Research Article |
Corresponding author: Tongzuo Zhang ( zhangtz@nwipb.cas.cn ) Academic editor: Marzio Zapparoli
© 2019 Penghai Qiao, Wen Qin, Huiqin Ma, Gonghua Lin, Tongzuo Zhang.
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, Lin G, Zhang T (2019)
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Lithobius (Ezembius) hualongensis sp. nov. and Lithobius (Ezembius) sui sp. nov. (Lithobiomorpha, Lithobiidae) recently discovered from the Qinghai-Tibet Plateau, China are described. Morphologically, the two new species are very similar but can be distinguished by the number of coxosternal teeth: L. (E.) hualongensis sp. nov. has 2 + 2 while L. (E.) sui sp. nov. has 3 + 3. The two new species resemble L. (E.) multispinipes
myriapod, Qinghai-Tibet Plateau, stone centipede, taxonomy
The myriapod fauna of China is still poorly known and this is especially the case with centipedes of the order Lithobiomorpha. Only about 84 species/subspecies of lithobiomorphs are known from the country (
The centipede subgenus Ezembius was erected by
Ezembius is characterized by antennae with ca 20 articles; ocelli 1 + 4 to 1 + 20; forcipular coxosternal teeth usually 2 + 2, sometimes 2 + 3, 3 + 2 or 3 + 3; porodonts generally setiform but sometimes stout; tergites generally without posterior triangular projections, occasionally with; tarsal articulation of legs 1–13 distinct; female gonopods with uni-, bi- or tridentate claw, 2 + 2 or 3 + 3, rarely 4 + 4, spurs (
All specimens were hand-collected under leaf litter or stones. The material was examined with the aid of a Motic-C microscope of which the measuring accuracy is +/- 0.01 mm. The color description is based on specimens in 75% ethanol, and body length is measured from the anterior margin of the cephalic plate to the posterior end of the postpedal tergite. Type specimens are preserved in 75% ethanol and deposited in Northwest Institute of Plateau Biology, Chinese Academy of Sciences. The terminology of the external anatomy follows
Subfamily Lithobiinae Newport, 1844
Genus Lithobius Leach, 1814
Subgenus Ezembius Chamberlin, 1919
Holotype : ♂ (HL9), Hualong Hui Autonomous County, Qinghai Province, 36.18848333N, 102.2971333E, 3185 m, a.s.l., 14 April 2012, leg. Lin Gong-Hua, Li Wei-Ping. Paratypes: 1 ♀ (HL7), 2 ♂♂ (HL1, HL4), all from the same locality.
A Lithobius (Ezembius) species with body length 12.31–16.15 mm, antennae of 20 + 20 articles; 8–11 ocelli on each side, arranged in 3 irregular rows, terminal two ocelli comparatively large; Tömösváry’s organ distinctly larger than the adjoining ocelli; 2 + 2 coxosternal teeth; porodonts posterolateral and ventral to the lateral-most tooth; posterior angles of all tergites without triangular projections; 4–7 coxal pores oval to round, arranged in one row; female gonopods with 3 + 3 moderately large, coniform spurs; terminal claw of the third article simple, with a very small triangular protuberance on basal ventral side; male gonopods short and small, with one long setae on the terminal segment.
Male (Fig.
Lithobius (Ezembius) hualongensis sp. nov. A, C, D holotype, male: A habitus, dorsal view C forcipular coxosternite, ventral view D male posterior segments and gonopods, ventral view B, E, F paratype, female: B ocelli and Tömösváry’s organ (To), lateral view E female gonopods, dorsal lateral view F female posterior segments and gonopods, ventral view. Scale bars: 1 mm (A, C); 500 μm (D, F); 250 μm (B, E).
Coloration. Body and antennae reddish brown. Pleural region pale grey. Sternites yellow-brown. Distal part of forcipules red-brown, with basal and proximal parts of forcipules and forcipular coxosternite yellow-brown. Legs 1–15 pale yellow-brown.
Antennae composed of 20 + 20 articles extending back to anterior part of T3, basal article about the same width as length, second article longer than wide, third article slightly longer than wide, with following articles tapering, distal-most article 2.7 times as long as wide; abundant setae on antennal surface, gradual increase in density of setae to about 4th article, then more or less constant.
