Research Article |
Corresponding author: Tongzuo Zhang ( zhangtz@nwipb.cas.cn ) Corresponding author: Jianping Su ( jpsu@nwipb.cas.cn ) Academic editor: Marzio Zapparoli
© 2018 Penghai Qiao, Wen Qin, Huiqin Ma, Tongzuo Zhang, Jianping Su, Gonghua 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:
Qiao P, Qin W, Ma H, Zhang T, Su J, Lin G (2018) Two new species of Lithobius on Qinghai-Tibetan plateau identified from morphology and COI sequences (Lithobiomorpha: Lithobiidae). ZooKeys 785: 11-28. https://doi.org/10.3897/zookeys.785.28580
|
Lithobius (Ezembius) longibasitarsussp. n. and Lithobius (Ezembius) datongensissp. n. (Lithobiomorpha: Lithobiidae), recently discovered from Qinghai-Tibet Plateau, China, are described. A key to the species of the subgenus Ezembius in China is presented. The partial mitochondrial cytochrome c oxidase subunit I barcoding gene was amplified and sequenced for eight individuals of the two new species and the dataset was used for molecular phylogenetic analysis and genetic distance determination. Both morphology and molecular data show that the specimens examined should be referred to Lithobius (Ezembius).
China, COI, Ezembius , Lithobiidae , Qinghai-Tibet Plateau, taxonomy
The myriapod fauna of China has been poorly investigated and this is especially the case with centipedes of the order Lithobiomorpha, with only approximately 80 species/subspecies of lithobiomorphs are known from the country. Qinghai-Tibet Plateau is among the very poorly studied regions of China (
The centipede subgenus Ezembius was erected by
Specimen collection and preparation: the specimens were all collected by hand, preserved in 95% ethanol, and deposited in the collections of Northwest Institute of Plateau Biology (NWIPB), Chinese Academy of Sciences. Characters were examined using an Olympus SZ61 stereoscope. Terminology for external anatomy follows
a anterior;
C coxa;
D dorsal;
DT Datong.
F femur;
GH Gonghe,
m median;
P prefemur;
p posterior;
T, TT tergite, tergites;
S, SS sternite, sternites;
Ti tibia;
To Tömösváry’s organ;
Tr trochanter;
Ts I tarsus I;
Ts II tarsus II;
V ventral.
DNA extraction and sequencing protocols: standard DNA extraction and amplification methods were performed. Total DNA was extracted from a single leg removed from each specimen sample using MicroElute Genomic DNA kit (OMEGA), after overnight incubation at 65°C. Polymerase chain reactions (PCRs) were conducted using Mastercycler pros PCR (Eppendorff) in total reaction volumes of 39μL volumes containing 5–60 ng template DNA, 1μL; ddH2O 28μL; 10×Buffer 5μL (Takara, Dalian, China); 0.5mm/L dNTPs 2.5μL (Takara, Dalian, China); 5U/μL Taq polymerase 0.5μL (Takara, Dalian, China); Forward Primer 1μL; Reverse Primer 1μL (synthesized by Sangon Biotech from Shanghai). A 686 bp fragment of COI was amplified using the primers LCO1490/LCO2198 (
The GenBank accession numbers of all eight new sequences were MH05602–MH045609 (Ezembius COI). Sequence identities were confirmed with BLAST searches (
Phylogenetic analyses: the sequences were aligned with Clustal X2.0 (
Distance analysis: the analysis involved 26 nucleotide sequences (App. 1). Codon positions included were 1st+2nd+3rd. All ambiguous positions were removed for each sequence pair. There were a total of 613 positions in the final dataset. Evolutionary analyses were conducted in MEGA6 (
Holotype: female labelled GH3 (Figure
Specimens were collected under stones on steppes covered with legume shrubs and grass composed mainly of Poaceae. The sampling point belongs to the Gonghe Basin region of the Tibet plateau severely affected by desertification.
The specific name refers to the new species with a long tarsus I of leg XV, tarsus I approx. 1.7 times longer than tarsus II.
