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Research Article
First report of Lepidosira (Collembola, Entomobryidae) from China, with description of four new species under the aid of COI barcoding
expand article infoXiaowei Qian‡, Meidong Jing‡, Yitong Ma‡
‡ Nantong University, Nantong, China

Corresponding author: Yitong Ma ( mayitong@ntu.edu.cn )

Academic editor: Alexssandro Camargo
© 2025 Xiaowei Qian, Meidong Jing, Yitong Ma.
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: Qian X, Jing M, Ma Y (2025) First report of Lepidosira (Collembola, Entomobryidae) from China, with description of four new species under the aid of COI barcoding. Deutsche Entomologische Zeitschrift 72(2): 341-365. https://doi.org/10.3897/dez.72.153961
ZooBank: urn:lsid:zoobank.org:pub:E4AB50E6-895B-44FC-A994-078B3FF27271
Open Access

Abstract

In this study, species of Lepidosira Schött, 1925, from China are studied based on molecular markers and morphological characters. Four new species, Lepidosira apigmenta sp. nov., Lepidosira similis sp. nov., Lepidosira wuxiensis sp. nov., and Lepidosira chongqingensis sp. nov., are described, and their COI barcodes are provided. This is the first report of the genus from China. Willowsia flava Zhou, Pan & Ma, 2021, is transferred to Lepidosira, and the replacement name Lepidosira baojiensis comb. et nom. nov. is proposed since the specific epithet is preoccupied by the senior homonym Lepidosira flava (Salmon, 1938). The transfer of Willowsia trifascia Zhou & Ma, 2022 to Lepidosira is also herein proposed. A key to the scaled genera of the subfamily Entomobryinae is provided.

Key Words

Chaetotaxy, Chongqing, cryptic species, molecular markers, new species, springtails

Introduction

With the development of molecular biology and the discovery of new morphological characters, great changes have taken place in the classification of the family Entomobryidae in the past decade. The tribe Willowsiini was abandoned (Zhang and Deharveng 2015), and the family Entomobryidae was divided into six subfamilies, Entomobryinae, Lepidocyrtinae, Paronellidinae, Paronellinae, Salininae, and Seirinae (Godeiro et al. 2023). Among these six subfamilies, Entomobryinae is the largest, comprising 31 genera. Of these, 15 genera are devoid of scales on their body, while 16 genera possess scales. Among the scaled genera, the shape and distribution of scales are important for taxonomy at the genus level.

The genus Lepidosira was established by Schött in 1925, and its main characters include scales on the manubrium and dentes, dorsal chaetotaxy of the head and trunk polymacrochaetotic, a specialized microchaeta on abdominal segment I external to the m5 macrochaeta, a manubrial plate without blunt chaetae, and a bidentate mucro with a basal spine (Nunes et al. 2019). It is very close to the genus Lepidocyrtoides Schött, 1917, and the main difference between them is the blunt macrochaetae on the manubrium, which are absent in the former and present in the latter. Lepidosira is also similar to Willowsia Shoebotham, 1917, but scales are present on the dens in Lepidosira and absent in Willowsia.

To date, 57 species of the genus have been described worldwide (Bellinger et al. 1996–2024), and about 40 of them are from Oceania. The remaining species are from Brazil (L. neotropicalis Nunes & Bellini, 2019), India (L. unguserrata Salmon, 1970 and L. pallida (Ritter, 1911)), Indonesia (L. calolepis (Börner, 1913), L. javana (Börner, 1913), and L. sundana Yoshii & Suhardjono, 1989), Japan (L. gigantea (Börner, 1909)), Vietnam (L. alba (Nguyên, 2005) and L. nigropunctata (Nguyên, 2005)), and Rwanda and Burundi (L. congoia Salmon, 1956). The original descriptions of many of the early described species do not include body chaetotaxy and were mainly based on their color pattern. Here, we describe four new species of Lepidosira from Chongqing Municipality, China, representing the first report of the genus from China. Detailed descriptions of body chaetotaxy and COI sequences are provided for the new species.

Materials and methods

Survey area

Specimens were collected in the Yintiaoling National Nature Reserve, located in the northeast of Chongqing Municipality, which belongs to southwestern China. It has a subtropical humid monsoon climate with an annual average temperature of about 10 °C and an annual average precipitation of 1,500 mm. The total area of the reserve is about 22,000 hectares, and its highest point is 2,798 m above sea level (Zhang 2025).

Taxon sampling and specimen examination

Specimens were collected with an aspirator from leaf litter and stored in 99% alcohol. They were mounted on glass slides in Marc André II solution (Christiansen 1990) and studied with a Leica DM2500 phase-contrast microscope. Photographs were taken using a Leica DFC300 FX digital camera mounted on the microscope and enhanced with Photoshop CS2 (Adobe Inc.). Type specimens were deposited in the School of Life Sciences, Nantong University, Jiangsu, China.

The nomenclature of the dorsal macrochaetotaxy of the head and interocular chaetae follows Jordana and Baquero (2005) and Mari-Mutt (1979, 1986). Labial chaetae are designated following Gisin (1964). Labral chaetae follow Szeptycki (1973), and trunk dorsal chaetotaxy follows Szeptycki (1979), adapted from Zhang et al. (2019).

Abbreviations

Ant. antennal segment(s),

Th. thoracic segment(s),

Abd. abdominal segment(s),

mac macrochaeta(e),

mes mesochaeta(e),

ms specialized microchaeta(e),

sens specialized ordinary chaeta(e),

Gr. group.

DNA extraction and amplification

DNA was extracted using an Ezup Column Animal Genomic DNA Purification Kit (Sangon Biotech, Shanghai, China) following the manufacturer’s standard protocols. Amplification of a 658 bp fragment of the mitochondrial COI gene was carried out using a Prime Thermal Cycler (TECHNE, Bibby Scientific Limited, Stone, Staffordshire, UK) in 25 μl reaction volumes with the Premix Taq polymerase system (Takara Bio, Otsu, Shiga, Japan). The primers and polymerase chain reaction (PCR) programs followed Greenslade et al. (2011). All PCR products were checked using 1% agarose gel electrophoresis. Successful products were purified and sequenced on an ABI 3730XL DNA Analyzer (Applied Biosystems, Foster City, CA, USA). All experiments were completed by Shenggong (Shanghai, China). The GenBank accession numbers are provided in Table 1.