Cephalic plate
smooth, cordiform. Frontal marginal ridge of head with shallow anterior median furrow. Lateral marginal ridge discontinuous. Posterior margin continuous, convex (Fig.
On each side of head, 1 + 3, 3, 1 oval to rounded ocelli arranged in three irregular rows; posterior ocellus large; ocelli adjacent to the Tömösváry organ slightly smaller. Seriate ocelli domed, translucent, usually darkly pigmented. Tömösváry organ at anterolateral margin of the cephalic plate, larger than the adjacent ocelli.
Coxosternite subtrapezoidal
(Fig.
All tergites with numerous minute setae scattered on surface, several setae on anterior and posterior angles of each tergite and lateral borders, dorsum slightly convex; T1 slightly narrower posterolaterally than anterolaterally, generally trapezoidal, slightly narrower than the cephalic plate and T3, cephalic plate slightly about the same size as T3. Lateral marginal ridges of all tergites continuous. Posterior marginal ridges of TT 1, 3 and 5 concave, continuous, posterior marginal ridges of TT 7, 8, 10, 12 and 14 slightly concave, discontinuous. Posterior angles of all tergites rounded, without triangular projections.
Sternites smooth, posterior part of sternites narrower than anterior, generally trapezoidal. Sternites with 8 short to long setae on anterior corners and anterior lateral borders, 2 setae on posterior lateral borders.
Legs
slender, tarsal articulation well defined on legs 1–15. All legs with fairly long curved claws; pretarsus of legs 1–13 with a slightly curved, long, principal claw and anterior and posterior accessory spines, anterior accessory spines slightly longer and slender, ca 0.56 the length of principal claw, posterior one stouter, ca 0.43 the length of principal claw, forming slightly larger angles with tarsal claws; leg 14 with only posterior spines; leg 15 lacking accessory spines. Dense glandular pore on surface of femur, tibia and tarsi of legs 14 and 15. Long setae sparsely scattered over surface of prefemur, femur, tibia, and tarsi of legs 14–15, more setae on the tarsal surface. 6–7 thicker setae arranged in one row on the ventral surface of tarsus 1 of legs 1–13, 6–7 pairs of thicker setae arranged in two rows on the ventral surface of tarsus 2 of legs 1–13. Legs 14 and 15 thicker and stronger than the anterior pairs. 15th leg 40.6% of body length, tarsus 1 3.3 times longer than wide, tarsus 2 44.4% length of tarsus on leg 15. No modification on legs 14 and 15 in males. Leg plectrotaxy as in Table
Lithobius (Ezembius) hualongensis sp. nov.: plectrotaxy of legs. Letters in brackets indicate spines on one leg of pair, or in one specimen.
Legs | Ventral | Dorsal | ||||||||
C | Tr | P | F | Ti | C | Tr | P | F | Ti | |
1 | – | – | mp | amp | am | – | – | mp | ap | a |
2–4 | – | – | (a)mp | amp | am | – | – | (a)mp | ap | ap |
5 | – | – | amp | amp | am | – | – | (a)mp | ap | ap |
6–11 | – | – | amp | amp | am | – | – | amp | ap | ap |
12 | – | m | amp | amp | am | a | – | amp | ap | ap |
13 | – | m | amp | amp | am | a | – | amp | ap | ap |
14 | – | m | amp | amp | am | a | – | amp | p | p |
15 | – | m | amp | amp | a | a | – | amp | p | – |
Coxal pores 4655 round or slightly oval, variable in size, arranged in a row. 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.
Male posterior segment. Male S15 subtrapeziform, posterior margin narrower than anterior, sparsely covered with short to long setae on ventral side of S15 and lateral and posterior borders (Fig.
Female posterior segment. Female S15 anterior margin broader than posterior, generally trapezoidal, posteromedially straight, S15 with short to long setae on the ventral surface and lateral and posterior borders. Posterior margin of genital sternite deeply concave between condyles of gonopods, except for a small, median rhomboid bulge. Short to long setae sparsely scattered on ventral surface of genital segment. Gonopods: first article fairly broad, bearing 17–20 short to moderately long setae, arranged in four irregular rows; with 3 + 3, moderately long and slender spurs, inner spur smaller than the outer (Fig.
