Type species: Neanura mandarina Yosii, 1954

Diagnosis

Neanura-like Collembola, body pale yellow to red in life and white in alcohol. With reduced Neanura- type mouthparts. Antennal tubercles isolated from frontal tubercle on head. Organ of Ant. III flanked by 2 guard setae. Tuberculation of post cephalic tergites (Di, De, DL, L), with some of them fused. Tubercles are well individualized and distinct in living specimens. Tergites of Abd. V with 2 + 2 tubercles (De + Di, DL and L tubercle migrated ventrally) or more rarely 1 + 1 + 1 (Di tubercles merged at midline). Di sometimes vestigial on Abd. IV. Most species have 2 or 3 pairs of eyes.

Remarks on Morphology

As mentioned by Deharveng (1983), Deharveng and Weiner (1984) and Smolis and Deharveng (2006), differences between species of this genus are primarily based on mouthparts (such as mandible and maxilla), number of setae on ocular tubercle, unguicular tooth, chaetotaxy, and tubercles Di on head fused or not. Besides that, types of macrosetae maybe important for species separation, but types of short setae on body vary among different individuals of the same species.

Material Examined

HOLOTYPE. Female, eastern China, Shanghai City, Shanghai Botanic Garden, bamboo forest (Bambusoideae spp.), 23-IX-1997, leg. Q.Y. Yue. Paratypes. 4 females and 3 males, same locality as holotype, collected by Q.Y. Yue among 1997–1998 (2 males on 27-VIII-1997, 2 females on 23-IX-1997, 1 male on 27-IX-1997, 2 females on 18-V-1998); 1 female (voucher SH2012009) and 2 males (vouchers SH2012011 and SH2012012), with DNA barcodes sequenced, same locality as holotype, 12-III-2012, leg. Y. Bu and Y. Gao; 2 females, Shanghai, Songjiang County, Zhongjiashan, 6-XI-2002, and 15-VI-2004, leg. Y. M. Yang. Holotype and 10 paratypes are deposited in SIPPE; two paratypes are deposited in UNAM.

Description. Adult. Length (n = 13) 1.7 mm (range: 1.1–2.5 mm). Color red when alive for adults and pink for early juveniles and with barbulate macrosetae (Fig. 1; for color image see Supplementary Fig. 1), white in alcohol. Eye patches dark. Postantennal organ absent. Antenna shorter than head diagonal (ratio as 4:5). Tubercles with heavy granulation (Fig. 8). Macrosetae (ML, Ml and Mc) with heavy barbulation. Ml acuminated, present on lateral tubercles, ML and Mc with blunt tips present almost on all tubercles (Fig. 9). Measurements of setae: long and acuminated barbulate macrosetae (Ml) 85–130 µm, long and thick barbulate macrosetae (ML) 80–113 µm, short barbulate macrosetae (Mc) 25–63 µm, mesosetae (me) thin and smooth 20–50 µm, long smooth setae 100 µm, present on last abdomen segment, microsetae (mi) tiny, 10 µm, and sensorial setae slender, 38–50 µm (Fig. 9).

Ant. I with 7 setae: 2 long, thick and ciliate (Ml), others smooth, and few subcuticular reticulations. Ant. II with 11 setae: 2 long, thick and ciliate setae (Ml), and others smooth. Ant. III and IV dorsally fused, sensorial organ of Ant. III with Sgv and Sgd similar, 2 microsensilla on separate dorso-lateral groove and ventral microsensillum very close to Sgv (Fig. 6). Ant. IV with 8 sensilla and 12 large smooth setae, i seta, trilobed apical vesicle present, subapical organite (or) with global tip, without ventral modified setae (Fig. 6).

Mouth cone short (Fig. 10). Labrum with 0/2, 2 smooth setae of different sizes. Labium with 7 pairs of setae, labial organ, seta A and L absent (Fig. 10), papilla × absent. Mandible with 3 big teeth, apical one with 4–5 denticules (Fig. 4). Maxilla styliform, with 2 lamellae, one of them trifurcated (Fig. 3).

Head with 2 + 2 weakly pigmented eyes. Ocular tubercle with 3 ocular setae, Oca is microseta (mi), Ocm (ML) and Ocp (Mc) are barbulate macrosetae (Fig. 2).