Table 1.
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Included species and their respective GenBank accession numbers.

Species Voucher GenBank Data
Lepidosira apigmenta sp. nov. M1311-8 M1321-14 M1321-13 M1321-12 PV282589 PV282590 PV282591 PV282592 This study
L. similis sp. nov. M1311-7 PV282594 This study
L. wuxiensis sp. nov. M1321-9 M1321-10 M1321-11 PV282586 PV282587 PV282588 This study
L. chongqingensis sp. nov. M1311-4 PV282593 This study
L. baojiensis comb. et nom. nov. 1102-3 PV282595 This study
L. calolepis (Börner, 1913) OP094719 GenBank
L. neotropicalis Nunes & Bellini, 2019 MF716603 Personal communication*

* Sequence obtained directly from the authors.

DNA sequences were assembled using Sequencher 4.5 (Gene Codes Corp.) and then deposited in GenBank. Sequences were aligned using ClustalW implemented in MEGA 5.1 (Tamura et al. 2011) with default settings. Pairwise genetic distances were calculated in MEGA 5.1 employing the Kimura 2-parameter (K2-P) model (Kimura 1980). The neighbor-joining tree was constructed in MEGA 5.1 using the neighbor-joining algorithm (Fig. 1).

Figure 1. 

Neighbor joining tree (using K2-P model) of seven Lepidosira species based on COI sequences.

Results

Molecular analysis

In the present study, the shortest interspecific distances were 13.9–15.6% (between L. apigmenta sp. nov. and L. wuxiensis sp. nov.) and the longest were 23.8–25.0% (between L. wuxiensis sp. nov. and L. calolepis) (Table 2). Therefore, the interspecific distances of COI among the seven species were greater than the accepted barcoding gap recently reported for species of Tomocerus (Zhang et al. 2014) and Coecobrya (Zhang et al. 2018). The molecular distances coincided with the morphological divergences, thus further supporting the separation of the four new species.

Table 2.
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Genetic distances (mean K2-P divergence) between Lepidosira species in this study.

Species L. apigmenta sp. nov. L. wuxiensis sp. nov. L. similis sp. nov. L. chongqingensis sp. nov. L. baojiensis nom. nov. L. calolepis (Börner, 1913) L. neotropicalis Nunes & Bellini, 2019
L. apigmenta sp. nov.
L. wuxiensis sp. nov. 13.9–15.6
L. similis sp. nov. 19.8–20.3 21.3–22.5
L. chongqingensis sp. nov. 18.5–19.0 20.0–21.2 19.7
L. baojiensis nom. nov. 19.5–20.0 21.0–22.2 20.7 16.4
L. calolepis (Börner, 1913) 22.3–22.8 23.8–25.0 24.9 23.6 24.6
L. neotropicalis Nunes & Bellini, 2019 20.7–21.2 22.2–23.4 23.3 22.0 23.0 21.0

Taxonomy

Class Collembola Lubbock, 1870

Order Entomobryomorpha Börner, 1913

Family Entomobryidae Tömösvary, 1882

Subfamily Entomobryinae Schäffer, 1896

Key to the scaled genera of the subfamily Entomobryinae

Based mainly on Bellinger et al. (1996–2024).

1 Abd. VI with finger-like projection Epimetrura
– Abd. VI without finger-like projection 2
2 Scales present on dens 3
– Scales absent on dens 7
3 Dental spines present Acanthocyrtus
– Dental spines absent 4
4 Basal ribs of scales longer than distal ones Lepidodens
– Basal ribs of scales almost as long as distal ones 5
5 Scales narrow and tip pointed Lepidobrya
– Scales not narrow and tip not pointed 6
6 Manubrium with distal thick blunt macrochaetae Lepidocyrtoides
– Manubrium without distal thick blunt macrochaetae Lepidosira
7 Dental spines present 8
– Dental spines absent 9
8 Prelabral bifurcate Amazhomidia
– Prelabral not bifurcate Sinhomidia
9 Eyes absent 10
– Eyes present 12
10 Mucro bidentate Szeptyckiella
– Mucro falcate 11
11 Body macrochaetae well developed Hawinella
– Body macrochaetae strongly reduced Lepidosinella
12 Mucro falcate 13
– Mucro bidentate 14
13 Mucronal basal spine absent Desertia
– Mucronal basal spine present Drepanosira
14 Scales chaeta-like and scales of posterior row of tergites strongly elongate Janetschekbrya
– Scales not chaeta-like and scales of posterior row of tergites not strongly elongate 15
15 Scales narrow and with 2 uninterrupted lateral ribs Americabrya
– Scales of different types, but always with internal cilia Willowsia

Lepidosira Schött, 1925

Type species.

Lepidosira terraereginae (Ellis & Bellinger, 1973: 28), by subsequent designation.

Diagnosis.

Scales ciliate and present on the body, including manubrium and dentes; dorsal chaetotaxy of head and trunk polymacrochaetotic; ms on Abd. I external to m5; manubrial plate without blunt chaetae; a bidentate mucro with a basal spine (Nunes et al. 2019).

Lepidosira apigmenta sp. nov.

https://zoobank.org/C78B645B-EAC3-4FB6-ACF9-94F668405948
Figs 2, 3, 4, 5, 6, 7, Table 3

Type material.

Holotype. CHINA • ♀ on slide; Chongqing Municipality, Wuxi County, the Yintiaoling National Nature Reserve, the Guanshan Protection Station, the Stone Pillar; 2,168.92 m a.s.l.; 31°32'15"N, 109°41'54"E; 28 Jul. 2024; Y.T. Ma leg.; sample number 1321 (CQWX0717).

Paratypes. CHINA • 3 ♀ on slides, same data as holotype.

Diagnosis.

Each segment of Ant. I–III distally violet pigmented; Ant. IV and base of Ant. I with scattered violet pigment; Ant. IV with bilobed apical bulb; head with one mac on Gr. II; Th. II with one medio-medial, two medio-sublateral and 15–16 posterior mac; Th. III to Abd. IV respectively with 10, 4, 4, 2, and 9–11 central mac; Abd. III with four lateral mac.

Description.

Body (head + trunk) length up to 1.92 mm, holotype 1.92 mm.

Color pattern : ground color pale yellow; eye patches dark blue; each segment of Ant. I–III distally violet pigmented, Ant. IV and base of Ant. I with scattered violet pigment; a little violet pigment present in post-ocular region and on legs (Fig. 2A, B).