Body length 17.0–18.0 mm; head slightly widened; antennae of 20 articles; 10–14 ocelli arranged in three irregular rows; To oval to round, slightly smaller in size to neighbouring ocelli; lateral margins of forcipular coxosternite slanting; anterior margin with 2+2, 3+2 or 2+3 blunt teeth and with strong setiform porodonts; tergites without triangular posterolateral process; legs XIV and XV thicker and longer than anterior ones in both sexes; coxal pores 4–6, round to ovate arranged in one row; female gonopods with two moderately long, bullet-shaped spurs; terminal claw of the third article simple, with a small triangular protuberance on basal ventral side; male gonopods short and small.
Holotype (♀), body 17.0 mm long, cephalic plate width 2.1 mm, length 2.0 mm.
Colour: antennae light yellow; tergites pale yellow-brown; cephalic plate and terminal tergite yellow-brown; pleural region and sternites pale yellow; distal part of forcipules dark brown, maxillipede coxosternum and SXV yellow; legs pale yellow with grey hue, pretarsal claw brown.
Antennae composed of 20+20 articles (Figure
Ocelli area translucent with dark pigment, 1+5, 3, 2 ocelli on each side of cephalic plate, arranged in three irregular rows. The posterior ocellus is the biggest. To oval, smaller than the adjacent ocelli, situated ventrally on anterolateral margin of cephalic plate.
Cephalic plate smooth, slightly broader than long; as broad as TIII or slightly broader. Frontal marginal of head with clear transverse suture. Posterior margin slightly concave; projection of lateral marginal conspicuously discontinuous; posterior marginal ridge slightly concave with median thickening.
Coxosternite subtrapezoidal, anterior margin narrow, lateral margins of the coxosternite slightly longer than medial margins. Median diastema shallow, U-shaped; anterior margin with 3+2 blunt nipple-like teeth (Figure
Tergites all smooth, without wrinkles, TI narrower posterolaterally than anterolaterally, generally trapezoidal, narrower than the cephalic plate and TIII, the cephalic plate almost the same width as TIII. Posterior marginal ridge of TI straight; of TT III, V shallow concave; of TT VIII, X, XII slightly concave; of TXIV deeply concave; TT VI– XIV bordered laterally only (Figure
Sternites: posterior part of sternites narrower than anterior, generally trapezoidal, smooth; 2–8 setae on anterior angle, anterior lateral side, posterior angle and posterior lateral side; some minute setae on SS XIV and XV, most of which distributed on posterior lateral margins and posterior borders.
Legs: tarsal articulation well defined on legs I-XV. All legs with fairly long curved claws. Legs I–XIV with anterior and posterior accessory spurs, anterior accessory spur moderately long and slender ca. 33%-50% the length of principle claw, the posterior one stouter forming slightly larger angles with tarsal claws, ca. 0.25 the length of principal claw. Legs XV lacking anterior and posterior accessory spurs. Dense glandular pores on the surface of prefemur, femur, tibia, and tarsi of legs XIV and XV. Short to long setae sparsely scattered over the surface of prefemur, femur, tibia, and tarsi of legs I-XIII, more setae on the tarsal surface, with two rows of comb-like setae along ventral side, fewer setae on legs XIV and XV. Legs XIV and XV moderately thicker and longer than anterior legs, tarsus I ca. 6.6 times as long as wide, tarsus II ca. 37% length of the whole tarsus on leg XV. Leg plectrotaxy as presented in Table
Coxal pores circular on legs XII–XV, separated by a distance 1–2 times larger than diameter of pore; inner pores smaller; formula 6, 5, 5, 5. Coxal pores set in a shallow groove arranged in a row with short to long setae scattered over the surface of apophysis (Figure
Female posterior segment: S XV generally trapeziform, straight posteromedially; sternite of genital segment wider than long with posterior margin moderately concave between condyles of gonopods, except for a small, median bulge; distal part lightly sclerotised; short to long setae scattered over the surface of genital segment and lateral margins. The first article of gonopod moderately broad bearing 22–24 short to moderately long setae arranged in three rows with 2+2 moderately long, bullet-shaped spurs, inner spur slightly smaller and more anterior than the outer (Figure
Male posterior segment: S XV subtrapeziform, long setae scattered sparsely over its surface and posterior margins. Male genital sternite slightly wider than long; posterior margin quite deeply concave between the gonopods, no bulge medially; ca. 69 short to medium setae scattered sparsely over its surface and at lateral margins; gonopods of a single small semicircular article with 3-5 seta on its surface (Figure
Lithobius (Ezembius) longibasitarsus sp. n.: leg plectrotaxy; letters in brackets indicate variable spines.