Body setae types as in Fig. 9. Cephalic tubercles and chaetotaxy in Fig. 2 and Table 1. Clypeal tubercle (Cl) with 2 barbulate macrosetae (ML), 2 smooth setae (mi). Antennal tubercles (An) separately, with 4 setae, 2 barbulate macrosetae (ML, Mc) and 2 smooth setae (mi). Frontal tubercle (Fr) with 2 long barbulate macrosetae (ML) and one smooth microseta (mi). Cephalic dorsal dorsointernal (Di) very close each other, coalescent, with 2 long barbulate macrosetae (ML), dorsoexternal (De) with one barbulate macroseta (ML) and 2 smooth microsetae (mi), dorsolateral (DL), lateral (L) and subocular tubercles (So) fused with about 12 setae: 2 long barbulate macrosetae (ML), 1 long and acuminated barbulate macroseta (Ml), 2 short barbulate macrosetae (Mc) and about 7 smooth setae (me, mi).

Number of tubercles from Th. I to Abd. VI on half notum is: 3, 4, 4/4, 4, 4, 4, 2, 1 (Figs. 2 and 11). Thoracic and abdominal chaetotaxy as in Fig. 2 and Table 2. Tubercles setae formula on Th. I-Abd. V as Di: 033/22223; De: 1,3+s,4+s/3+s, 3+s, 3+s, 2+s; DL: 2,3+s+ms, 3+s/2222; L: 133/33364. Sensory setae formula on Th. I-Abd. V as 0, 2+ms, 2/11111. Tubercles De+DL of Abd. V with 1 sensillum and 3 setae (2ML, m). Each tubercle on Abd. VI with 7 setae (2ML, Mc, 4m) (Fig. 11).

Subcoxae with 0, 2, 2 setae, coxae with 3, 7, 8 setae, trochanters with 6, 6, 6 setae; femora with 12–13, 11–12, 10 setae, and tibiotarsi with 19, 19 and 18 setae on legs I, II and III respectively (Table 2). Tibiotarsi without tenent hairs (Fig. 7). Unguis without tooth (Fig. 7).

Chaetotaxy of legs and sterna as shown in Table 2. Ventral tube with 4+4 setae (Fig. 11). Anal tubercle with 14 setae and 3 setulae (Fig. 11). Furcular vestige with 4–5 setae (Fig. 11). Female with 3–4 pairs pregenital setae, 15–27 circumgenital setae and 1+1 eugenital setae (Fig. 12); male with 4 pairs of pregenital setae, about 40 circumgenital setae and 4 + 4 eugenital setae (Fig. 13).

Variation. There is a variation of the type of shortest setae on each body tubercles. In some individuals shortest setae are short barbulate macrosetae (Mc), in others those shortest setae are smooth microsetae (mi). This variation does not depend on age, sex or body size of the specimens. Chaetotaxy of Abd. V of 2 females from Zhongjiashan are 2, 5*, 4 (Others are 3, 4*, 4, Table 2), other variations for the chaetotaxy of ventral side of the abdomen segments are in Table 2.

Etymology

The species is named after the long heavy barbulate macrosetae on the body.

Ecology

In moist litter and leaves under bamboo (Bambusoideae spp.) or lily magnolia, Magnolia liliiflora Desrous (Magnoliales: Magnoliaceae) in Shanghai Botanic Garden.

DNA Barcodes

The standard DNA barcodes (658 base pairs) from Mitochondrial COI gene was amplified and sequenced for 4 individuals from the Shanghai Botanical Garden. There is no divergence between individuals of this population. The sequences were deposited at GenBank with accession number JX027483–JX027486.

Remarks

The new species is similar to V. tubercula Lee and Kim, 1990 from Taibei with big barbulate macrosetae, well developed tubercles and weakly fused Di tubercles on head. But they can be separated by the mouthparts (mandible with 3 smooth teeth in V. tubercula vs. 3 big teeth and the apical one with 4–5 small denticules in V. setaebarbulata sp. nov.; maxilla with 2 simple lamellae in V. tubercula vs. 2 lamellae with one of them trifurcate in V. setaebarbulata sp. nov.), and body chaetotaxy and size (Table 5).

Material Examined

HOLOTYPE. female: eastern China, Shanghai City, Shanghai Botanic Garden, sweet olive plantation (Osmanthus spp.), 28-X-1997, leg. Q. Y. Yue. Paratypes. 2 females and 1 male, same locality as holotype, collected on 28-X-1997, 5-XI-1997, and 10-XI-1997, respectively; 2 females on 23-IX-1997, 1 male on 27-IX-1997, 2 females on 18-V-1998, leg. Q. Y. Yue. 8 females (vouchers SH2012001–SH2012005, SH2012022– SH2012024), garden of Shanghai Institute of Plant Physiology and Ecology, bamboo plantation (Bambusoideae spp.), with DNA barcodes sequenced, 9-III-2012, leg.Y. Bu and Y. Gao; 2 females (voucher SH2012018, SH2012020) and 2 males (voucher SH2012019, SH2012021), with DNA barcodes sequenced, Shanghai Zoo, 24-III-2012, leg.Y. Bu and Y. Gao. Holotype and 18 paratypes are deposited in SIPPE; two paratypes are deposited in UNAM.