Figure 2. 

Lepidosira apigmenta sp. nov. A, B. Habitus (lateral view). Scale bars: 500 μm.

Scales : ciliate type, present on head, Ant. I–II, legs (Fig. 3A), both faces of ventral tube, ventral side of manubrium and dentes (Fig. 3B, C), and terga (Figs 3D, 4A).

Figure 3. 

SEM photomicrographs of scales of Lepidosira apigmenta sp. nov. A. Leg; B. Manubrium; C. Dens; D. Trunk. Scale bars: 20 μm.

Figure 4. 

Lepidosira apigmenta sp. nov. A. Scale on dorsal trunk; B. Apex of Ant. IV (dorsal view); C. Distal Ant. III (lateral view); D. Distal Ant. II (dorsal view); E. Dorsal head (right side, solid circle representing mac present or absent); F. Prelabral and labral chaetotaxy (dorsal view); G. Maxillary palp and outer lobe (right side); H. Labial palp (right side); I. Labial and post-labial chaetotaxy (right side). Scale bars: 20 μm.

Head : antenna not annulated and 0.45–0.56 (holotype 0.50) times length of body. Ratio of Ant. I–IV as 1.00/1.35–1.82/1.20–1.67/2.25–3.00 (holotype 1.00/1.78/1.65/2.67). Distal part of Ant. IV with many sensory chaetae and normal ciliate chaetae, apical bulb bilobed (Fig. 4B). Ant. III sense organ with two rods, two spiny guard sensilla, smooth blunt sens, and ciliated chaetae (Fig. 4C). Ant. II with two rods apically (Fig. 4D). Eyes 8+8, G and H smaller than others, interocular chaetae as p, q, r, s, t mes, and r absent in one specimen. Dorsal chaetotaxy of head with 10–14 antennal (An) mac or mes, four anterior (A0, A2, A3, A5), four median (M1–4), eight sutural (S0–7) mac, one post-sutural mac (Ps2) and mes (Ps5), 12 posterior mac plus one (Pa1) in Gr. II (Fig. 4E). Prelabral and labral chaetae as 4/5, 5, 4, prelabral chaetae ciliate and other smooth, a2 and b2 slightly shorter than middle ones, labral papillae not clearly seen (Fig. 4F). Basal chaeta on maxillary outer lobe almost as thick as apical one; sublobal plate with four smooth chaetae-like processes (Fig. 4G). Lateral process (l. p.) of labial palp E finger-shaped with tip not or almost reaching apex of papilla E (Fig. 4H). Labial base with M1M2REL1L2, all ciliate (Fig. 4I).

Thorax : tergal ms formula on Th. II–Abd. V as 1, 0/1, 0, 1, 0, 0, sens as 2, 2/1, 2, 2, 2, 3 (Figs 5A, 6A–C). Th. II with about ten anterior, one medio-medial (m1), two medio-sublateral (m4, m4i), 15–16 posterior mac. Th. III with ten central and six lateral mac (Fig. 5A). Coxal chaetal formula as 7–10/8–11 (anterior), 9–12 (posterior)/9–14 (Fig. 5B–D). Trochanteral organ with 36–46 smooth chaetae (Fig. 5E). Tenent hair smooth and clavate, 1.27–1.30 length of inner edge of unguis; unguis with a pair of laterobasal outer teeth and four inner teeth, basal paired inner teeth located at 0.38–0.43 distance from base of inner edge of unguis, distal unpaired inner teeth at 0.67–0.70 and 0.84–0.85 distance from base, respectively; unguiculus lanceolate, outer edge slightly serrate (Fig. 5F).

Figure 5. 

Lepidosira apigmenta sp. nov. A. Chaetotaxy of Th. II−III (right side); B–D. Coxal chaetotaxy of fore, middle, and hind legs (outer view); E. Trochanteral organ (posterior side); F. Hind foot complex (lateral view). Scale bars: 20 μm.

Figure 6. 

Lepidosira apigmenta sp. nov. A. Chaetotaxy of Abd. I−III (right side); B. Chaetotaxy of Abd. IV (right side and partially left side); C. Chaetotaxy of Abd. V (right side); D. Anterior face of ventral tube (T-shape representing scale). Scale bars: 20 μm.

Abdomen : range of Abd. IV length as 4.77–5.50 (holotype 5.50) times as dorsal axial length of Abd. III. Abd. I with four (m2–4, m4i) mac. Abd. II with four (a2, m3, m3e, m3ep) central, one (m5) lateral mac. Abd. III with two (a2, m3) central, four (am6, pm6, m7a, p6) lateral mac (Fig. 6A). Abd. IV with two normal sens, 9–11 central and about 13 lateral mac (Fig. 6B). Abd. V with three sens (Fig. 6C). Anterior face of ventral tube with 3+3 large and about 13 small ciliate chaetae, line connecting proximal (Pr) and external-distal (Ed) mac oblique to median furrow (Fig. 6D); posterior face with two apical smooth chaetae besides about 60 ciliate chaetae of different sizes (Fig. 7A); each lateral flap with 5–6 smooth and 9–10 ciliate chaetae (Fig. 7B). Manubrial plate dorsally with 12–17 ciliate mac and 3–4 pseudopores (Fig. 7C); ventrally with 21–28 ciliate chaetae on each side (Fig. 7D). Mucro bidentate; tip of basal spine reaching apex of subapical tooth; distal smooth section of dens almost equal to mucro in length (Fig. 7E).

Figure 7. 

Lepidosira apigmenta sp. nov. A. Apical chaetae of posterior face of ventral tube; B. Lateral flap of ventral tube; C. Manubrial plaque (dorsal view); D. Ventro–apical part of manubrium; E. Distal part of dens and mucro (lateral view). Scale bars: 20 μm.

Etymology.

The name is composed of the Greek word “a,” meaning absence, and the Latin word “pigmentum,” meaning color. Thus, “apigmenta” refers to the unpigmented trunk.

Habitat.

Found in litter of subtropical forests, mainly composed of leaves of Buxus sinica, Ilex yunnanensis, Pinus armandii and Rosa corymbulosa.

Remarks.

