Protura from Hainan Island, China: new species, checklist and distribution.

More than 1500 proturan specimens from Hainan Island are systematically studied. An annotated list of all species of Protura from Hainan Island is provided and their geographical distribution is discussed. The genus Paracondeellum is reported from Hainan Island for the first time, and Paracondeellum paradisum sp. nov. is described. The type species Paracondeellum dukouense (Tang & Yin, 1988) is redescribed based on syntype, and the lectotype and paralectotype are designated. The characters of the genus Paracondeellum are redefined, and the two known species are compared in detail. The Protura fauna of Hainan Island is mainly composed of species from the Oriental region, with 91% of the species belonging to the families Berberentulidae and Eosentomidae. Yun Bu, Yan Xiong, Yun-Xia Luan, Wen-Ying Yin.

Introduction

is a group of tiny soil-dwelling arthropods with more than 800 described species (Bu et al. 2012, 2017; Galli et al. 2018). The diagnosis, distribution, and key to 76 known genera and seven families of worldwide were recently given by Galli et al. (2018). So far, there are 214 species belonging to 43 genera recorded in China (Bu et al. 2012, 2017; Qian et al. 2018).

Hainan Island is the second largest island of China and is located off the southernmost point of the mainland (18°10'–20°10'N, 108°37'–111°03'E; Fig. 1). The tropical forest landscape on Hainan Island is one of the hotspots for biodiversity in China, with a high floral diversity and over 6000 species of insects recorded (Huang 2002). In recent years, many rare insects, such as belonging to , have been found on Hainan Island (Yin et al. 2015).

Figure 1.

The sampling localities in Hainan Island. Numbers 1–13 indicate the localities listed in Tables 1, 5.

Table 1.

The sampling localities of in Hainan Island.

Number	Locality	Coordinates	Altitude (m)	Sampling years
1	Ledong County, Jianfengling National Natural Reserve	18°23'– 18°52'N, 108°44'–109°02'E	120–330	1984, 1993, 2003, 2004
2	Wuzhishan City, Wuzhishan National Natural Reserve	18°49'–18°59'N, 109°32'–109°43'E	800–1200	1984, 1985, 2004, 2011
3	Wuzhishan City, Diaoluoshan National Natural Reserve	18°43'–18°58'N, 109°43'–110°03'E	500–1000	1985, 2004
4	Haikou City, Dongzhaigang National Natural Reserve	19°51'–20°01'N, 110°32'–110°37'E	20	2004
5	Haikou City, Crater National Geological Park	19°55'N, 110°12'E	223	2003
6	Tunchang County, Meiling Mountain	19°22'N, 110°04'E	150–230	2003
7	Tunchang County, Limu Mountain	19°17'N, 109°77'E	600–1000	2003
8	Baoting County, Ganshenling Provincial Natural Reserve	18°39'N, 109°66'E	500	2003
9	Changjiang County, Qizi bay	19°21'N, 108°40'E	15	2011
10	Baoting, Qixianling National Forest Park	18°42'N, 109°40'E	150	2017
11	Sanya City, Yalong Bay Tropical Paradise Forest Park	18°15'N, 109°38'E	200	2017
12	Sanya City, Luhuitou Park	18°13'N, 109°29'E	80	2017
13	Sanya City, Sanya bay	18°17'N, 109°22'E	5	2017

Table 5.

The list of proturan species from Hainan Island and their distribution in Hainan Island and three neighboring mainland provinces.