legs | ventral | dorsal | ||||||||
---|---|---|---|---|---|---|---|---|---|---|
C | Tr | P | F | Ti | C | Tr | P | F | Ti | |
1 | (a)mp | amp | am | ap | ap | ap | ||||
2–3 | amp | amp | am | ap | ap | ap | ||||
4–7 | amp | amp | am | a(m)p | ap | ap | ||||
8–11 | amp | amp | am | amp | ap | ap | ||||
12 | amp | amp | am | (a) | amp | ap | ap | |||
13 | amp | amp | am | a | amp | p | ap | |||
14 | m | amp | amp | am | a | amp | p | p | ||
15 | m | amp | am | a | a | amp | p | (p) |
Body length 17.0–18.0 mm; ocelli 1+5, 4, 4 or 1+5, 3, 2 or 1+4, 3, 2 (Figure
Lithobius (E.) longibasitarsus sp. n. can be distinguished from all the other known Chinese species of subgenus Ezembius Chamberlin, 1919 by 2+2, 2+3 or 3+2 moderately blunt teeth on the forcipular coxosternite and the terminal claw of the female gonopod simple, slender and sharp, having a small triangular protuberance on its ventral side. It has a larger body (17.7–18.0 mm), more ocelli (10–14), more coxal pores (5544, 5554 or 6555), and DaC spine on legs XII and XV.
Morphologically it resembles L. (E.) tetraspinus but can be readily distinguished by the following characters: more ocelli (10–14 vs. 9–10), more coxosternal teeth (2+3, 3+2 vs. 2+2), more coxal pores (4–6 vs. 2–5), and less spurs on female gonopods (2+2 contrary to 2+3 or 3+2).
Lithobius (Ezembius) longibasitarsus sp. n., A, D, E, H–J holotype, female: A dorsal view D ocelli and To, lateral view E forcipular coxosternite, ventral view H posterior segments and gonopods, dorsal view I claw of female gonopod, inboard view J female posterior segments and gonopods, ventral view B, F paratype, female, labelled GH4: B ocelli and To, lateral view F forcipular coxosternite, ventral view C, G, K paratype, male, labelled GH8: C ocelli and To, lateral view G forcipular coxosternite, ventral view K posterior segments and gonopods, ventral view. Scale bars 1 mm A, E, F, G; 300 μm B, C, D, H, I; 500 μm J.
Holotype: female labelled DT5 (Figure
Specimens were collected under stones of slope-lands covered with grass mainly of Pedicularis chinensis and shrub mainly of Potentilla fruticosa along the riverside in coniferous forest composed mainly of Picea crassifolia.
The name is derived from the locality Datong County where the new species was discovered.
Body length 12.3–14.2 mm; antennae composed of 20+20 articles; 10 ocelli on each side arranged in 3 irregular rows, terminal one ocellus comparatively large; To larger than the adjoining ocelli; 2+2 coxosternite teeth and setiform porodonts posterolateral to the lateralmost tooth; posterior angles of all tergites without triangular projections; tarsal articulation well-defined on all legs; legs XII–XV with DaC, leg XV with posterior accessory claw; coxal pores 4–7, round, arranged in one row; female gonopods with 2+2 moderately large, coniform spurs; claw of the third article simple, with a small triangular protuberance on basal ventral side; male gonopods short and small.
Holotype (♀), body 14.2 mm long, cephalic plate width 1.54 mm, length 1.54 mm.
Colour red-brown, with a distinct, darker, axial stripe on cephalic plate and tergites. Legs pale yellow-brown. Sternite yellow-brown with distal part brown with reddish hue.