According to Fig. 1 and Table 3, the new species is close to Lepidosira wuxiensis sp. nov. Their antennal color pattern and the chaetotaxy of the dorsal head, Th. II, Abd. I, and Abd. III–IV are also very similar. However, the color pattern of Abd. IV and the chaetotaxy of Th. III and Abd. II are different. The new species is also similar to the Vietnamese species L. alba (Nguyên, 2005) in color pattern, but labral papillae and smooth setae on the posterior face of the ventral tube are present in the former and absent in the latter. The main differences among the three species in chaetotaxy are listed in Table 3.

Table 3.
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Variation of dorsal chaetotaxy of Lepidosira species from Asia and South America (? = not known).

Species Species number Head Th. II Th. III Abd. I Central Abd. II Abd. III Abd. IV
An interocular setae Gr. II Mm posterior central lateral central lateral central lateral
L. apigmenta sp. nov. 1321–11 11 pqrst 1 1 15 10 6 4 4 2 4 10 11+13
1321–12 14 pqrst 1 1 15 10 6 4 4 2 4 11 ?
1321–13A 10+11 pqrst 1 1 16 9+10 6 4 4 2 4 11 ?
1321–13B ? pqrst 1 1 15 10 6 4 4 2 4 9 16
L. similis sp. nov. 1306–27 8 pvt 1 3 15 11 6 3 5 3 5 16+18 20
1306–40A ? ? 1 ? 17 11 6 3 5 3 5 15 17+18
1306–40B 8 pvt 1 3 17 11 6 3 5 3 5 16+18 22
1306–40C 9 pvt 1 3 17 11 6 3 5 3 5 18+19 ?
L. wuxiensis sp. nov. 1321–6A ? pqrst ? 1 10 8 6 4 3 2 4 8+9 13
1321–6B 7 pqrst 1 1 12 8 6 4 3 2 4 8 16
1321–14 9 pqrst 1 1 11+12 8 6 ? ? 2 ? 8 ?
1321–15 7+9 ? 1 1 12 8 6 4 3 2 4 9 15
L. chongqingensis sp. nov. 1311–1A 8+9 pvt 3 3 18 12 7 4 6 2−3 5 19+23 20
1311–1B 8 pvt 3 3 17+18 12 7 4 6 3 5 18+20 ?
1311–1C 9 pvt 3 2+3 18 12 7 4 6 3 5 17+20 18
1311–1D 11 pvt 3 3 19 12 7 4 6 3 5 22+25 20
L. baojiensis nom. nov. 9 pvt 3 3 22−23 12−13 7 7 6 3 5 16−20 21−22
L. trifascia 8−12 pvt 3 3 25−27 16 8 7 6 3 5 16−20 18−22
L. alba ? ? ? 2 ? 9 ? 2 5 3 4 ? ?
L. calolepis* 7 ? 0 3 7 4 ? 0 3 1 0 0 0
L. gigantea** ? ? ? 3 10 9 ? 2 3 3 ? ? ?
L. javana* 9 ? 0 3 9 6 ? 0 3 1 ? ? ?
L. neotropicalis 8−10 pqrst 0 2 16−17 8 6−7 6 6 2 3 ? ?
L. nigropunctata ? ? ? 2 15 20 ? 8 4 2 ? 13 ?
L. sundana 7 ? 0 3 16 20 ? 5 5 3 ? ? ?
L. pallida ? ? ? ? ? ? ? ? ? ? ? ? ?
L. unguserrata ? ? ? ? ? ? ? ? ? ? ? ? ?

* based on Yoshii and Suhardjono (1989); **based on Yoshii (1992).

Lepidosira similis sp. nov.

https://zoobank.org/7685D6DC-F92A-4A00-9CE5-25AD80C65116
Figs 8, 9, 10, 11, 12, 13, Table 3

Type material.

Holotype. CHINA • ♀ on slide; Chongqing Municipality, Wuxi County, the Yintiaoling National Nature Reserve, the Hongqi Protection Station; 1,129.10 m a.s.l.; 31°30'33"N, 109°49'10"E; 21 Jul. 2024; Y.T. Ma leg.; sample number 1306 (CQWX0701).

Paratypes. CHINA • 2 ♀ on slides, same data as holotype.

Diagnosis.

Violet pigment present on Ant. III–IV and post-ocular region; coxae and distal part of femora of mid and hind legs with a little violet pigment; Ant. IV with bilobed apical bulb; head with one mac on Gr. II; Th. II with three medio-medial, one medio-sublateral and 17 posterior mac; Th. III to Abd. IV respectively with 11, 3, 5, 3, and 15–19 central mac; Abd. III with five lateral mac.

Description.

Body (head + trunk) length up to 2.45 mm, holotype 2.24 mm.

Color pattern : ground color pale yellow; eye patches dark blue; violet pigment present on Ant. III–IV and post-ocular region; coxae and distal part of femora of mid and hind legs with a little violet pigment (Fig. 8A, B).

Figure 8. 

Lepidosira similis sp. nov. A, B. Habitus (lateral view). Scale bars: 500 μm.

Scales : ciliate type, present on Ant. I–II (Fig. 9A), basal Ant. III, legs (Fig. 9B), both faces of ventral tube, ventral side of manubrium and dentes (Fig. 9C, D), head, and terga (Figs 9E, 10A).

Figure 9. 

SEM photomicrographs of scales of Lepidosira similis sp. nov. A. Antenna; B. Leg; C. Manubrium; D. Dens; E. Trunk. Scale bars: 20 μm.

Figure 10. 

Lepidosira similis sp. nov. A. Scale on trunk; B. Apex of Ant. IV (dorsal view); C. Distal Ant. III (lateral view); D. Distal Ant. II (dorsal view); E. Dorsal head (right side); F. Prelabral and labral chaetotaxy (dorsal view); G. Maxillary palp and outer lobe (right side); H. Labial palp (right side); I. Labial and post-labial chaetotaxy (right side). Scale bars: 20 μm.