Classification	Species	Hainan	Guangdong	Guangxi	Yunnan
Acerentomata Yin, 1996
Protentomidae Ewing, 1936
Paracondeellum Yin, Xie & Zhang, 1994	P. paradisum sp. n.*	11**
Berberentulidae Yin, 1983
Baculentulus Tuxen,1977	B. tienmushanensis (Yin, 1963)	2, 3, 5, 11			+
Kenyentulus Tuxen, 1981	K. ciliciocalyci Yin, 1987	1, 5, 6, 9, 11, 12			+
	K. dolichadeni Yin, 1987	2
	K. hainanensis Yin, 1987	1, 3	+
	K. henanensis Yin, 1983	2			+
	K. japonicus (Imadate, 1961)	2			+
	K. jianfengensis Yin, 1987	1, 2
	K. jinghongensis Yin, 1983	1			+
	K. minys Yin, 1983	1		+	+
Amphientulus Tuxen, 1981	A. sinensis Xiong, Xie & Yin, 2005	1, 3	+
Sinentomata Yin, 1996
Sinentomidae Yin, 1965
Sinentomon Yin, 1965	S. erythranum Yin, 1965	1, 5, 12, 13	+	+	+
Fujientomidae Yin, 1996
Fujientomon Yin, 1977	F. dicestum Yin, 1977	5
Eosentomata Yin, 1996
Eosentomidae Berlese, 1909
Eosentomon Berlese, 1909	E. actitum Zhang, 1983	1, 3, 8, 11	+
	E. hainanense Yin, 1986*	1, 3, 5, 6, 8	+		+
	E. iban Imadate, 1965	1
	E. jinhongense Yin, 1982	2			+
	E. margarops Yin & Zhang, 1982	1, 2, 3, 5, 10	+
	E. novemchaetum Yin, 1965	11
	E. orientale Yin, 1965	1	+	+
	E. sakura Imadate & Yosii, 1959	1, 2, 4–8, 11	+	+	+
	E. shanum (Zhang, 1984)	1, 4		+
	E. spanum Yin, 1986*	1
	E. tropicum Yin, 1986*	1, 8, 11
	E. xishaense Yin, 1988	1, 5, 9–11
	E. yanshanense Yin & Zhang, 1982	1, 4, 6	+	+	+
	E. zhanjiangense Zhang, 1983	1	+
Anisentomon Yin, 1977	A. hainanense Xiong, Bu & Yin, 2008*	1
	A. quadrisetum Zhang & Yin, 1981	1, 11	+	+
Neanisentomon Zhang & Yin, 1984	N. yuenicum Zhang & Yin, 1984	1, 10	+
Paranisentomon Zhang & Yin, 1984	P. tuxeni (Imadate & Yosii, 1959)	3
Pseudanisentomon Zhang & Yin, 1984	P. paurophthalmum Zhang & Yin, 1984	11		+
	P. molykos Zhang & Yin, 1984	1	+	+
	P. sininotiale Zhang & Yin, 1984	1, 4		+

* Species known only from Hainan Island so far. ** Numbers indicate the localities given in Table 1.

There are several previous publications on the from Hainan Island. The first study reported 14 species of from Hainan (Yin 1986). Then, eight species of the genus () were described (Yin 1987). Later, 24 species of were recorded in Hainan Province with Yin, 1977 and Yin, 1988 included (Yin 1999, 2002). In 2004, the from Jianfengling Mountain were investigated again. In 2005, Xiong, Xie & Yin, 2005 was described and seven new records and three undetermined species were newly added (Xiong 2005; Xiong et al. 2005). One of these undetermined species was subsequently described as Xiong, Bu & Yin, 2008 (Xiong et al. 2008).

In 2011 and 2017, we investigated the soil fauna of Hainan Island on several occasions and collected many proturan specimens. In the present paper, is recorded for the first time and one new species of genus Yin, Xie & Zhang, 1994 is identified and described. We checked the syntypes of the type species of (Tang & Yin, 1988), designated a lectotype and paralectotype, and redescribed it in detail. In addition, based on more than 1500 proturans collected in Hainan Island from 1984 to 2017, a checklist is presented and the distribution of on Hainan Island is summarized.

Materials and methods

Most of specimens were collected between 1984 and 2004, and more recent specimens were collected during the expeditions in 2011 and 2017. All localities sampled so far are listed in Table 1 and shown in Figure 1. All specimens were extracted by means of the Tullgren funnels from soil and humus samples and preserved in 75% ethanol. They were mounted on slides using Hoyer’s solution and dried in an oven at 50 °C.