Antennae tapering, ca. 4.3 mm long, reaching the anterior part of T V, composed of 20 elongate articles (Figure
Ocelli area: ten on each side, dark, arranged in three broken rows; posterior ocellus slightly larger than posterosuperior ocellus and other seriated ocelli. To slightly larger than nearest ocellus, rounded.
Cephalic plate: breath/length ratio 1.0 (1.54 mm); smooth, longer setae scattered along the entire surface sparsely and the marginal ridge of the cephalic plate. Transverse suture distinct, lateral marginal ridge discontinuous, posterior margin continuous, slightly concave (Figure
Coxosternite: dental margin slightly concave, with 2+2 slightly acute teeth and setiform porodonts separated from the lateral tooth laterally, median diastema U-shaped; shoulders of coxosternite strongly sloping, as in Figure
Tergites almost smooth. The anterior part of T I is approx. the same width as cephalic plate and T III; T I and T III approximately the same width. Posterior angles of all tergites rounded without triangular projections. Posterior margin of TI straight; posterior margin of TT III, V, VIII, X, XII, and XIV concave; posterior margin of TVII convex; posterior margin of intermediate T straight; TT VI–XIV bordered laterally only (Figure
Sternites: posterior side of sternites narrower than anterior, generally trapezoidal, smooth; SS XIII–XV with miniscule setae scattered sparsely over the surface; genital sternite more densely setose, as in Figure
Legs: tarsal articulation of all legs distinct. Legs XIV and XV incrassate, without visible modification. Length of legs XV: F = 0.85 mm, Ti = 1.00 mm, Ts I = 0.77 mm, Ts II = 0.54 mm. Legs XII– XV with DaC. All legs with fairly long curved claws; legs I– XIV with anterior and posterior accessory spurs, anterior accessory spur moderately long and slender, posterior accessory spur slightly more robust; the anterior accessory spines form relatively small angles with the main claw, the posterior accessory spines form relatively large angles with the main claw; posterior accessory spines on legs XIV. Numerous glandular pores scattered on the surface of prefemur, femur, tibia, tarsus of legs XIV and XIV; short to long comparatively long setae scattered very sparsely over the surface of all segments of legs I– XIII, more setae scattered on the whole surface of tarsus, slightly thick setae arranged in two rows on the ventral side of tarsus. Plectrotaxy as presented in Table
Coxal pores present on legs XII–XV, rounded and separated by distance 0.2–2.5 times greater than their own diameter; inner pores smaller than neighbouring ones; formula 4655 and 5575. Coxal pores 4654 and 4554 in male. Coxal pore field set in a relatively shallow groove, the coxal pore-field fringe with prominence. Prominence with short to moderately long setae sparsely scattered over the surface.
Female posterior segment S XV generally trapeziform, straight posteromedially; sternite of genital segment wider than long with posterior margin moderately concave between condyles of gonopods, except for a small, median bulge; distal part lightly sclerotised; short to long setae scattered over the surface of genital segment and lateral margins. Basal article of gonopod bearing 22–25 setae, with two blunt spurs of approximately equal size at distal end of slender, elongate process and three long spines on dorsolateral side; second article of gonopod with 5-6 setae and five long curved spines on dorsolateral side; third article with two setae (Figure
Male posterior segment (Figure
Lithobius (Ezembius) datongensis sp. n.: leg plectrotaxy; letters in brackets indicate variable spines.
legs | ventral | dorsal | ||||||||
---|---|---|---|---|---|---|---|---|---|---|
C | Tr | P | F | Ti | C | Tr | P | F | Ti | |
1 | mp | (a)mp | am | ap | a(p) | ap | ||||
2–9 | amp | amp | am | ap | ap | ap | ||||
10 | amp | amp | am | ap | ap | ap | ||||
11 | amp | amp | am | a(m)p | ap | ap | ||||
12–14 | m | amp | amp | am | a | amp | p | p | ||
15 | m | amp | am | a | a | amp | p |
Body length 12.3–14.2 mm; 9–10 ocelli; coxal pores 4655, 5575, or 5544 in female, 4654 and 4554 in male.