Head : antenna not annulated and 0.62–0.71 (holotype 0.65) times length of body, Ratio of Ant. I–IV as 1.00/1.57–1.67/1.33–1.42/2.62–2.75 (holotype 1.00/1.57/1.41/2.65). Distal part of Ant. IV with many sensory chaetae and normal ciliate chaetae, apical bulb bilobed (Fig. 10B). Ant. III sense organ with two rods, two spiny guard sensilla, smooth blunt sens, and ciliated chaetae (Fig. 10C). Ant. II with 2–3 rods apically (Fig. 10D). Eyes 8+8, G and H smaller than others, interocular chaetae as p, r, t mes. Dorsal chaetotaxy of head with 8–9 antennal (An), four anterior (A0, A2, A3, A5), four median (M1–4), seven sutural (S0–3, S5–7) mac, one post-sutural mac (Ps2) and mes (Ps5), 12 posterior mac plus one (Pa1) in Gr. II (Fig. 10E). Prelabral and labral chaetae as 4/5, 5, 4, prelabral chaetae ciliate and other smooth, a2 and b2 slightly shorter than middle ones, labral papillae conical (Fig. 10F). Basal chaeta on maxillary outer lobe almost as thick as apical one; sublobal plate with four smooth chaetae-like processes (Fig. 10G). Lateral process (l. p.) of labial palp E finger-shaped with tip not reaching apex of papilla E (Fig. 10H). Labial base with M1M2REL1L2, all ciliate (Fig. 10I).

Thorax : tergal ms formula on Th. II–Abd. V as 1, 0/1, 0, 1, 0, 0, sens as 2, 2/1, 2, 2, 2, 3 (Figs 11A, 12A–C). Th. II with 10–12 anterior, three medio-medial (m1, m2, m2i), one medio-sublateral (m4), 17 (rarely 15) posterior mac. Th. III with 11 central and six lateral mac (Fig. 11A). Coxal chaetal formula as 3–5/10–14 (anterior), 3–9 (posterior)/17–23 (Fig. 11B–D). Trochanteral organ with 75–103 smooth chaetae (Fig. 11E). Tenent hair smooth and clavate, 1.21–1.39 length of inner edge of unguis; unguis with a pair of laterobasal outer teeth and four inner teeth, basal pair inner teeth located at 0.42–0.45 distance from base of inner edge of unguis, distal unpaired inner teeth at 0.69–0.73 and 0.87–0.88 distance from base, respectively; unguiculus lanceolate, outer edge slightly serrate (Fig. 11F).

Figure 11. 

Lepidosira similis sp. nov. A. Chaetotaxy of Th. II−III (right side); B–D. Coxal chaetotaxy of fore, middle, and hind legs (outer view); E. Trochanteral organ (posterior side); F. Hind foot complex (lateral view). Scale bars: 20 μm.

Figure 12. 

Lepidosira similis sp. nov. A. Chaetotaxy of Abd. I−III (right side); B. Chaetotaxy of Abd. IV (right side and partially left side); C. Chaetotaxy of Abd. V (right side); D. Anterior face of ventral tube. Scale bars: 20 μm.

Abdomen : range of Abd. IV length as 5.38–7.72 (holotype 7.11) times as dorsal axial length of Abd. III. Abd. I with three (m2–4) mac. Abd. II with five (a2, a3, m3, m3e, m3ep) central, one (m5) lateral mac. Abd. III with three (a2, a3, m3) central, five (am6, pm6, m7a, p6, p7) lateral mac (Fig. 12A). Abd. IV with two normal sens, 15–19 central, 17–22 lateral mac (Fig. 12B). Abd. V with three sens (Fig. 12C). Anterior face of ventral tube with 3+3 large and about one small ciliate chaeta, line connecting proximal (Pr) and external-distal (Ed) mac oblique to median furrow (Fig. 12D); posterior face with two apical smooth chaetae besides about 30 ciliate chaetae of different sizes (Fig. 13A); each lateral flap with 6–8 smooth and about 19 ciliate chaetae (Fig. 13B). Manubrial plate dorsally with 13–18 ciliate mac and 3–4 pseudopores (Fig. 13C); ventrally with 23–33 ciliate chaetae (Fig. 13D). Mucro bidentate; tip of basal spine reaching apex of subapical tooth; distal smooth section of dens almost equal to mucro in length (Fig. 13E).

Figure 13. 

Lepidosira similis sp. nov. A. Apical chaetae of posterior face of ventral tube; B. Lateral flap of ventral tube; C. Manubrial plaque (dorsal view); D. Ventro–apical part of manubrium; E. Distal part of dens and mucro (lateral view). Scale bars: 20 μm.

Etymology.

The Latin adjective word “similis,” meaning similar, refers to its similarity with L. chongqingensis sp. nov.

Habitat.

Found in litter of subtropical forests, mainly composed of leaves of Buxus sinica, Dendrobenthamia japonica var. chinensis, Ilex yunnanensis, Rosa corymbulosa, and Viburnum betulifolium.

Remarks.

This new species is very similar to L. chongqingensis sp. nov. and L. baojiensis nom. nov. in color pattern, and their chaetotaxy of Th. II–III and Abd. III–IV is also similar, but their chaetotaxy of Abd. I and central Abd. II is different. The main differences among the three species in chaetotaxy are listed in Table 3. The new species is also similar to the Indonesian species L. javana (Börner, 1913) in color pattern, but the labial base formula is M1M2REL1L2 in the former and mRel1l2 in the latter (Yoshii and Suhardjono 1989). Their dorsal body chaetotaxy is also quite different (Table 3).

Lepidosira wuxiensis sp. nov.

https://zoobank.org/4112D50C-A946-46DD-886F-0CB0D670DD3D
Figs 14, 15, 16, 17, 18, 19, Table 3

Type material.

Holotype. CHINA • ♀ on slide; Chongqing Municipality, Wuxi County, the Yintiaoling National Nature Reserve, the Guanshan Protection Station, the Stone Pillar; 2,168.92 m a.s.l.; 31°32'15"N, 109°41'54"E; 28 Jul. 2024; Y.T. Ma leg.; sample number 1321 (CQWX0717).

Paratypes. CHINA • 3 ♀ on slides, same data as holotype.

Diagnosis.

Posterior parts of Abd. IV–V violet pigmented; Ant. IV with bilobed apical bulb; head with one mac on Gr. II; Th. II with one medio-medial, two medio-sublateral and 10–12 posterior mac; Th. III to Abd. IV respectively with 8, 4, 3, 2 and 8–9 central mac; Abd. III with four lateral mac.

Description.

Body (head + trunk) length up to 2.01 mm, holotype 2.01 mm.

Color pattern : ground color pale yellow; eye patches dark blue; each segment of Ant. I–III distally violet pigmented, Ant. IV and base of Ant. I with scattered violet pigment; a little violet pigment present on post-ocular region; posterior parts of Abd. IV–V violet pigmented (Fig. 14A). Scattered violet pigment sometimes present on Th. II–Abd. III and legs (Fig. 14B).

