 – lateral setae, a0 – antero-central seta, arrows indicate macrosetae) I leg12, dorso-lateral view. Scale bars: 20 μm.
Tergites. Tergite 1 reduced, with 4+4 setae of different length (Fig.
Chaetotaxy of tergites of Symphylella macrochaeta sp. nov. (holotype in brackets).
Tergite | Lateromarginal setae | Inserted setae | Central setae | Other setae |
---|---|---|---|---|
1 | 4+4 | |||
2 | 6–7 (6) | 1–2 (2) | 1–2 (2) | 6–10 (6) |
3 | 8–10 (8) | 1–2 (2) | 1–3 (2) | 14–25 (14) |
4 | 5–7 (6) | 1–2 (1) | 3–5 (3) | 10–15 (11) |
5 | 5–7 (6) | 1–2 (2) | 2–4 (3) | 7–13 (12) |
6 | 8–10 (9) | 1–2 (2) | 2–4 (4) | 17–28 (20) |
7 | 5–7 (6) | 1–2 (2) | 4–6 (4) | 10–14 (11) |
8 | 5–7 (6) | 1–3 (2) | 3–5 (4) | 10–14 (10) |
9 | 8–10 (9) | 1–3 (1) | 3–5 (3) | 20–27 (22) |
10 | 5–6 (6) | 1–2 (1) | 4–6 (4) | 9–14 (11) |
11 | 5–8 (6) | 1–2 (1) | 3–5 (4) | 7–14 (9) |
12 | 7–10 (8) | 1–2 (2) | 3–5 (3) | 15–24 (15) |
13 | 4–7 (5) | 0–2 (1) | 2–5 (3) | 8–14 (8) |
14 | 17–26 (21) | |||
15 | 6–9 (7) | 0/2 (0) | 2–4 (3) | 14–19 (14) |
16 | 12–17 (14) | |||
17 | 27–38 (29) |
Measurements of tergites and processes of Symphylella macrochaeta sp. nov. (mean ± se, n = 11, in μm) (holotype in brackets).
Tergite | Length | Width | Length of processes | Basal width of processes | Basal distance between processes |
---|---|---|---|---|---|
1 | 24 ± 7 (23) | 141 ± 13 (138) | |||
2 | 48 ± 5 (45) | 144 ± 7 (150) | 37 ± 3 (33) | 41 ± 3 (43) | 28 ± 4 (32) |
3 | 102 ± 14 (100) | 186 ± 25 (180) | 41 ± 4 (42) | 45 ± 3 (47) | 33 ± 3 (37) |
4 | 57 ±7 (55) | 194 ± 17 (205) | 33 ± 3 (35) | 49 ± 5 (52) | 60 ± 8 (65) |
5 | 71 ± 12 (65) | 183 ± 10 (190) | 41± 5 (45) | 43 ± 4 (45) | 62 ± 7 (67) |
6 | 121 ± 9 (125) | 223 ± 38 (235) | 47± 5 (47) | 48 ± 4 (47) | 62 ± 6 (67) |
7 | 70 ± 9 (65) | 229 ± 18 (242) | 36 ± 5 (40) | 48 ± 6 (50) | 85 ± 10 (95) |
8 | 74 ± 5 (82) | 204 ± 13 (205) | 45 ± 4 (50) | 45 ± 3 (50) | 77 ± 10 (85) |
9 | 114 ± 26 (120) | 253 ± 21 (250) | 46 ± 4 (50) | 46 ± 4 (50) | 72 ± 6 (75) |
10 | 76 ± 14 (82) | 235 ± 23 (250) | 34 ± 3 (37) | 48 ± 4 (50) | 92 ± 16 (100) |
11 | 73 ± 7 (70) | 207 ± 13 (210) | 41 ± 4 (45) | 43 ± 3 (42) | 78 ± 9 (85) |
12 | 115 ± 7 (115) | 255 ± 12 (260) | 41 ± 4 (45) | 47 ± 7 (50) | 74 ± 9 (77) |
13 | 68 ± 10 (60) | 233 ± 24 (245) | 28 ± 5 (32) | 48 ± 6 (55) | 89 ± 12 (90) |
14 | 68 ± 10 (60) | 205 ± 20 (210) | |||
15 | 93 ± 9 (90) | 226 ± 21 (247) | 28 ± 3 (32) | 45 ± 5 (52) | 67 ±8 (75) |
16 | 72 ± 6 (80) | 185 ± 25 (200) | |||
17 | 110 ± 8 (125) | 172 ± 21 (175) |
Legs. First pair of legs reduced to 2 small hairy cupules, each with 1 long seta (9–10 μm) (Fig.
Coxal sacs
present at bases of legs 3–9, fully developed, each with 4 or 5 setae on surface (Fig.