The new species can be easily distinguished from the other species of the subgenus of Ezembius of China except Ezembius anabilineatus by the apical claw of female gonopods simple with a small subtriangular protuberance on the ventral side. It differs from E. anabilineatus in many aspects, such as a larger body, fewer antennal articles (20+20, vs. 23+23 in E. anabilineatus), more ocelli, a DaC spine on legs XII–XV (only on legs XIV and XV in E. anabilineatus), and a posterior accessory spur present on legs XV present. It differs from Lithobius (Ezembius) longibasitarsus sp. n. by having posterior accessory spur on XV legs, fewer ocelli (10 versus up to 14 in E. longibarsitarsus) and different plectrotaxy (VmTr absent on legs XII and XIII vs. present).
Lithobius (Ezembius) datongensis sp. n., A, C–E holotype, female: A habitus, dorsal view; C forcipular coxosternite, ventral view D female gonopods, dorsal lateral view E female posterior segments and gonopods B, F paratuype, male: B ocelli and Tömösváry’s organ (To), lateral view F posterior segments and gonopods, ventral view. Scale bars 1 mm A, C; 300 μm B, D; 500 μm E, F.
1 | Posterior angles of tergites with triangular projections | 2 |
– | Posterior angles of tergites rounded, without projections | 3 |
2 | Posterior angles of TT VII, IX, XI, XIII with triangular projections | L. (E.) kiayiensis Wang, 1959 |
– | Posterior angles of TT XIV with slightly triangular projections | L. (E.) sulcipes Attems, 1927 |
3 | At most four ocelli on each side of cephalic plate | L. (E.) parvicornis (Porat, 1893) |
– | At least five ocelli on each side of cephalic plate | 4 |
4 | Cephalic plate with scattered, rough punctae and tergites with distinct punctae | L. (E.) rhysus Attems, 1934 |
– | Cephalic plate and tergites without any punctae | 5 |
5 | All ocelli subequal in size | 6 |
– | All ocelli not subequal in size | 7 |
6 | Six ocelli on each side of cephalic plate | L. (E.) sulcifemoralis Takakuwa & Takashima, 1949 |
– | Eight to twelve ocelli on each side of cephalic plate | L. (E.) sibiricus Gerstfeldt, 1858 |
7 | Posterior ocellus small | L. (E.) lineatus Takakuwa, 1939 |
– | Posterior ocellus large | 8 |
8 | The terminal two ocelli comparatively large | 9 |
– | The terminal one ocellus comparatively large | 12 |
9 | Ocelli arranged in two rows | L. (E.) laevidentata Pei, Ma, Hou, Zhu & Gai, 2015 |
– | Ocelli arranged in three rows | 10 |
10 | 3+3 coxosternal teeth | L. (E.) multispinipes Pei, Lu, Liu, Hou, Ma & Zapparoli, 2016 |
– | 2+2 coxosternal teeth | 11 |
11 | Tömösváry’s organ larger than the adjoining ocellus | L. (E.) bilineatus Pei, Ma, Zhu & Gai, 2014 |
– | Tömösváry’s organ smaller than the adjoining ocellus | L. (E.) anabilineatus Ma, Pei, Hou, Zhu & Gai, 2015 |
12 | Only five ocelli on each side of cephalic plate | L. (E.) chekianus Chamberlin & Wang, 1952 |
– | At least six ocelli on each side of cephalic plate | 13 |
13 | Tömösváry’s organ smaller than the adjoining ocellus. | 14 |
– | Tömösváry’s organ larger than the adjoining ocellus | 16 |
14 | Ocelli arranged in three rows | L. (E.) longibasitarsus sp. n. |
– | Ocelli arranged in two rows | 15 |