Observations were made with a phase contrast microscope (Leica DM 2500). Photos were taken by a digital camera (Leica DMC 4500). Line drawings were made using a drawing tube. All specimens are deposited in the collections of Shanghai Natural History Museum (SNHM) and Shanghai Entomological Museum (SEM), Shanghai, China.

Abbreviations used in the text follow the paper by Bu and Yin (2007). Head setae and pores are named according to Rusek et al. (2012) and Shrubovych (2014). The arrangement of the taxa follows the system proposed by Yin (1999).

Results

Taxonomy

Family Ewing, 1936

Yin, Xie & Zhang, 1994

AAF9B8EF-C7D8-5C6B-A2AC-539D9FDF5DF9

Diagnosis.

Habitus short and robust. Pseudoculi circular without lever. Calyx of maxillary glands globular and smooth. Foretarsal sensilla of the exterior side reduced; interior sensilla b’ absent. Abdominal appendages I–II two-segmented each with four setae, III uni-segmented with two setae. Tergites II–VII without or with few anterior setae. Sternites II–III each with three posterior setae. Sternites IV–VII each with nine posterior setae; sternite VIII with four setae in one row. Female squama genitalis short, with pointed acrostyli (Yin 1999; Galli et al. 2018).

Distribution.

South China (Sichuan, Yunnan, Hainan).

Remarks.

Yin, Xie & Zhang, 1994 was originally separated from the genus Tuxen, 1963. They have similar shapes of pseudocellus and maxillary gland, and the presence of setae Pc on sternites IV–V, but they can be easily separated by the chaetotaxy of tergite I (seta P5 absent in but present in ) and sternite VIII (four setae in vs six setae in ). In addition, can be distinguished from the genus Tuxen & Yin, 1982 by the shape of pseudocellus (posterior lever absent in but present in ) and the chaetotaxy of sternites IV–V (setae Pc present in but absent in ).

Bu & Yin sp. nov.

6F18A7F6-9512-58F8-BB0C-AF685065A6D6

http://zoobank.org/A723F8F3-18BF-420F-885E-29EA34F782D7

Figures 2 , 3 , 4 ; Tables 2 , 4

Figure 2.

sp. nov., holotype A head, dorsal view (cp = clypeal pore, fp = frontal pore) B pseudoculus C canal of maxillary gland D labial palpus E maxillary palpus F foretarsus, exterior view G foretarsus, interior view H dorsal thorax, right side I tergite I, right side (al = anterolateral pore, psm = posterosubmedial pore) J tergite VI, right side K comb L female quama genitalis. Arrows indicate pores. Scale bars: 10 μm (B, K); 20 μm (A, C–J, L).

Figure 3.

A–C sp. nov., holotype A sternite VI (spm = sternal posteromedial pore) B tergite VII, right side C sternite VII (spsm = sternal posterosubmedial pore). D–E (Tang & Yin, 1988) holotype D foretarsus, exterior view E foretarsus, interior view. Arrows indicate pores. Scale bars: 20 μm.

Figure 4.

sp. nov., holotype A habitus B canal of maxillary gland C tergites VIII–XII D sternites VIII–XII E tergites V–VII F sternites VI–VII. Scale bars: 100 μm (A); 20 μm (B–F).

Table 2.

Adult chaetotaxy of sp. nov.