Styli
present at base of legs 3–12, slender (length 6–9 μm, width 4–6 μm), basal part with dense straight hairs; distal quarter hairless and with blunt apex (3–5 μm) (Fig.
Sense calicles located on 2 ventral protuberances of last tergite, posterior to base of leg 12, with smooth margin around pit. Sensory seta inserted in cup center, extremely long (110–140 μm).
Cerci
about 0.5–0.6 of head length, 2.5–3 times as long as its greatest width (125–170 μm, 50–63 μm), densely covered with 75–90 subequal setae (Figs
From the Greek words “macro” meaning “large” and “chaeta” meaning “seta”. The species name “macrochaeta” is feminine and refers to extremely long setae on the frons.
China (Shanghai, Zhejiang).
Symphylella macrochaeta sp. nov. has 10 extremely long macrosetae on the frons, which can distinguish it from all other congeners. It is similar to S. communa from East China and S. asiatica Scheller, 1971 from India and Sri Lanka in the shapes of the central rod, tergites, and leg 12, but the new species differs in the chaetotaxy of the first tergite (4+4 setae in S. macrochaeta sp. nov. and S. communa vs 3+3 setae in S. asiatica) and in the shape of stylus (slender in S. macrochaeta sp. nov. vs subconical in S. communa and conical in S. asiatica). The new species can also be compared to S. macropora from Tibet in the shape of tergites and processes, but it can be easily separated by the shape and the size of the opening of the Tömösváry organ (moderate and round in S. macrochaeta sp. nov. vs large and elongate in S. macropora).
Symphylella longispina sp. nov. is characterized by apparently thickened labrum, distinctly long proximal spines on the pars molaris of the mandible, eight macrosetae arranged as 4/2/2 on the frons, 3+3 setae on the first tergite and narrow triangular processes on tergites.
Holotype : female (slide no. XJ-SY20160003) (SNHM), China, Xinjiang, Bole City, Hariturege National Forest Park, extracted from soil samples from the forest of Populus euphratica, alt. 1125 m, 40°08'N, 81°46'E, 31-VIII-2016, coll. C. W. Huang.
Paratypes : 5 females (slides no. XJ-SY20160001, XJ-SY20160002, XJ-SY20160004, XJ-SY20160005, XJ-SY20160006) (SNHM), same data as holotype.
Adult body 2.4 mm long in average (1.8–2.6 mm, n = 6), holotype 2.4 mm (Fig.
Head
length 210–225 μm, width 190–225 μm, with widest part on equal level of points of articulation of mandibles. Central rod well developed but thin, divided into 2 portions by node-like sub-median interruption, with anterior 48–50 μm and posterior 60–70 μm. (Fig.
Symphylella longispina sp. nov. A habitus, dorsal view B head, dorsal view C labrum and mandible D tergites 1–3 E Tömösváry organ and antennomeres 1–4 F tergite 2, right side (als – anterolateral seta, lms – lateromarginl setae, as – apical seta, is – inserted seta, ibs – inner basal seta) G tergites 4–5 H first pair of legs (arrows indicate the reduced legs) I styli and coxal sacs on base of leg 12 (arrows indicate styli). Scale bars: 100 μm (A); 20 μm (B–I).
Symphylella longispina sp. nov. A mandible, lateral view (pi – pars incisivus, pm – pars molaris, lm – lacinia mobilis) B first maxilla C left 1–3 antennomeres, dorsal view D terminal three antennomeres, dorsal view (bo – bladder-shaped organ, co – cavity-shaped organ, rso – rudimentary spined sensory organ, so – spined sensory organ) E terminal three antennomeres, ventral view F left cercus, dorsal view (arrows indicate long and erect outer setae) G left cercus, ventral view H frons (L1–L5 – lateral setae, a0 – antero-central seta, arrows indicate macrosetae) I leg 12, dorso-lateral view. Scale bars: 20 μm.
Tömösváry organ
globular, diameter 12–16 μm, shorter than half of greatest diameter of third antennomere (33–35 μm), opening small and round (4–6 μm), with distinct vertical inner striae (Fig.
Mouthparts
. Labrum apparently thickened and protruding (Figs
Antennae
with 16–20 antennomeres (holotype with 18), about 0.2 of body length. First antennomere cylindrical, almost same as wide as long (width 24–28 μm, length 25–28 μm), with 5–7 setae in 1 whorl, longest inner seta 14–15 μm (Figs
Numbers of setae and sensory organs on antennae of Symphylella longispina sp. nov. (holotype).