Body length 12.31–16.15 mm; ocelli 1 + 3, 3, 1 or 1 + 4, 3, 3; coxal pores 6666 and 6777 in female and 4654 and 4655 in male.
Morphologically, the new species can be easily distinguished from the seven species in the subgenus from Qinghai Province, L. (E.) asulcutus, L. (E.) rarihirsutipes, L. (E.) femorisulcutus, L. (E.) longibasitarsus, L. (E.) datongensis, L. (E.) maqinensis and L. (E.) dulanensis, by the 3 + 3 coniform spurs on female gonopods contrary to 2 + 2 coniform spurs (Table
Main morphological characters of Chinese species of subgenus Lithobius (Ezembius) Chamberlin, 1919 from Qinghai Province.
Characters | asulcutus | rarihirsutipes | femorisulcutus | longibasitarsus | datongensis | maqinensis | dulanensis | hualongensis sp. nov. | sui sp. nov. |
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Sources |
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this paper | this paper |
Body length (mm) | 13–15 | 11–12 | 15–18 | 17–18 | 12.3–14.2 | 13.1–14.6 | about 20.5 | 12.31–16.15 | 12.15–18.85 |
Number of antennal articles | 20 + 20 | 20 + 20 | 20 + 20 | 20 + 20 | 20 + 20 | 20 + 20 | 20–21 | 20 + 20 | 20 + 20 |
Number, arrangement of ocelli | 10, in 3 rows | 11, in 3 rows | 10–14, in 3 rows | 10–14, in 3 rows | 9–10 ocelli, in 3 broken rows | 10–12, in 3 rows | 11, in 3 rows | 8–11, in 3 rows | 9–10, in 3 rows |
Posterior ocellus | round, comparatively large | oval to round, large | comparatively large | posterior ocellus is biggest | slightly larger than posterosuperior ocellus | posterior ocellus is biggest | posterior ocellus and posterosuperior ocellus comparatively large | terminal two ocelli comparatively large | terminal two ocelli comparatively large |
Seriate ocelli | ones near ventral margin moderately small | ones near ventral margin moderately small | ones near ventral margin moderately small | ones near ventral margin moderately small | ones near ventral margin moderately small | ones near ventral margin moderately small | second row smaller than first, third smallest | ones near ventral margin moderately small | ones near ventral margin moderately small |
Tömösváry’s organ | round, smaller than adjoining ocelli | rounded, slightly, smaller than adjoining ocelli | slightly larger than adjoining ocelli | slightly smaller than adjoining ocelli | larger than adjoining ocelli | almost same size as adjacent ocelli | oval and slightly smaller than adjoining ocelli | obviously larger than adjoining ocelli | obviously larger than adjoining ocelli |
Number and shape of coxosternal teeth | 2 + 2 subtriangular teeth | 2 + 2, subtriangular teeth | 2 + 2 | 2 + 2 or 3 + 2 | 2 + 2 | 2 + 2, small coniform teeth | 2 + 2, coniform, moderately robust teeth | 2 + 2, small coniform teeth | 3 + 3, inner tooth smaller than outer tooth |
Porodont | long and slender, lying posterolateral to lateral-most teeth | long and slender, lying posterolateral to lateral-most tooth | long and strong, lying posterolateral to lateral-most tooth | thick and strong separated from lateral tooth ventrolaterally | setiform, separated from lateral tooth laterally | setiform, lying posterolateral to lateral-most tooth | long and strong, lying posterolateral to lateral-most tooth | thick and long, lying posterolateral to lateral-most tooth | thick and long, lying posterolateral to lateral-most tooth |
Tergites | smooth | smooth | smooth | smooth | smooth | smooth | smooth | with numerous minute setae scattered on surface | with numerous minute setae scattered on surface |
Number of coxal pores | 4–7, 4544, 4554, 5665, 5766 | not reported | 5544, 5554, 5555, 5564 | 4–6 | 4–7 | 4–6 | 5–7, 5667, 5666 | 6666 and 6777 in female and 4654 and 4655 in male | 5664, 5665, 7775 and 8875 in female and 6886 and 7665 in male. |
Shape of coxal pores | not reported | not reported | round | circular | round | round | circular to ovate | round or slightly ovate | round or slightly ovate |
Tarsus 1–tarsus 2 articulation on legs 1–13 | not well defined | well defined | well defined | well defined | well defined | not well defined | not well defined | well defined | well defined |