15 | First article of female gonopods with 3+3 spurs | L. (E.) insolitus Eason, 1993 |
– | First article of female gonopods with 2+2 or 2+3 spurs | L. (E.) irregularis Takakuwa & Takashima, 1949 |
16 | First article of female gonopods with 1 + 1 spurs | L. (E.) gantoensis Takakuwa & Takashima, 1949 |
– | First article of female gonopods with more than 1+1 spurs | 17 |
17 | First article of female gonopods with 3+3 or 4+4 spurs | 18 |
– | First article of female gonopods with 2+2 or 2+3 spurs | 19 |
18 | Terminal claw of female gonopods simple without a small subtriangular tooth on inner margin | L. (E.) mandschreiensis Takakuwa, 1939 |
– | Terminal claw of female gonopods simple with a small subtriangular tooth on inner margin | L. (E.) bidens Takakuwa, 1939 |
19 | Terminal claw of female gonopods bipartite | L. (E.) anasulcifemoralis Ma, Pei, Wu & Gai, 2013 |
– | Terminal claw of female gonopods not bipartite | 19 |
20 | Terminal claw of female gonopods tridentate | L. (E.) zhui Pei, Ma, Shi, Wu & Gai, 2011 |
– | Terminal claw of female gonopods simple | 20 |
21 | Terminal claw of female gonopods simple without a small subtriangular teeth on inner margin | L. (E.) giganteus Sseliwanoff, 1881 |
– | Terminal claw of female gonopods simple with a small subtriangular teeth on inner margin | L. (E.) datongensis sp. n. |
Sequence characterisation. Alignment of the PCR fragment sequences from COI showed that in the 613 bp there were 271 variable sites and 258 parsimony informative characters. The base composition of the fragments showed a strong bias of A + T (29.0+32.3). The results of the substitution saturation test showed that the index of substitution saturation 0.2562 (Iss) is significantly lower than the critical value of the index of substitution saturation 0.7345 (Iss. c).
Genetic distance. Calculation of the distances (Table
Phylogenetic relationship. Bayesian inference (BI) analysis (Figure
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | 24 | 25 | |
1. DT4 | |||||||||||||||||||||||||
2. DT6 | 0.0016 | ||||||||||||||||||||||||
3. DT5 | 0.0016 | 0.0000 | |||||||||||||||||||||||
4. GH04 | 0.1794 | 0.1794 | 0.1794 | ||||||||||||||||||||||
5. GH05 | 0.1794 | 0.1794 | 0.1794 | 0.0000 | |||||||||||||||||||||
6. GH03 | 0.1794 | 0.1794 | 0.1794 | 0.0000 | 0.0000 | ||||||||||||||||||||
7. GH011 | 0.1794 | 0.1794 | 0.1794 | 0.0000 | 0.0000 | 0.0000 | |||||||||||||||||||
8. GH06 | 0.1794 | 0.1794 | 0.1794 | 0.0000 | 0.0000 | 0.0000 | 0.0000 | ||||||||||||||||||
9. MF123702.1 | 0.1990 | 0.1974 | 0.1974 | 0.2055 | 0.2055 | 0.2055 | 0.2055 | 0.2055 | |||||||||||||||||
10. HM453305.1 | 0.1876 | 0.1876 | 0.1876 | 0.1811 | 0.1811 | 0.1811 | 0.1811 | 0.1811 | 0.2039 | ||||||||||||||||
11. MF123710.1 | 0.1876 | 0.1892 | 0.1892 | 0.2202 | 0.2202 | 0.2202 | 0.2202 | 0.2202 | 0.2153 | 0.1811 | |||||||||||||||
12. HM453306.1 | 0.1958 | 0.1941 | 0.1941 | 0.1794 | 0.1794 | 0.1794 | 0.1794 | 0.1794 | 0.2088 | 0.1794 | 0.1941 | ||||||||||||||
13. HM453307.1 | 0.2104 | 0.2104 | 0.2104 | 0.2055 | 0.2055 | 0.2055 | 0.2055 | 0.2055 | 0.2235 | 0.1713 | 0.1974 | 0.1697 | |||||||||||||