Segment		Dorsal		Ventral
		Formula	Setae	Formula	Setae
Th.	I	4	1, 2	(2+2)/6	A1, M
					P1, 2, 3
	II	6/14	A2, 4, M	(4+2)/4	A1, 2, M
			P1, 1a, 2, 3, 4, 5, 5a		P1, 2
	III	6/12	A2, 4, M	(6+2)/4	A1, 2, 3, M
			P1, 1a, 2, 3, 4, 5		P1, 2
Abd.	I	4/12	A1, 5	4/2	A1, 2
			P1, 1a, 2, 2a, 3, 4		P1
	II–III	2/16	A1	4/3	A1, 2
			P1, 1a, 2, 2a, 3, 4, 4a, 5		Pc, 2
	IV–VI	2/16	A1	4/9	A1, 2
			P1, 1a, 2, 2a, 3, 4, 4a, 5		Pc, 1, 1a, 2, 3
	VII	0/16		4/9	A1, 2
			P1, 1a, 2, 3, 3a, 4, 4a, 5		Pc, 1, 1a, 2, 3
	VIII	4/14	A1, 3
			P1, 1a, 2, 2a, 3, 3a, 4	4	1, 2
	IX	12	1, 1a, 2, 2a, 3, 4	4	1, 2
	X	10	1, 2, 2a, 3, 4	4	1, 2
	XI	6		6	1, 2, 3
	XII	9		6

Table 4.

Comparison between sp. nov. and (Tang & Yin, 1988).

	Paracondeellum paradisum sp. nov.	P. dukouense
body length (μm)	570	720–880
pseudoculus (μm)	8	13
foretarsus (μm)	31	46–50
sensilla b and f	short, rod-like	longer, sward-like
sensillum t-3	short and spatulate	longer, sward-like
A-setae on tergite I	4 (A1, A5)	2 (A5)
A-setae on tergites II–VI	2 (A1)	0
P-setae on tergites II–VI	16 (P1a present)	14 (P1a absent)
P-setae on tergite VII	16 (P2a absent)	18 (P2a present)
A-setae on tergite VIII	4 (A1, A3)	6 (A1, A3, A5)
setae on tergite IX	12 (P3a absent)	14 (P3a present)
setae on tergite X	10 (P3a absent)	12 (P3a present)
A-setae on prosternum	2 (A2 absent)	4 (A2 present)

sp. nov. is characterized by two pairs of A-setae on tergite I, one pair of A-setae, and eight pairs of P-setae on tergites II–VI, absence of A-setae and P2a seta on tergite VII, tergites IX and X with 12 and 10 setae, respectively, absence of seta d4 on dorsal side of head, and female squama genitalis short, with conical acrostylus.

Material examined.

Holotype, female (slide no. HN-SY-P2017016) (SNHM), China, Hainan, Sanya City, Yalong Bay Tropical Paradise Forest Park, soil of the tropical rain forest, 200 m elev., , 22-III-2017, Y. Bu collector. Paratypes, 1 female (slide no. HN-SY-P2017071) (SNHM), same data as holotype.

Description.

Holotype: body length 570 μm, yellow-brown, foretarsus darker (Fig. 4A).

Head. Elliptic, length 80 μm, width 50 μm (Fig. 2A). Head setae short, rostrum slightly protruded. Setae d6 and sd6 present, d4 and sd4 absent, d6 and d7 length 6 μm and 7 μm respectively. Pores cp and fp present. Pseudoculus oval, without lever, length 8 μm, width 6.5 μm. PR = 10 (Fig. 2B). Canal of maxillary gland short, with globular calyx and short sausage-like posterior dilation. CF = 10 (Figs 2C, 4B). Labial palpus well developed, with four setae and apical tuft, without basal sensillum (Fig. 2D). Maxillary palpus with two subequal seta-like sensilla (Fig. 2E).

Foretarsus. Length 31 μm, claw length 9 μm, TR = 3.4; empodium length 2 μm, EU = 0.22. Dorsal sensilla t-1 and t-2 slender and long, BS = 0.63; t-3 short and spatulate, not reaching base of claw (Fig. 2G). Exterior side with only sensilla a, b and f present; a spatulate, b and f short (Fig. 2F). Interior sensilla a’ and c’ short sward-like, b’ absent. Relative length of sensilla: t-2 > t-1 > c' > t3 > a > a' > (b = f) (Fig. 2F, G). Length of middle tarsus 15 μm; claw length 10 μm. Length of hind tarsus 17 μm; claw length 12 μm.