Male 14th leg | slightly thicker and stronger than other legs | markedly thicker and stronger than 1–13 legs | slightly thick and strong than 1–13 legs | moderately thick and strong than 1–13 legs | moderately thick and strong than 1–13 legs | moderately thick and strong than 1–13 legs | slightly thick and strong than 1–13 legs | slightly thick and strong than 1–13 legs | slightly thick and strong than 1–13 legs |
Male 15th leg | slightly thicker and stronger than other legs | markedly thicker and stronger than 1–13 legs | slightly thick and strong than 1–13 legs | moderately thick and strong than 1–13 legs | moderately thick and strong than 1–13 legs | moderately thick and strong than 1–13 legs | slightly thick and strong than 1–13 legs | slightly thick and strong than 1–13 legs | slightly thick and strong than 1–13 legs |
Dorsal sulci on male 15th legs | absent | two distinct, dorsal sulci on femur |
distinct, dorsal sulcus on tibia and tarsus 1 |
absent | absent | absent | absent | absent | absent |
DaC spine | on 12th–15th legs | on 12th–15th legs | on 11th–15th legs | on 12th–15th legs | on 12th–15th legs | on 11th–15th legs | on 11th–15th legs | on 12th–15th legs | on 12th–15th legs |
14th accessory spur | present | present | present | present | present | anterior accessory spur absent, posterior accessory spur present |
anterior accessory spur absent, posterior accessory spur present |
anterior accessory spur absent, posterior accessory spur present |
anterior accessory spur absent, posterior accessory spur present |
15th accessory spur | absent | absent | present | present | present | absent | absent | absent | anterior accessory spur absent, posterior accessory spur present |
Number and shape of spurs on female gonopods | 2 + 2 coniform spurs | 2 + 2 conical spurs | 2 + 2 conical spurs | 2 + 2 conical spurs | 2 + 2 conical spurs | 2 + 2 conical spurs | 2 + 2 conical spurs | 3 + 3 coniform spurs, inner spur moderately smaller than outer one | 3 + 3 coniform spurs, inner spur moderately smaller than outer one |
Dorsal side of second article of female gonopods | not reported | not reported | not reported | four short setae and three long setae on dorsolateral ridge | five long curved spines on dorsolateral side | not reported | with six dorsolateral setae | with 10 short to long dorsal lateral setae, stouter than general setae | with 10 short dorsal lateral setae, stouter than general setae |
Apical claw of female gonopods (and lateral denticles) | simple | simple, having small sharp teeth on the inner side | simple | simple, having small triangular protuberance on ventral side | simple, bearing very small triangular protuberance on ventral side | simple, having very small triangular protuberance on ventral side | simple | simple, having very small triangular protuberance on ventral side | simple, having very small triangular protuberance on ventral side |
Male gonopods | short and small bulge, with two long setae | with small bulge, with 3 long setae | not reported | single small semicircular article with 3–5 setae on its surface | hemispherical, with three setae | small, oblique apically, with 2 setae | small, one-segmented, with two long setae | short and small bulge, having long seta, apically slightly sclerotized | short and small, with two long setae on terminal segment |
The new species is named from the type locality.
The four specimens here examined (3 ♂♂, 1 ♀) were collected under granular gravel on the alpine meadows composed mainly of Poaceae, Cyperaceae, Fabaceae, Polygonaceae, Chenopodiaceae, Asteraceae, Rosaceae, Liliaceae and Cucurbitaceae. The region is located on the upper reaches of the Yellow River Valley and features an arid climate, with mean annual precipitation 451.2 mm and average annual temperature 2.8 °C (http://data.cma.cn/data/weatherBk.html).
Holotype : ♀ (MHA8), Minhe County, Qinghai Province, 36.12076N, 102.7809E, 2280 m, a.s.l., 24 October 2010, leg. Lin Gong-Hua. Paratypes: 1 ♂ (MHA6), 2♀♀ (MHA1, MHA5), all from the same locality.