Figure 14. 

Lepidosira wuxiensis sp. nov. A, B. Habitus (lateral view). Scale bars: 500 μm.

Scales : ciliate type, present on Ant. I–II (Fig. 15A), head (Fig. 15B), legs (Fig. 15C), both faces of ventral tube, ventral side of manubrium and dentes (Fig. 15D), terga (Figs 15E, 16A).

Figure 15. 

SEM photomicrographs of scales of Lepidosira wuxiensis sp. nov. A. Basal Ant. I; B. Leg; C. Manubrium; D. Dorsal head; E. Trunk. Scale bars: 20 μm.

Figure 16. 

Lepidosira wuxiensis sp. nov. A. Scale on trunk; B. Apex of Ant. IV (dorsal view); C. Distal Ant. III (lateral view); D. Distal Ant. II (dorsal view); E. Dorsal head (dorsal view); F. Prelabral and labral chaetotaxy (dorsal view); G. Maxillary palp and outer lobe (right side); H. Labial palp (right side); I. Labial and post-labial chaetotaxy (right side). Scale bars: 20 μm.

Head : antenna not annulated and 0.56–0.57 (holotype 0.57) times length of body. Ratio of Ant. I–IV as 1.00/1.50–1.73/1.30–1.44/2.22–2.77 (holotype 1.00/1.67/1.40/2.77). Distal part of Ant. IV with many sensory chaetae and normal ciliate chaetae, apical bulb bilobed (Fig. 16B). Ant. III sensory organ with two rods, two spiny guard sensilla, smooth blunt sens, and ciliate chaetae (Fig. 16C). Ant. II with three rods apically (Fig. 16D). Eyes 8+8, G and H smaller than others, interocular chaetae as p, q, r, s, t mes. Dorsal chaetotaxy of head with 7–9 antennal (An), four anterior (A0, A2, A3, A5), four median (M1–4), eight sutural (S0–7) mac, one post-sutural mac (Ps2) mac and mes (Ps5), 12 posterior mac plus one (Pa1) in Gr. II (Fig. 16E). Prelabral and labral chaetae as 4/5, 5, 4, prelabral chaetae ciliate and other smooth, a2 and b2 slightly shorter than middle ones, labral papillae conical (Fig. 16F). Basal chaeta on maxillary outer lobe almost as thick as apical one; sublobal plate with four smooth chaetae-like processes (Fig. 16G). Lateral process (l. p.) of labial palp E finger-like with tip almost or not reaching apex of papilla E (Fig. 16H). Labial base with M1M2REL1L2, all ciliate (Fig. 16I).

Thorax : tergal ms formula on Th. II–Abd. V as 1, 0/1, 0, 1, 0, 0, sens as 2, 2/1, 2, 2, 2, 3 (Figs 17A, 18A–C). Th. II with about nine anterior, one medio-medial (m1), two medio-sublateral (m4, m4i), 10–12 posterior mac. Th. III with eight central and six lateral mac (Fig. 17A). Coxal chaetal formula as 4–6/8–9 (anterior), 4–10 (posterior)/9–10 (Fig. 17B–D). Trochanteral organ with 25–58 smooth chaetae (Fig. 17E). Tenent hair smooth and clavate, 1.12–1.25 length of inner edge of unguis; unguis with a pair of laterobasal outer teeth and four inner teeth, basal paired inner teeth located at 0.46–0.49 distance from base of inner edge of unguis, distal unpaired inner teeth at 0.71–0.74 and 0.85–0.88 distance from base, respectively; unguiculus lanceolate, outer edge slightly serrate (Fig. 17F).

Figure 17. 

Lepidosira wuxiensis sp. nov. A. Chaetotaxy of Th. II−III (dorsal view); B–D. Coxal chaetotaxy of fore, middle, and hind legs (outer view); E. Trochanteral organ (posterior side); F. Hind foot complex (lateral view). Scale bars: 20 μm.

Figure 18. 

Lepidosira wuxiensis sp. nov. A. Chaetotaxy of Abd. I−III (right side); B. Chaetotaxy of Abd. IV (right side and partially left side); C. Chaetotaxy of Abd. V (right side); D. Anterior face of ventral tube. Scale bars: 20 μm.

Abdomen : range of Abd. IV length as 3.29–5.67 (holotype 4.81) times as dorsal axial length of Abd. III. Abd. I with four (m2–4, m4i) mac. Abd. II with three (a2, m3, m3e) central, one (m5) lateral mac. Abd. III with two (a2, m3) central, four (am6, pm6, m7a, p6) lateral mac (Fig. 18A). Abd. IV with two normal sens, 8–9 central, 13–16 lateral mac (Fig. 18B). Abd. V with three sens (Fig. 18C). Anterior face of ventral tube with 3+3 large and 4–6 small ciliate chaetae, line connecting proximal (Pr) and external-distal (Ed) mac oblique to median furrow (Fig. 18D); posterior face with two apical smooth chaetae besides about 35 ciliate chaetae of different sizes (Fig. 19A); each lateral flap with five smooth and nine ciliate chaetae (Fig. 19B). Manubrial plate dorsally with 16–20 ciliate mac and three pseudopores (Fig. 19C); ventrally with 27–31 ciliate chaetae (Fig. 19D). Mucro bidentate; tip of basal spine reaching apex of subapical tooth; distal smooth section of dens slightly longer than mucro in length (Fig. 19E).

Figure 19. 

Lepidosira wuxiensis sp. nov. A. Apical chaetae of posterior face of ventral tube; B. Lateral flap of ventral tube; C. Manubrial plaque (dorsal view); D. Ventro–apical part of manubrium; E. Distal part of dens and mucro (lateral view). Scale bars: 20 μm.

Etymology.

The name is composed of the locality name Wuxi plus the Latin suffix “-ensis,” meaning pertaining or originating, referring to its type locality, Wuxi County.

Habitat.

Found in litter of subtropical forests, mainly composed of leaves of Buxus sinica, Ilex yunnanensis, Pinus armandii, and Rosa corymbulosa.

Remarks.

The new species can be easily distinguished from other species by its color pattern, although it is somewhat similar to the New Zealand species L. dorsalis (Salmon, 1941). However, only the posterior part of Abd. IV is brown pigmented in the former, and almost the whole part of Abd. IV is pigmented in the latter. The main differences between the new species and the other Chinese species of the genus in chaetotaxy are listed in Table 3.