Antennomere | Primary whorl setae | Secondary whorl setae | Rudimentary spined sensory organs | Cavity-shaped organs on dorsal side | Bladder-shaped organs |
---|---|---|---|---|---|
1 | 6 | ||||
2 | 8 | 1 | |||
3 | 8 | 1 | |||
4 | 9 | 1 | |||
5 | 10 | 1 | |||
6 | 10 | 1 | |||
7 | 10 | 1 | 1 | ||
8 | 10 | 0 | 1 | ||
9 | 10 | 0 | 1 | ||
10 | 10 | 1 | 0 | 1 | |
11 | 10 | 1 | 1 | 1 | |
12 | 11 | 3 | 1 | 1 | 1 |
13 | 11 | 4 | 0 | 1 | 3 |
14 | 11 | 4 | 0 | 1 | 5 |
15 | 11 | 5 | 2 | 9 | |
16 | 11 | 4 | 3 | 13 | |
17 | 10 | 5 | 2 | 13 |
Trunk
with 17 tergites. Tergites 2–13 and 15 each with 1 pair of triangular processes. Length from base to tip of processes slightly longer than its basal width except for tergites 4, 7, 10 and 13, in which processes almost as broad as long; basal distance between processes of tergites distinctly longer than their length from base to tip (Table
Chaetotaxy of tergites of Symphylella longispina sp. nov. (holotype in brackets).
Tergite | Lateromarginal setae | Inserted seta | Central setae | Other setae |
---|---|---|---|---|
1 | 3+3 | |||
2 | 5–6 (6) | 1 (1) | 1 (1) | 5–7 (6) |
3 | 7–8 (7) | 1 (1) | 1 (1) | 16–19 (16) |
4 | 5 (5) | 1 (1) | 1–2 (2) | 8–8 (8) |
5 | 5 (5) | 1 (1) | 1–2 (1) | 7–10 (8) |
6 | 7–8 (7) | 1 (1) | 2–3 (2) | 15–20 (20) |
7 | 5 (5) | 1 (1) | 2–3 (2) | 8–10 (8) |
8 | 5 (5) | 1 (1) | 2 (2) | 0–11 (9) |
9 | 7–8 (7) | 1 (1) | 2–3 (3) | 16–20 (18) |
10 | 5 (5) | 1 (1) | 2–3 (3) | 8–10 (8) |
11 | 5–6 (5) | 1 (1) | 2–3 (2) | 6–10 (9) |
12 | 6–7 (7) | 0/1 (1) | 2–3 (3) | 15–20 (17) |
13 | 4–5 (5) | 0/1 (1) | 1–2 (2) | 7–8 (7) |
14 | 15–18 (16) | |||
15 | 4–7 (5) | 0/1 (1) | 1–2 (2) | 8–14 (14) |
16 | 10–14 (14) | |||
17 | 10–14 (14) |
Tergites. Tergite 1 reduced, with 3+3 subequal setae (Fig.
Measurements of tergites and processes of Symphylella longispina sp. nov. (mean ± se, n = 6, in μm) (holotype in brackets).
Tergite | Length | Width | Length of processes | Basal width of processes | Basal distance between processes |
---|---|---|---|---|---|
1 | 27 ± 3 (25) | 126 ± 2 (125) | |||
2 | 45 ± 6 (50) | 123 ± 10 (130) | 32 ± 2 (34) | 28 ± 2 (30) | 33 ± 3 (33) |
3 | 97 ± 16 (90) | 151 ± 8 (155) | 34 ± 2 (37) | 28 ± 2 (32) | 39 ± 2 (40) |
4 | 60 ± 8 (62) | 162 ± 9 (170) | 31 ± 2 (29) | 34 ± 1 (33) | 47 ± 6 (50) |
5 | 60 ± 12 (75) | 138 ± 5 (140) | 35 ± 4 (39) | 28 ± 3 (30) | 52 ± 5 (50) |
6 | 115 ± 12 (125) | 181 ± 17 (192) | 39 ± 3 (41) | 31 ± 4 (33) | 60 ± 4 (63) |
7 | 72 ± 12 (85) | 191 ± 10 (202) | 38 ± 4 (36) | 39 ± 6 (35) | 65 ± 7 (75) |
8 | 71 ± 12 (80) | 160 ± 9 (170) | 35 ± 3 (38) | 27 ± 3 (29) | 65 ± 5 (70) |
9 | 124 ± 10 (138) | 199 ± 6 (205) | 37 ± 3 (41) | 30 ± 3 (34) | 68 ± 7 (75) |
10 | 74 ± 12 (90) | 196 ± 17 (207) | 36 ± 3 (37) | 36 ± 6 (36) | 74 ± 5 (80) |
11 | 72 ± 9 (80) | 171 ± 5 (175) | 35 ± 1 (34) | 29 ± 3 (25) | 70 ± 6 (78) |
12 | 121 ± 18 (125) | 201 ± 14 (217) | 37 ± 4 (38) | 30 ± 6 (32) | 66 ± 7 (70) |
13 | 71 ± 6 (75) | 184 ± 16 (207) | 29 ± 3 (27) | 36 ± 12 (30) | 66 ± 9 (72) |
14 | 68 ± 10 (75) | 164 ± 8 (175) | |||
15 | 99 ± 10 (100) | 180 ± 13 (200) | 27 ± 2 (30) | 26 ± 3 (26) | 55 ± 8 (65) |
16 | 71 ± 7 (75) | 148 ± 16 (162) | |||
17 | 92 ± 9 (92) | 131 ± 9 (135) |
Legs. First pair of legs reduced to 2 small hairy cupules, each with 1 long seta (9–11 μm) (Fig.