A Lithobius (Ezembius) species with body length 12.15–18.85 mm, antennae of 20 + 20 articles; 9–10 ocelli on each side, arranged in 3 irregular rows, terminal two ocelli comparatively large; Tömösváry’s organ distinctly larger than the adjoining ocelli; 3 + 3 coxosternal teeth; porodonts posterolateral and ventral to the lateral-most tooth; posterior angles of all tergites without triangular projections; 4–8 coxal pores oval to round, arranged in one row; female gonopods with 3 + 3 moderately large, coniform spurs; terminal claw of the third article simple, with a very small triangular protuberance on basal ventral side; male gonopods short and small, with two long setae on the terminal segment.
Female (Fig.
Lithobius (Ezembius) sui sp. nov. A, B, D, E, G holotype, female: A habitus, dorsal view B ocelli and Tömösváry’s organ (To), lateral view D forcipular coxosternite, ventral view E female posterior segments and gonopods, ventral view G female gonopods, dorsal lateral view C, F paratype, male: C ocelli and Tömösváry’s organ (To), lateral view F posterior segments and gonopods, ventral view.Scale bars: 1 mm (A, D); 500 μm (E, F); 250 μm (B, C, G).
Coloration. Body yellow-brown, cephalic plate and antennae light yellow-brown with reddish hue. Pleural region pale grey. Sternites yellow-brown. Distal part of forcipules red-brown, with basal and proximal parts of forcipules and forcipular coxosternite yellow-brown. Legs 1–15 light yellow-brown.
Antennae composed of 20 + 20 articles extending back to posterior part of T3, basal article about the same width as length, second article slightly longer than wide, third article slightly wider than long, with following articles tapering, distal-most article 2.9 times as long as wide; abundant setae on antennal surface, gradual increase in density of setae basally to distally to about fourth articles.
Cephalic plate
smooth, convex. Frontal marginal ridge of head with shallow anterior median furrow. Lateral marginal ridge discontinuous. Posterior margin continuous, slightly convex (Fig.
On each side of head, 1 + 4, 4, 1 oval to rounded ocelli (Fig.
Coxosternite subtrapezoidal
(Fig.
All tergites with numerous minute setae scattered on surface, several setae on anterior and posterior angles of each tergite and lateral borders, dorsum slightly convex; T1 narrower posterolaterally than anterolaterally, generally trapezoidal, narrower than the cephalic plate and T3, cephalic plate slightly wider than T3. Lateral marginal ridges of all tergites continuous. Posterior marginal ridges of TT 1 and 3 slightly concave, continuous, posterior marginal ridges of TT 5, 7, 8, 10, 12 and 14 concave, discontinuous. Posterior angles of all tergites rounded, without triangular projections.
Sternites smooth, posterior part of sternites narrower than anterior, generally trapezoidal. Sternites with 2–7 short to long setae on anterior corners and anterior lateral borders, the same with posterior lateral and posterior angles.
Legs
slender, tarsal articulation well defined on legs 1–15. All legs with fairly long curved claws; pretarsus of legs 1–13 with a slightly curved, long, principal claw and anterior and posterior accessory spines, anterior accessory spines slightly longer and slender, ca 0.50 the length of principal claw, posterior one stouter, ca 0.33 the length of principal claw, forming slightly larger angles with tarsal claws; legs 14 and 15 with only posterior spines. Dense glandular pore on surface of femur, tibia and tarsi of legs 14 and 15. Long setae sparsely scattered over surface of prefemur, femur, tibia, and tarsi of legs 14–15, more setae on the tarsal surface. 7–9 thicker setae arranged in one row on the ventral surface of tarsus 1 of legs 1–13, 7–8 pairs of thicker setae arranged in two rows on the ventral surface of tarsus 2 of legs 1–13. Legs 14 and 15 thicker and stronger than the anterior pairs. 15th leg 39.24% of body length, tarsus 1 3.8 times longer than wide, tarsus 2 44.4% length of tarsus on leg 15. Leg plectrotaxy as in Table
Lithobius (Ezembius) sui sp. nov.: plectrotaxy of legs. Letters in brackets indicate spines on one leg of pair, or in one specimen.