Thorax. Thoracic chaetotaxy given in Table 2. Setae 1 and 2 on pronotum subequal in length, 6 μm and 7 μm respectively (Fig. 2H); mesonotum with seven pairs of posterior setae, P5a minute; metanotum with six pairs of posterior setae, P5a absent; setae P1, P1a, P2 on mesonotum 6 μm, 1 μm, 7 μm, respectively; P1a on meso- and metanotum short, pin-shaped (Fig. 2H). Prosternum without seta A2. All setae on thoracic sternites of normal shape. Pores on thorax not observed.

Abdomen. Abdominal chaetotaxy given in Table 2. Tergite I with two pairs of anterior setae (A1, A5) and six pairs of posterior setae, A5 short, sensillum-shaped (Fig. 2I). Tergites II–VI with one pair of anterior (A1) and eight pairs of posterior setae, P2a present and P3a absent (Figs 2J, 3A, 4E, F). Tergite VII without anterior setae and with eight posterior setae, P2a absent and P3a present (Figs 3B, C, 4E, F). Accessory setae P1a on tergites I–V short pin-shaped (4 μm), on tergites VI–VII normal (5 μm). Accessory setae P2a and P4a always pin-shaped, 2 μm in length. P3a on tergite VII of normal shape and 5 μm long (Fig. 4E, F). Tergite VIII with two pairs of anterior setae (A1, A3) (Fig. 4C). Posterior central seta Pc present on sternites IV–VII, sensillum shaped, 4–5 μm long (Figs 3A, C, 4F). P1a on sternites IV–VI short, pin-shaped (Fig. 3A), on sternite VII setiform (Fig. 3C).

Tergites I–VII with pores psm and al (Fig. 2I, J), VIII with pores psm only, IX–XI without pores, XII single median pore. Sternites I–VI each with single posteromedial pore spm (Figs 3A, 4F), VII with three posteromedial pores (Figs 3C, 4F), VIII with single posteromedial pore (Fig. 4D), IX–XI without pores, XII with one pair of anterolateral sal pores.

Abdominal appendages typical of the genus. Subapical setae and apical setae on appendage III 11 μm and 5 μm long respectively. Striate band on abdominal segment VIII reduced to a single serrate line (Fig. 4D). Comb on abdomen VIII rectangular, with 10 teeth, 10 μm wide (Fig. 2K). Female squama genitalis short, with conical acrostylus (Fig. 2L).

Etymology.

Latin “paradisum”, after “Paradise Forest Park” where type specimens were collected.

China (Hainan)

The genus Yin, Xie & Zhang, 1994 is endemic to China and was previously known by a single species, , from Sichuan and Yunnan provinces. sp. nov. differs from in the shape of foretarsal sensilla, pseudoculus, and female squama genitalis, and in the body chaetotaxy. A comparison of the morphology of these two species is given in Table 4.

(Tang & Yin, 1988)

F8ED0826-3C27-5520-AC8B-6EA3D4CA092B

Figures 3 , 5 ; Tables 3 , 4

Figure 5.

(Tang & Yin, 1988), holotype A habitus B canal of maxillary gland C head, dorsal view D prosternum E mesosternum F tergite VI G tergites VII–VIII H sternite VI I sternite VII J sternite V K tergites VIII–XII L sternites VIII–XII. Arrows indicate pores. Scale bars: 20 μm.

Table 3.

Adult chaetotaxy of (Tang & Yin, 1988).