Lepidosira chongqingensis sp. nov.

https://zoobank.org/EC5D94EC-7F94-4499-9A0E-9B88F74154E5
Figs 20, 21, 22, 23, 24, 25, Table 3

Type material.

Holotype. CHINA • ♀ on slide; Chongqing Municipality, Wuxi County, the Yintiaoling National Nature Reserve, the Hongqi Protection Station, the Tianchi Dam; 1,762.16 m a.s.l.; 31°31'39"N, 109°47'18"E; 23 Jul. 2024; Y.T. Ma leg.; sample number 1311 (CQWX0706).

Paratypes. CHINA • 3 ♀ on slides, same data as holotype.

Diagnosis.

Ant. IV, distal Ant. III, and basal Ant. I. with scattered violet pigment; Ant. IV with bilobed apical bulb; head with three mac on Gr. II; Th. II with 3 (2) medio-medial, three medio-sublateral and 17–19 posterior mac; Th. III to Abd. IV respectively with 12, 4, 6, 3 (2), and 18–25 central mac; Abd. III with five lateral mac.

Description.

Body (head + trunk) length up to 1.84 mm, holotype 1.84 mm.

Color pattern : ground color pale yellow; eye patches dark blue; Ant. IV, distal Ant. III, and basal Ant. I. with scattered violet pigment; a little violet pigment present in post-ocular region and sometimes on coxae (Fig. 20A, B).

Figure 20. 

Lepidosira chongqingensis sp. nov. A, B. Habitus (A, lateral view; B, dorsal view). Scale bars: 500 μm.

Scales : ciliate type, present on Ant. I–II, basal Ant. III, legs (Fig. 21A), both faces of ventral tube, ventral side of manubrium and dentes (Fig. 21B, C), head, and terga (Figs 21D, 22A).

Figure 21. 

SEM photomicrographs of scales of Lepidosira chongqingensis sp. nov. A. Leg; B. Manubrium; C. Dens; D. Trunk. Scale bars: 20 μm.

Figure 22. 

Lepidosira chongqingensis sp. nov. A. Scale on trunk; B. Apex of Ant. IV (dorsal view); C. Distal Ant. III (lateral view); D. Distal Ant. II (dorsal view); E. Dorsal head (dorsal view); F. Prelabral and labral chaetotaxy (dorsal view); G. Maxillary palp and outer lobe (right side); H. Labial palp (right side); I. Labial and post-labial chaetotaxy (right side). Scale bars: 20 μm.

Head : antenna not annulated and 0.65–0.75 (holotype 0.65) times length of body. Ratio of Ant. I–IV as 1.00/1.33–2.00/1.24–1.57/2.00–3.10 (holotype 1.00/1.78/1.33/2.67). Distal part of Ant. IV with many sensory chaetae and normal ciliate chaetae, apical bulb bilobed (Fig. 22B). Ant. III sense organ with two rods, two spiny guard sensilla, smooth blunt sens, and ciliated chaetae (Fig. 22C). Ant. II with two rods apically (Fig. 22D). Eyes 8+8, G and H smaller than others, interocular chaetae as p, r, t mes. Dorsal chaetotaxy of head with 8–11 antennal (An), four anterior (A0, A2, A3, A5), four median (M1–4), eight sutural (S0–7) mac, one post-sutural mac (Ps2) and mes (Ps5), 13 posterior mac plus three mac (Pi1, Pa1, Pm1) in Gr. II (Fig. 22E). Prelabral and labral chaetae as 4/5, 5, 4, prelabral chaetae ciliate and other smooth, a2 and b2 slightly shorter than middle ones, labral papillae conical (Fig. 22F). Basal chaeta on maxillary outer lobe almost as thick as apical one; sublobal plate with four smooth chaetae-like processes (Fig. 22G). Lateral process (l. p.) of labial palp E finger-shaped with tip not reaching apex of papilla E (Fig. 22H). Labial base with M1M2REL1L2, all ciliate (Fig. 22I).

Thorax : tergal ms formula on Th. II–Abd. V as 1, 0/1, 0, 1, 0, 0, sens as 2, 2/1, 2, 2, 2, 3 (Figs 23A, 24A–C). Th. II with 11–12 anterior, 3 (2) medio-medial (m1, m2, m2i may be absent), three medio-sublateral (m4, m4i, m4p), 17–19 posterior mac. Th. III with 12 central and seven lateral mac (Fig. 23A). Coxal chaetal formula as 5–9/13–16 (anterior), 8–14 (posterior)/15–23 (Fig. 23B–D). Trochanteral organ with 70–97 smooth chaetae (Fig. 23E). Tenent hair smooth and clavate, 1.21–1.28 length of inner edge of unguis; unguis with a pair of laterobasal outer teeth and four inner teeth, basal paired inner teeth located at 0.42–0.43 distance from base of inner edge of unguis, distal unpaired inner teeth at 0.65–0.69 and 0.86–0.89 distance from base, respectively; unguiculus lanceolate, outer edge slightly serrate (Fig. 23F).

Figure 23. 

Lepidosira chongqingensis sp. nov. A. Chaetotaxy of Th. II−III (right side); B–D. Coxal chaetotaxy of fore, middle, and hind legs (outer view); E. Trochanteral organ (posterior side); F. Hind foot complex (lateral view). Scale bars: 20 μm.

Abdomen : range of Abd. IV length as 4.08–7.00 (holotype 6.67) times as dorsal axial length of Abd. III. Abd. I with four (m2–4, m4i) mac. Abd. II with six (a2, a3, m3, m3e, m3ea, m3ep) central, one (m5) lateral mac. Abd. III with 3 (2) (a2, m3, a3 absent on one side of one specimen) central, five (am6, pm6, m7a, p6, p7) lateral mac (Fig. 24A). Abd. IV with two normal sens, 18–25 central, 18–20 lateral mac (Fig. 24B). Abd. V with three sens (Fig. 24C). Anterior face of ventral tube with 3+3 large and seven small ciliate chaetae, line connecting proximal (Pr) and external-distal (Ed) mac oblique to median furrow (Fig. 24D); posterior face with 2–3 apical smooth chaetae besides about 35 ciliate chaetae of different sizes (Fig. 25A); each lateral flap with five smooth and 20–22 ciliate chaetae (Fig. 25B). Manubrial plate dorsally with 14–21 ciliate mac and four pseudopores (Fig. 25C); ventrally with 33–41 ciliate chaetae (Fig. 25D). Mucro bidentate; tip of basal spine reaching apex of subapical tooth; distal smooth section of dens slightly longer than mucro in length (Fig. 25E).