Legs | Ventral | Dorsal | ||||||||
C | Tr | P | F | Ti | C | Tr | P | F | Ti | |
1 | – | – | mp | amp | am | – | – | (a)mp | ap | a |
2–4 | – | – | mp | amp | am | – | – | (a)mp | ap | ap |
5–7 | – | – | mp | amp | am | – | – | amp | ap | ap |
8–11 | – | – | amp | amp | am | – | – | amp | ap | ap |
12 | – | m | amp | amp | am | a | – | amp | ap | ap |
13 | – | m | amp | amp | am | a | – | amp | (a)p | ap |
14 | – | m | amp | amp | am | a | – | amp | p | p |
15 | – | m | amp | amp | a | a | – | amp | p | – |
Coxal pores 4–8 round or slightly oval, variable in size, arranged in a row. 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. Female S15 anterior margin broader than posterior, generally trapezoidal, posteromedially straight, S15 with short to long setae on the ventral surface and lateral and posterior borders. Posterior margin of genital sternite deeply concave between condyles of gonopods, except for a small, median rhomboid bulge. Short to long setae sparsely scattered on ventral surface of genital segment. Gonopods: first article fairly broad, bearing 9–11 short to moderately long setae, arranged in three irregular rows; with 3 + 3, moderately long and slender, bullet-shape spurs, inner spur smaller than the outer (Fig.
Male posterior segment. Male S15 subtrapeziform, posterior margin narrower than anterior, sparsely covered with short to long setae on ventral side of S15 and lateral and posterior borders (Fig.
Body length 12.15–18.85 mm; ocelli 1 + 4, 4, 1, 1 + 4, 3, 2 or 1 + 4, 3, 1; coxal pores 5664, 5665, 7775 or 8875 in female and 6886, 7665 in male.
The two new species are very similar in morphology, especially in both having numerous minute setae scattered on surface of all tergites, but can be distinguished by the number of coxosternal teeth: L. (E.) hualongensis sp. nov. has 2 + 2 while L. (E.) sui sp. nov. has 3 + 3.
The specific name is a patronym in honor of the zoologist Dr Jianping Su, Academician at the Chinese Academy of Sciences.
The four specimens here examined (1 ♂, 3 ♀♀) were collected under granular gravel on the alpine meadows composed mainly of Gramineae, Cyperaceae, Leguminosae, Polygonaceae, Chenopodiaceae, Compositae, Rosaceae, Liliaceae and Cucurbitaceae. This region adjacent to Hualong County in the west features plateau continental, with mean annual precipitation 338.1 mm and average annual temperature 8.3 °C (http://data.cma.cn/data/weatherBk.html).
1 | Posterior angles of tergites with triangular projections | L. (E.) sulcipes Attems, 1927 |
– | Posterior angles of tergites rounded, without projections | 2 |
2 | 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 | 3 |
3 | Cephalic plate with scattered, rough puncta and tergite with distinct puncta | L. (E.) rhysus Attems, 1934 |
– | Cephalic plate and tergite without any puncta | 4 |
4 | All ocelli subequal in size | L. (E.) sulcifemoralis Takakuwa & Takashima, 1949 |
– | All ocelli not subequal in size | 5 |
5 | Terminal two ocelli comparatively large | 6 |
– | Terminal one ocellus comparatively large | 13 |
6 | Ocelli arranged in two rows | L. (E.) laevidentata Pei, Ma, Hou, Zhu & Gai, 2015 |
– | Ocelli arranged in three rows | 7 |
7 | 3 + 3 coxosternal teeth | 8 |
– | 2 + 2 coxosternal teeth | 9 |
8 | First article of female gonopods with 3 + 3 spurs | L. (E.) sui sp. nov. |
– | First article of female gonopods with 2 + 2 spurs | L. (E.) multispinipes Pei, Lu, Liu, Hou, Ma & Zapparoli, 2016 |
9 | Tömösváry’s organ larger than adjoining ocellus | 10 |
– | Tömösváry’s organ smaller than adjoining ocellus | 11 |
10 | First article of female gonopods with 3 + 3 spurs | L. (E.) hualongensis sp. nov. |