Segment		Dorsal		Ventral
		Formula	Setae	Formula	Setae
Th.	I	4	1, 2	(4+2)/6	A1, 2, M
					P1, 2, 3
	II	6/14	A2, 4, M	(4+2)/4	A1, 2, M
			P1, 1a, 2, 3, 4, 5, 5a		P1, 2
	III	6/12	A2, 4, M	(6+2)/4	A1, 2, 3, M
			P1, 1a, 2, 3, 4, 5		P1, 2
Abd.	I	2/12	A5	4/2	A1, 2
			P1, 1a, 2, 2a, 3, 4		P1
	II–III	0/14		4/3	A1, 2
			P1, 2, 2a, 3, 4, 4a, 5		Pc, 2
	IV–VI	0/14		4/9	A1, 2
			P1, 2, 2a, 3, 4, 4a, 5		Pc, 1, 1a, 2, 3
	VII	0/18		4/9	A1, 2
			P1, 1a, 2, 2a, 3, 3a, 4, 4a, 5		Pc, 1, 1a, 2, 3
	VIII	6/14	A1, 3, 5
			P1, 1a, 2, 2a, 3, 3a, 4	4	1, 2
	IX	14	1, 1a, 2, 2a, 3, 3a, 4	4	1, 2
	X	12	1, 2, 2a, 3, 3a, 4	4	1, 2
	XI	6		6	1, 2, 3
	XII	9		6

(Tang & Yin, 1988) is characterized by the one pair of A-setae on tergite I, absence of A-setae and P1a seta on tergites II–VI, absence of A-setae and nine pairs of P-setae (P2a present) on tergite VII, tergites IX and X with 14 and 12 setae respectively, absence of seta d4 on head, and female squama genitalis with pointed acrostylus.

Lectotype, female (slide no. 1), paralectotype, female (slide no. 2) (SEM), China, Sichuan, Dukou City (currently, Panzhihua City), Jinjiang County, soil under grass, 1155 m elev., , 26-IX-1985, B.W. Tang and G.T. Jin collectors. We designated as the lectotype the female on slide no. 1 and the other female on slide no. 2 as the paralectotype.

Redescription.

Body length of holotype 880 μm and paratype 720 μm; yellow-brown, with foretarsus darker (Fig. 5A).

Head. Elliptic, length 93–100 μm, width 70 μm. Dorsal setae longer than subdorsal and lateral ones, rostrum slightly protruded (Fig. 5C). Setae d6 and sd6 present, sd6 sensillum-shaped; d4 and sd4 absent; d6 and d7 11 μm and 6 μm long, respectively (Fig. 5C). Pores cp and fp present. Pseudoculus round, without lever, length 13 μm, width 11 μm. PR = 7.2–7.7 (Fig. 5C). Canal of maxillary gland short, with globular calyx and sausage-like posterior dilation. CF = 13.3–14.3 (Fig. 5B). Labial palpus well developed, with four setae and apical tuft, without basal sensillum. Maxillary palpus with two subequal sensilla.

Foretarsus. Length 46–50 μm, claw length 15–17 μm, TR=2.9–3.1; empodium length 4–5 μm, EU=0.24–0.33. Dorsal sensilla t-1 and t-2 slender and long, BS=0.66; t-3 short sward-like, nearly reaching base of claw (Fig. 3D). Exterior slide with only sensilla a, b and f present; a spatulate, b and f short sward-like (Fig. 3D). Interior sensilla a’ and c’ short sward-like, b’ absent (Fig. 3E). Relative length of sensilla: t-2 > t-1 > c' > a > t3 > a' > (b = f) (Fig. 3D, E). Length of middle tarsus 20 μm; claw length 12 μm. Length of hind tarsus 23 μm; claw length 15 μm.

Thorax. Thoracic chaetotaxy given in Table 3. Setae 1 and 2 on pronotum subequal in length, 10 μm long; mesonotum with seven pairs of posterior setae, P5a minute; metanotum with six pairs of posterior setae, P5a absent; setae P1, P1a, P2 on mesonotum 10 μm, 1.5 μm, 14 μm respectively; P1a on meso- and metanotum short, pin-shaped. Prosternum with anterior seta A2 (Fig. 5D), meso- and metasternum each with four posterior setae (Fig. 5E), metasternum with six anterior setae. All setae on sterna normal. Pores on thorax not detectable.