14. AF334311.1 | 0.2072 | 0.2072 | 0.2072 | 0.1925 | 0.1925 | 0.1925 | 0.1925 | 0.1925 | 0.2055 | 0.2007 | 0.2202 | 0.1794 | 0.1909 | ||||||||||||
15. JN269950.1 | 0.1974 | 0.1974 | 0.1974 | 0.1990 | 0.1990 | 0.1990 | 0.1990 | 0.1990 | 0.1990 | 0.1697 | 0.1843 | 0.1582 | 0.1860 | 0.1778 | |||||||||||
16. HM453308.1 | 0.2219 | 0.2202 | 0.2202 | 0.2039 | 0.2039 | 0.2039 | 0.2039 | 0.2039 | 0.2153 | 0.1925 | 0.1958 | 0.1990 | 0.2023 | 0.2007 | 0.1827 | ||||||||||
17. AY214425.1 | 0.2545 | 0.2561 | 0.2561 | 0.2496 | 0.2496 | 0.2496 | 0.2496 | 0.2496 | 0.2626 | 0.2235 | 0.2333 | 0.2202 | 0.2333 | 0.2577 | 0.2251 | 0.2365 | |||||||||
18. AF334316.1 | 0.2284 | 0.2268 | 0.2268 | 0.2300 | 0.2300 | 0.2300 | 0.2300 | 0.2300 | 0.2398 | 0.2088 | 0.2251 | 0.2251 | 0.2480 | 0.2545 | 0.2202 | 0.2349 | 0.1599 | ||||||||
19. AY214428.1 | 0.2512 | 0.2512 | 0.2512 | 0.2349 | 0.2349 | 0.2349 | 0.2349 | 0.2349 | 0.2365 | 0.2170 | 0.2153 | 0.2300 | 0.2186 | 0.2349 | 0.2072 | 0.2251 | 0.1746 | 0.1713 | |||||||
20.AF334315.1 | 0.2121 | 0.2137 | 0.2137 | 0.2365 | 0.2365 | 0.2365 | 0.2365 | 0.2365 | 0.2284 | 0.2088 | 0.2039 | 0.2055 | 0.2235 | 0.2414 | 0.2121 | 0.2268 | 0.1794 | 0.1697 | 0.1729 | ||||||
21. KX442654.1 | 0.2382 | 0.2398 | 0.2398 | 0.2300 | 0.2300 | 0.2300 | 0.2300 | 0.2300 | 0.2316 | 0.2219 | 0.2039 | 0.2316 | 0.2398 | 0.2414 | 0.2186 | 0.2349 | 0.1909 | 0.1843 | 0.1876 | 0.1876 | |||||
22. AF334330.1 | 0.2398 | 0.2414 | 0.2414 | 0.2398 | 0.2398 | 0.2398 | 0.2398 | 0.2398 | 0.2219 | 0.2382 | 0.2186 | 0.2496 | 0.2414 | 0.2463 | 0.2170 | 0.2594 | 0.2202 | 0.2023 | 0.1925 | 0.1958 | 0.1892 | ||||
23. AF334332.1 | 0.2529 | 0.2512 | 0.2512 | 0.2431 | 0.2431 | 0.2431 | 0.2431 | 0.2431 | 0.2431 | 0.2349 | 0.2251 | 0.2186 | 0.2349 | 0.2349 | 0.1925 | 0.2072 | 0.2055 | 0.1778 | 0.1876 | 0.1990 | 0.2072 | 0.1925 | |||
24. AF334313.1 | 0.2284 | 0.2284 | 0.2284 | 0.2170 | 0.2170 | 0.2170 | 0.2170 | 0.2170 | 0.2121 | 0.1892 | 0.2039 | 0.2170 | 0.1892 | 0.2219 | 0.1762 | 0.2121 | 0.2055 | 0.1958 | 0.1664 | 0.1827 | 0.1925 | 0.1941 | 0.1860 | ||
25. AF334320.1 | 0.2316 | 0.2316 | 0.2316 | 0.2219 | 0.2219 | 0.2219 | 0.2219 | 0.2219 | 0.2235 | 0.2104 | 0.2480 | 0.2251 | 0.2284 | 0.2300 | 0.2251 | 0.2300 | 0.2219 | 0.1974 | 0.1941 | 0.2153 | 0.2072 | 0.1990 | 0.2039 | 0.1794 | |
26. DQ201428.1 | 0.2202 | 0.2186 | 0.2186 | 0.2235 | 0.2235 | 0.2235 | 0.2235 | 0.2235 | 0.2186 | 0.1958 | 0.1974 | 0.1974 | 0.2202 | 0.2007 | 0.1860 | 0.2055 | 0.1876 | 0.1925 | 0.1860 | 0.1974 | 0.1925 | 0.2153 | 0.1860 | 0.1876 | 0.1892 |
Both molecular analysis (Figure
The intraspecific distance (less than 1%), is significantly less than interspecific distance (more than 10%), so the COI can be used for species identification. The intraspecific genetic polymorphism of Lithobius (E.) longibasitarsus sp. n. and Lithobius (E.) datongensis sp. n.) is very low and could indicate weak migration and diffusion capacity with strong natural selection. Assuring the monophyly and interrelationships of the many genera and subgenera belong to Lithobiidae needs further intensive study including more diverse sampling and molecular evidence, the direction of our future effects.