Description. Length of adult (n = 21) 2.1 mm (range: 0.9–3.3 mm). Color red in living specimens (Fig. 14; for color image see Supplementary Fig. 14) and with well-distributed pink tubercles, pink for early juveniles, white in alcohol. Eye patches dark (Fig. 14). Postantennal organ absent. Antenna shorter than diagonal of head (ratio as 3:5). Tubercles with granulation distributed around long macrosetae and subcuticular reticulation. All body setae smooth. Blunt macrosetae (ML and Mc) with middle axle, present on most body tubercles, acuminated macrosetae (Ml) present on lateral tubercles. Measurements of setae: long and acuminated macrosetae (Ml) 80–115 µm, long and thick macrosetae (ML) 75–100 µm, short macrosetae (Mc) 30–50 µm, mesosetae (me) 20–50 µm, and sensorial setae slender, 50–75 µm (Fig. 18).

Ant. I with 7 setae, 2 longer than others; Ant. II with 11 setae, 2 longer than others. Ant. III and IV dorsally fused, sensorial organ of Ant. III with Sgv and Sgd similar, 2 microsensilla on separate dorso-lateral groove and ventral microsensillum very close to Sgv (Figs. 16 and 17). Ant. IV with 8 sensilla and 12 large smooth setae, i seta, trilobed apical vesicle and subapical organ (or) present, without ventral modified setae (Fig. 17).

Labrum with 0/2, 2 smooth setae of different sizes. Mouth cone long (Fig. 20), labium with 7 pairs of setae, labial organ, seta A absent, papilla × present (Fig. 20). Mandible with 3 big teeth as V. setaebarbulata sp. nov. (Fig. 21). Maxilla styliform, with 2 simple lamellae (Fig. 21).

2 + 2 weakly pigmented eyes. Ocular tubercle with 3 ocular setae; Oca is a microseta (me), Ocm (ML) and Ocp (Mc) are macrosetae.

Types of setae on body as in Fig. 18. Cephalic tubercles and chaetotaxy in Fig. 15 and Table 3. Head with 14 tubercles. Clypeal tubercle (Cl) with 2 macrosetae (ML), 2 smooth setae (mi). Antennal tubercles (An) isolated, each with 4 setae: one macroseta (ML) and 3 mesosetae (me). Frontal tubercle (Fr) with 2 long macrosetae (ML) and one short macroseta (Mc). Cephalic dorsal dorsointernal tubercle (Di), with one long macroseta (ML), dorsoexternal (De) with one long macroseta (ML) and 2 mesosetae (me), dorsolateral (DL) with 5 setae (ML, 4me), lateral (L) with 3 setae (Ml, 2me). Subocular area not tubercle shaped (So), with about 10 setae: one long macroseta (Ml) and about 9 mesosetae (me).

Number of tubercles from Th. I to Abd. VI on half notum is: 3, 4, 4/4, 4, 4, 4, 3, 1 (Figs. 15, 22). Thoracic and abdominal chaetotaxy as in Fig. 15 and Table 4. Tubercles setae formula on Th. I-Abd. V as Di: 033/22223; De: 1, 3+s, 4+s/3+s, 3+s, 3+s, 2+s; DL: 2, 3+s+ms, 3+s/2223; L: 133/33344. Sensory setae formula on Th. I-Abd. IV as 0, 2+ms, 2/11111. Tubercles De+DL of Abd. V composed by two adjacent tubercles, with one sensillum on small tubercle and 4 setae (2ML, Mc, mi) on large tubercle. Each tubercle on Abd. VI with 7 setae (2ML, 2Ml, 3m) (Fig. 22).

Subcoxae with 0, 2, 2 setae, coxae with 3, 7, 8 setae, trochanters with 6, 6, 6 setae, femora with 12, 11, 10 setae, tibiotarsi with 19, 19 and 18 setae on legs I, II and III, respectively (Table 4), tenent hairs absent (Fig. 23). Unguis without tooth (Fig. 23).

Chaetotaxy of legs and sterna as in Table 4. Ventral tube with 4+4 setae (Fig. 22). Anal tubercle with 14 setae and 3 setulae (Fig. 22). Furcular vestige with 4–6 setae (Fig. 22). Male with 4 pairs pregenital setae, about 36 circumgential setae and 4+4 eugenital setae (Fig. 24). Female with 3–4 pairs pregenital setae, 17–20 circumgenital setae and 1+1 eugenital setae (Fig. 25).

Variation. Lateral tubercles on Abd. V of one female from Shanghai Zoo with 5 setae. Other variations for the chaetotaxy of ventral side of the abdomen segments as shown in Table 4.