Figure 24. 

Lepidosira chongqingensis sp. nov. A. Chaetotaxy of Abd. I−III (right side); B. Chaetotaxy of Abd. IV (right side and partially left side); C. Chaetotaxy of Abd. V (right side); D. Anterior face of ventral tube. Scale bars: 20 μm.

Figure 25. 

Lepidosira chongqingensis sp. nov. A. Apical chaetae of posterior face of ventral tube; B. Lateral flap of ventral tube; C. Manubrial plaque (dorsal view); D. Ventro–apical part of manubrium; E. Distal part of dens and mucro (lateral view). Scale bars: 20 μm.

Etymology.

The name is composed of the locality name Chongqing plus the Latin suffix “-ensis,” meaning pertaining or originating, referring to its type locality, Chongqing Municipality.

Habitat.

Found in litter of subtropical forests, mainly composed of leaves of Buxus sinica, Dendrobenthamia japonica var. chinensis, Ilex yunnanensis, Rosa corymbulosa, and Viburnum betulifolium.

Remarks.

The new species is similar to the Indian species L. pallida (Ritter, 1911) and L. unguserrata Salmon, 1970, and the Vietnamese species L. alba (Nguyên, 2005), because all these species lack violet or blue pigment on the trunk. However, L. pallida (Ritter, 1911) is characterized by one inner tooth on the unguis, L. unguserrata Salmon, 1970, by the unilobed apical bulb on Ant. IV, and L. alba (Nguyên, 2005) by two mac on Abd. I. The new species is also similar to the Chinese species L. baojiensis nom. nov. and L. trifascia (Zhou & Ma, 2022), comb. nov. in the chaetotaxy of the dorsal head, Th. II–III, and Abd. II–IV, but Abd. I has four mac in the former and seven in the latter two species. Detailed comparisons among them are listed in Table 3.

Lepidosira baojiensis comb. et, nom. nov.

Willowsia flava Zhou, Pan & Ma, 2021: 74, junior homonym of Lepidosira flava (Salmon, 1938).

Etymology.

The name is composed of the locality name Baoji plus the Latin suffix –“-ensis,” meaning pertaining or originating, referring to its locality, Baoji City.

Diagnosis.

Blue pigment absent on dorsal body; Ant. IV with bilobed apical bulb; head with three mac on Gr. II; Th II with three (m1, m2, m2i) medio-medial, three medio-sublateral (m4, m4i, m4p), and 22–23 posterior mac; Th. III to Abd. IV respectively with 12–13, 7, 6, 3, and 16–20 central mac; Abd. III with five lateral mac.

Remarks.

After checking the type material of this species, we confirm that there are scales on the dentes and no blunt chaetae on the manubrial plates. Hence, we propose its transfer to Lepidosira herein. As a consequence, Willowsia flava Zhou, Pan & Ma, 2021, becomes a junior secondary homonym of Lepidosira flava (Salmon, 1938). Thus, the new replacement name, Lepidosira baojiensis comb. et nom. nov., is being proposed herein.

Lepidosira trifascia (Zhou & Ma, 2022), comb. nov.

Willowsia trifascia Zhou & Ma, 2022: 372.

Diagnosis.

Three longitudinal blue stripes present on dorsal body; Ant. IV with bilobed apical bulb; head with three mac on Gr. II; Th. II with three (m1, m2, m2i) medio-medial, three medio-sublateral (m4, m4i, m4p), and 25–27 posterior mac; Th. III to Abd. IV respectively with 16, 7, 6, 3, and 16–20 central mac; Abd. III with five lateral mac.

Remarks.

After examining the type material of this species, we confirm that scales are present on the dentes and no blunt chaetae are found on the manubrial plate. Hence, we propose its transfer to Lepidosira herein. It can be distinguished from the other known Lepidosira species by its three longitudinal blue stripes on the dorsal body.

Discussion

There are about 9,600 species of Collembola worldwide (Bellinger et al. 1996–2024), but many of these species were diagnosed primarily based on color pattern. Some cryptic species occur in several Collembola genera, such as Tomocerus (Zhang et al. 2014), Coecobrya (Zhang et al. 2018), and Entomobrya (Katz et al. 2015). Molecular methods are very useful for separating species with similar color patterns (Schneider et al. 2011; Porco et al. 2012).

Among the fifteen species of Lepidosira from Asia and South America, six (L. calolepis (Börner, 1913), L. gigantea (Börner, 1909), L. javana (Börner, 1913), L. nigropunctata (Nguyên, 2005), L. sundana Yoshii & Suhardjono, 1989, and L. trifascia (Zhou & Ma, 2022), comb. nov.) have a distinct irregular longitudinal stripe along the lateral side of the trunk from Th. II to Abd. IV. The species L. wuxiensis sp. nov. is characterized by violet pigment on the posterior part of Abd. IV, and L. neotropicalis Nunes & Bellini, 2019, by blue pigment on the dorsal and ventral head anteriorly. There is no great difference in color pattern among the other seven species. However, L. pallida (Ritter, 1911) is characterized by one inner tooth on the unguis, and L. unguserrata Salmon, 1970, by the unilobed apical bulb on Ant. IV. Lepidosira baojiensis comb. et nom. nov. has seven mac on Abd. I; L. alba (Nguyên, 2005) has two mac; L. similis sp. nov. has three mac; and both L. apigmenta sp. nov. and L. chongqingensis sp. nov. have four mac on Abd. I. L. apigmenta sp. nov. has four central mac on Abd. II, and L. chongqingensis sp. nov. has six mac (Table 3).

The genetic distances in the sequences of the COI gene among the seven species were significantly large (Table 2). They corroborated the observed morphological differences, thus providing strong evidence supporting the new species described in the present study.

Acknowledgments

This study was supported by the National Natural Science Foundation of China (32370447) and the Large Instruments Open Foundation of Nantong University (KFJN2475). We also thank Dr. Zhang Zhisheng of Southwest University, China, for assistance with specimen collection.

References

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