Coxal sacs
present at bases of legs 3–9, fully developed, each with 4 setae on surface. Corresponding area of leg 2, 10, 11 and 12 replaced by 1–3 setae (Fig.
Styli
present at base of legs 3–12, subconical (length 5 μm, width 3 μm), basal part with straight hairs; distal quarter hairless and with blunt apex (3 μm) (Fig.
Sense calicles with smooth margin around pit. Sensory seta inserted in cup center, extremely long (115–120 μm).
Cerci
about half length of head, 3.3–3.8 times as long as its greatest width (108–115 μm, 30–34 μm), sparsely covered with 33–39 subequal setae (Fig.
The species name is derived from the Latin words “longus” and “spina” meaning “long spine”. It is feminine and refers to the extremely long proximal spines on the pars molaris of the mandible.
Known only from the type locality.
Symphylella longispina sp. nov. has a thickened and prominent labrum and irregular bladder-shaped organs on antennae, which separate it from all other congeners. It is most similar to S. asiatica Scheller, 1971 from India and Sri Lanka in the shape and chaetotaxy of the tergites, but the new species differs in the distal part of the processes (distinctly swollen in S. longispina sp. nov. vs small and slender in S. asiatica), in the shape and chaetotaxy of cerci (subcylindrical and with sparse setae in S. longispina sp. nov. vs conical and with dense setae in S. asiatica), and in the shape of the palp of the first maxilla (slightly curved in S. longispina sp. nov. vs straight in S. asiatica). The new species is also similar to S. brincki Scheller, 1971 from Sri Lanka in the chaetotaxy of the tergites, but they can be easily separated by the central rod (with a middle node-like interruption in S. longispina sp. nov. vs with a narrow transverse interruption in S. brincki), by the end of the processes (with round end-swellings in S. longispina sp. nov. vs spatulate end-swellings in S. brincki), and by the shape and chaetotaxy of cerci (3.3–3.8 times as long as wide and with sparse setae in S. longispina sp. nov. vs 2.3 times as long as wide and with dense setae in S. brincki).
Symphylella is one of the most common and diverse group of symphylans with a wide global distribution (
The mandible structure of Symphyla was carefully studied and compared with other arthropods by former colleagues (
In our study of Symphylella specimens from Zhejiang and Shanghai, we observed that the extremely long setae on the frons of S. macrochaeta sp. nov. differ from other Chinese congeners (Fig.
According to our observations, the frons of Symphylella spp. often has well-differentiated macrosetae located on the 2/3 anterior part and 5+5 setae on the lateral margin. The quantity, length, arrangement, and ratio to antero-central seta of the macrosetae vary among species but vary little among conspecific individuals (Table
Characters | S. macrochaeta sp. nov. | S. longispina sp. nov. | S. macropora | S. zhongi | S. communa | S. minuta |
---|---|---|---|---|---|---|
Number of macrosetae (M) | 10 | 8 | 6 | 8 | 8 | 4 |
Formula of M-setae | 4/4/2 | 4/2/2 | 2/2/2 | 4/2/2 | 4/2/2 | 0/2/2 |
length of M-setae (μm) | 58–73 | 21–28 | 25–37 | 22–37 | 20–30 | 12–20 |
length of a0 setae (μm) | 12–15 | 7–8 | 12 | 12–15 | 10–16 | 7–11 |
M/a0 | 4–5.6 | 2.8–3.7 | 2–3 | 1.5–2.7 | 1.6–2.7 | 1.4–2.4 |
We sincerely thank Dr Jing-Yang Li, Mrs Si-Qi Yang and Mr Cheng-Wang Huang for their generous help in the collection. We appreciate Dr Nerivania Nunes Godeiro for the linguistic corrections to the manuscript. Special thanks are given to Dr Derek Hennen (USA) and an anonymous reviewer for their valuable comments in review of the manuscript. This research was supported by the National Natural Science Foundation of China (no. 32170471) and the Research Foundation of Shanghai Science and Technology Museum.