Abdomen. Abdominal chaetotaxy given in Table 3. Tergite I with one pair of anterior setae (A5) and six pairs of posterior setae, A5 short, sensillum-shaped. Tergites II–VI without anterior setae and seven pairs of posterior setae, P2a present, P1a and P3a absent (Fig. 5F). Tergite VII without anterior setae and with nine pairs of posterior setae, both P2a and P3a present (Fig. 5G). Accessory setae P2a and P4a on tergites II–VII short, sensillum-shaped, 4 μm in length, P1a and P3a on tergites VII normal, 9–10 μm in length (Fig. 5F, G). Tergite VIII with three pairs of anterior setae (A1, A3, A5) and seven pairs of posterior setae, P3a short (5 μm) (Fig. 5G, K, L). Posterior central seta Pc present on sternites IV–VII slender, 8–9 μm long (Fig. 5H–J). P1a on sternites IV–VI short pin-shaped, 2 μm long (Fig. 5H, J), on sternite VII as normal seta, 9 μm long (Fig. 5I). Sternites IX and X with short P3a seta (Fig. 5L), which had been omitted in original description.

Tergites I–VII with pores psm and al, VIII with pores psm only, IX–XI without pores, XII with single median pore. Pores on sternites I–VI not observed due to the opacity of the old specimens (Fig. 5H, J); three posteromedial pores observed on sternite VII (Fig. 5I), VIII with posteromedial pore (Fig. 5L), IX–XI without pores, XII with one pair of sal pore.

Abdominal appendages typical of the genus. Subapical setae and apical setae on appendage III 12–13 μm and 6–8 μm long, respectively. Striate band on abdominal segment VIII reduced to a single serrate line (Fig. 5G, K, L). Comb on abdomen VIII rectangular, with 10 teeth, 12–13 μm wide (Fig. 5K). Female squama genitalis short, with pointed acrostylus.

Named for Dukou City (now Panzhihua City, Sichuan Province) where type specimens were collected.

China (Sichuan, Yunnan).

was originally described based on two syntypes (Tang and Yin 1988). In the original description (Tang and Yin 1988) and in the monograph of Yin (1999), most important characters such as foretarsal sensilla, pseudoculus, maxillary gland, as well as body chaetotaxy were briefly described and illustrated. After careful study of type specimens under a modern phase contrast microscope with higher resolution, we find that sensillum c’ is present on the foretarsus and that some of the setae on the body were previously ignored due to the lower resolution of the microscope used. We correct here these mistakes in the original description and supplement the description of head chaetotaxy, the porotaxy, and the shapes of setae on the body. Table 4 compares with the new species.

List of species from Hainan Island

Family Ewing, 1936

sp. nov.

14AEAD6E-C784-589E-984F-F347E2926AE1

Description.

The description is given above.

Family Yin, 1983

(Yin, 1963)

BC4E1EF5-F991-5ECB-BC70-7894EC060EDC

Material examined.

1 male, 1 mj, locality 3, 19-I-1985, coll. G. T. Jin & Z. Y. Liu. 4 females, 1 mj, locality 5, 26-II-2003; 1 female, locality 1, 14-I-2004; 10 females, 6 mj, locality 2, 27-I-2004, coll. Y. Xiong. 1 mj, locality 11, 22-III-2017, coll. Y. Bu.

Distribution.

Widely distributed in China (Hainan, Zhejiang, Shanghai, Jiangxi, Anhui, Hubei, Sichuan, Chongqing, Guizhou, Yunnan, Ningxia, Gansu, Shaanxi, Henan, Hebei, Liaoning, Neimenggu).

Yin, 1987

DAEA8EC0-B0B8-599B-8A1F-430EEA90D8D0

5 females, locality 1, 27-XI-1984, coll. G. T. Jin & Z. Y. Liu. 1 female, VI-1993; 1 female, IV-1994, locality 1, coll. C. H. Liao. 9 females, 2 males, locality 5, 26-II-2003; 1 female, locality 6, 2-III-2003; 1 female, locality 5, 15-VI-2003; 6 females, 1 male, locality 1, 6-X-2003; 9 females, 2 males, 13 mj, locality 1, 14-I-2004; 7 females, 3 males, locality 1, 15-I-2004; 1 female, locality 1, 14-IV-2004; 2 females, 5 males, 1 mj, locality 1, 15-VII-2004, coll. Y. Xiong. 2 females, 1 male, 2 mj, locality 9, 20-III-2011, coll. Y. Bu & C. W. Huang. 14 females, 10 male, 1 mj, locality 11, 22-III-2017; 3 males, locality 12, 17-X-2017, coll. Y. Bu.