Etymology

The species is named V. quartadecima for the 14 tubercles on the head.

Ecology

In moist litter and leaves under bamboo (Bambusoideae spp.) or sweet olive (Osmanthus spp.; Lamiales: Oleaceae) of several gardens in Shanghai.

DNA Barcode

The standard DNA barcodes sequences (658 base pairs) from Mitochondrial COI gene was amplified and sequenced for 12 paratypes (No. N1–5, Nx1–3, N 18–21) of Vitronura quartadecima sp. nov. The gene sequences between SIPPE population and Shanghai Zoo populations are completely identical. All sequences were deposited at GenBank with the accession number JX027487– JX027498.

Remarks

The new species is close to V. giselae (Gisin, 1950) (cosmopolitan species), with smooth axle macrosetae, ventral tube with 4+4 setae and some dorsal chaetotaxy. But they can be separated by the tubercles numbers on the head (12 in V. giselae vs. 14 in V. quartadecima sp. nov.) and mandible (3 smooth teeth in V. giselae vs. 3 big teeth and the apical one with denticules in V. quartadecima sp. nov.) (Table 5).

The different numbers of nucleotides for barcode sequences between Vitronura setaebarbulata sp. nov and Vitronura quartadecima sp. nov. are 122 sites, and the genetic divergence is 18.54%. The different numbers of coded amino acids for this sequence between these 2 new species is 3, and divergence of coded amino acids is 1.37%.

1. Cephalic tubercle Oc with 1 or 2 setae 2

— Cephalic tubercle Oc with 3 setae 12

2. Cephalic tubercle Oc with 1 seta 3

— Cephalic tubercle Oc with 2 setae 4

3. Cephalic tubercle An with 1 seta V. mascula Smolis & Deharveng, 2006; Vietnam

— Cephalic tubercle An with 2 setae V. shaanxiensis Jiang & Yin, 2011; China: Shaanxi

4. Tubercles Di on head fused along midline V. tubercula Lee & Kim, 1990; China: Taiwan

— Tubercles Di on head separate 5

5. Seta O on head present 9

— Seta O on head absent 6

6. Tubercles Di on Abd. V fused along midline V. sinica Yosii, 1976;China: Hongkang

— Tubercles Di on Abd. V separate 7

7. Tubercle De of Th. II with 5 setae V. luzonica Yosii, 1976; Philippines

— Tubercle De of Th. II with 4 setae 8

8. Tubercle An of head with C and E setae V. gressitti Cassagnau & Deharveng, 1981; Papua New Guinea, Brunei

— Tubercle An of head without C and E setae V. kunigamiensis Tanaka & Hasegawa, 2010; Japan

9. Tubercles Di on Th. I with 2 setae V. latior (Rusek, 1967); China:Guangdong

— Tubercles Di on Th. I with 1 seta 10

10. Claw with inner tooth 11

— Claw without inner tooth V. mandarina (Yosii, 1954); Japan, Solomon Islands

11. Tubercle Di on Abd. V with 2 setae V. namhaeiensis Lee, 1974; Korea

— Tubercle Di on Abd. V with 3 setae V. pygmaea (Yosii, 1954); China:Taiwan, Japan, Indonesia

12. Cephalic tubercle De with 2 setae V. Joanna (Coates, 1968); South Africa

— Cephalic tubercle De with 3 setae 13

13. Cephalic tubercles Di fused or weakly fused along midline 14

— Cephalic tubercles Di distinctly separate 16

14. Cephalic seta O on head present 15

— Cephalic seta O on head absent V. singaporiensis (Yosii, 1959); China:taiwan, Southeast Asia

15. Tubercle Di of Abd. I present V. macgillivrayi (Denis, 1933); Costa Rica

— Tubercle Di of Abd. I absent V. setaebarbulata sp. nov.; China: Shanghai

16. Claw with inner tooth 17

— Claw without inner tooth 18

17. Mandible with 5 denticles, tubercle De+DL of Abd. V with 5 setae V. qingchengensis Jiang & Yin, 2012; China: Sichuan

— Mandible with 3 denticles, tubercle De+DLof Abd. V with 4 setae dentata Deharveng & Weiner, 1984; North Korea

18. Head with 14 tubercles V. quartadecima sp. nov.; China: Shanghai

— Head with 12 tubercles 19

19. Tubercle Ve of Abd. IV with 7–9 setae; labrum distally ogival V. acuta Deharveng & Weiner, 1984; North Korea

— Tubercle Ve of Abd. IV with 10 setae; labrum distally non-ogival V. giselae (Gisin, 1950); Switzerland, Kenya, Mexico