We are grateful to the National Natural Science Foundation of China (No. 31370405) and the Construction Fund for Qinghai Key Laboratories (2017-ZJ-Y23), which supported our Project. We thank Weiping Li and Gonghua Lin for collecting the specimens. We are grateful to reviewers Gregory D. Edgecombe, Ivan H. Tuf, and Marzio Zapparoli for their hospitality and continued assistance during our research. We thank editor Nathalie Yonow for help in preparing the paper.
Species used for CO1 sequence analysis, sequence references, GenBank accession numbers, voucher, and locality.
Taxa | Sequence reference | GenBank No. | Voucher | Locality |
---|---|---|---|---|
Lithobiidae | ||||
Lithobiinae | ||||
Lithobius (Monotarsobius) crassipes |
|
MF123710.1 |
|
France, |
Lithobius (L.) forficatus |
|
MF123702 |
|
Germany |
Lithobius (L.) variegatus rubriceps |
|
AF334311 |
|
Spain |
Lithobius (L.) castaneus |
|
HM453305 |
|
Italy |
Lithobius (Ezembius) giganteus |
|
HM453306 |
|
Kazakhstan |
Lithobius (E.) longibasitarsus sp. n. | this paper | MH05602 | GH04 | China |
Lithobius (E.) longibasitarsus sp. n. | this paper | MH05603 | GH05 | China |
Lithobius (E.) longibasitarsus sp. n. | this paper | MH05604 | GH03 | China |
Lithobius (E.) longibasitarsus sp. n. | this paper | MH05605 | GH011 | China |
Lithobius (E.) longibasitarsus sp. n. | this paper | MH05606 | GH06 | China |
Lithobius (E.) datongensis sp. n. | this paper | MH05607 | DT4 | China |
Lithobius (E.) datongensis sp. n. | this paper | MH05608 | DT6 | China |
Lithobius (E.) datongensis sp. n. | this paper | MH05609 | DT5 | China |
Lithobius (M.) holsti |
|
HM453307 |
|
Japan |
Australobius scabrior |
|
DQ201428 | Australia | |
Ethopolyinae | ||||
Eupolybothrus tridentinus |
|
JN269950.1 | BC |
Croatia |
Bothropolys xanti |
|
HM453308 | Bmultide | USA |
Henicopidae | ||||
Anopsobiinae | ||||
Anopsobius neozelanicus |
|
AF334313.1 |
|
New Zealand |
Henicopinae | ||||
Henicopini | ||||
Henicops maculatus |
|
AF334316.1 |
|
Australia |
Lamyctes coeculus |
|
AF334315.1 |
|
Australia |
Lamyctes emarginatus |
|
KX442654.1 |
|
Germany |
Lamyctes inermipes |
|
AY214425.1 |
|
Argentina |
Lamyctes hellyeri |
|
AY214428.1 |
|
Australia |
Paralamyctes (P.) harrisi |
|
AF334320 |
|
New Zealand |
P. (Thingathinga) validus |
|
AF334330 |
|
New Zealand |
Zygethobiini | ||||
Cermatobius japonicus |
|
AF334332 |
|
Japan |