– | First article of female gonopods with 2 + 2 spurs | L. (E.) bilineatus Pei, Ma, Zhu & Gai, 2014 |
11 | Apical claw of female gonopods simple, without inner small subtriangular teeth | L. (E.) tetraspinus Pei, Lu, Liu, Hou & Ma, 2018 |
– | Apical claw of female gonopods simple, with inner small subtriangular teeth | 12 |
12 | Number of antennal articles 23 + 23 | L. (E.) anabilineatus Ma, Pei, Hou, Zhu & Gai, 2015 |
– | Number of antennal articles 20 + 20–21 + 21 | L. (E.) dulensis Qiao, Ma, Pei, Zhang & Su, 2019 |
13 | 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 | 14 |
14 | Tömösváry’s organ smaller than adjoining ocellus. | 15 |
– | Tömösváry’s organ larger or subequal in size as adjoining ocellus | 21 |
15 | First article of female gonopods with 3 + 3 or 4 + 4 spurs | 16 |
– | First article of female gonopods with 2 + 2 spurs | 17 |
16 | Apical claw of female gonopods simple, with innersmall subtriangular teeth | L. (E.) bidens Takakuwa, 1939 |
– | Apical claw of female gonopods simple, without inner small subtriangular teeth | L. (E.) insolitus Eason, 1993 |
17 | Terminal claw of female gonopods bipartite | L. (E.) anasulcifemoralis Ma, Pei, Wu & Gai, 2013 |
– | Terminal claw of female gonopods simple | 18 |
18 | Terminal claw of female gonopods simple, with inner small triangular teeth | 19 |
– | Terminal claw of female gonopods simple, without inner small triangular teeth | 20 |
19 | Body length 11–12 mm, 15th accessory spur absent | L. (E.) rarihirsutipes Zhang, 1996 |
– | Body length 17–18 mm, 15th accessory spur present | L. (E.) longibasitarsus Qiao, Qin, Ma, Zhang, Su & Lin, 2018 |
20 | DaC spine on 12th –15th legs | L. (E.) asulcutus Zhang, 1996 |
– | DaC spine absent | L. (E.) giganteus Sseliwanoff, 1881 |
21 | Six ocelli on each side of cephalic plate | L. (E.) gantoensis Takakuwa & Takashima, 1949 |
– | At least seven ocelli on each side of cephalic plate | 22 |
22 | Ocelli arranged in two rows | L. (E.) irregularis Takakuwa & Takashima, 1949 |
– | Ocelli arranged in three rows | 23 |
23 | First article of female gonopods with 3 + 3 spurs | 24 |
– | First article of female gonopods with 2 + 2 spurs | 25 |
24 | DaC spine on 14th–15th legs | L. (E.) lineatus Takakuwa, 1939 |
– | DaC spine on 12th–15th legs | L. (E.) mandschreiensis Takakuwa, 1940 |
25 | Terminal claw of female gonopods tridentate | L. (E.) zhui Pei, Ma, Shi, Wu & Gai, 2011 |
– | Terminal claw of female gonopods simple | 26 |
26 | Terminal claw of female gonopods simple, without inner small triangular teeth | L. (E.) femorisulcutus Zhang, 1996 |
– | Terminal claw of female gonopods simple, with inner small triangular teeth | 27 |
27 | Tarsal articulation on legs 1–13 well defined, 14th accessory spur present | L. (E.) datongensis Qiao, Qin, Ma, Zhang, Su & Lin, 2018 |
– | Tarsal articulation on legs 1–13 not well defined, only 14th posterior accessory spur present | L. (E.) maqinensis Qiao, Qin, Ma & Zhang, 2019 |
This work was financially supported by the the Strategic Priority Research Program of the Chinese Academy of Sciences [XDA23060602, XDA2002030302]; National Key R&D Program of China [2017YFC0506405]; and Qinghai Key R&D and Transformation Program [2019-SF-150].
We are grateful to Dr Gregory D. Edgecombe, London, U. K., Dr Ivan H. Tuf, Olomouc, Czech Republic, Dr Pavel Stoev, Sofia, Bulgaria and Dr Marzio Zapparoli, Viterbo, Italy for their hospitality and everlasting help during our research, respectively. We thank Dr Rowland M. Shelley, North Carolina, USA, and Dr His-Te Shih, Taichung, China, for providing us with invaluable literature.