Widely distributed in China (Hainan, Zhejiang, Hunan, Sichuan, Chongqing, Guizhou, Yunnan, Shaanxi).

Yin, 1987

1949F23F-73CC-5029-9781-479910DE8677

3 females, locality 2, 14-XI-1984, coll. G. T. Jin & Z. Y. Liu.

China (Hainan, Zhejiang, Guangxi, Guizhou, Hubei, Sichuan, Jiangxi).

Yin, 1987

069666C6-53E9-5ED5-ABA9-CF36A40539A9

4 females, 1 male, locality 1, 30-XI-1984; 2 females, 2 males, locality 2, 14-XI-1984; 1 male, 2 mj, locality 3, 19-I-1985, coll. G. T. Jin & Z. Y. Liu.

China (Hainan, Guangdong).

Yin, 1983

C409945E-EC68-5719-9684-C7D26DCF0CB2

2 female, 1 male, locality 2, 19-XI-1984, coll. G. T. Jin & Z. Y. Liu.

Widely distributed in China (Hainan, Zhejiang, Jiangxi, Henan, Hubei, Guizhou, Yunnan, Ningxia).

(Imadate, 1961)

15C9072A-885C-5DDE-9165-525C29C3FC2A

2 females, locality 2, 14-XI-1984; 5 females, 5 males, locality 1, 25-XI-1984, coll. G. T. Jin & Z. Y. Liu.

Widely distributed in China (Hainan, Zhejiang, Jiangsu, Shanghai, Jiangxi, Anhui, Hunan, Sichuan, Guizhou, Yunnan, Shaanxi); Japan.

Yin, 1987

2283ED13-750D-521E-9E58-6C9A6E8C237A

4 females, locality 1, 1-XII-1984, coll. G. T. Jin & Z. Y. Liu. 2 females, 1 male, 1 mj, locality 1, 6-X-2003; 7 females, 5 males, 2 mj, locality 1, 15-I-2004; 1 mj, locality 3, 27-I-2004; 3 females, 1 males, 1 mj, locality 1, 14-IV-2004; 7 females, 9 males, locality 1, 15-VII-2004, coll. Y. Xiong.

China (Hainan, Guizhou).

Yin, 1983

4C410C8D-934C-58F0-B6BC-C74724136BA0

3 females, locality 1, 25-XI-1984, coll. G. T. Jin & Z. Y. Liu.

China (Hainan, Yunnan, Guizhou).

Yin, 1983

FFAB7526-BB9F-58A9-828D-930BB2FA5E98

2 females, 2 males, locality 1, 19-XI-1984, coll. G. T. Jin & Z. Y. Liu.

China (Hainan, Yunnan, Guangxi, Jiangxi)

Xiong, Xie & Yin, 2005

674EC540-2878-571A-9BF8-006304F5895B

1 female, locality 1, 17-XII-2002; 1 female, locality 2; 8 females, 4 males, 1 mj, locality 3, 27-I-2004, coll. Y. Xiong.

China (Hainan, Guangdong).

Family Yin, 1965

Yin, 1965

C38F3FC0-C2C3-5D67-BA34-02520599B07F

1 female, 1male, 1 mj, locality 5, 26-II-2003; 1 mj, locality 1, 6-X-2003; 7 females, locality 1, 15-VII-2004, coll. Y. Xiong. 1 female, 1 LI, locality 13, 16-X-2017; 1 LI, locality 12, 17-X-2017, coll. Y. Bu.

Widely distributed in South China (Hainan, Shanghai, Jiangsu, Zhejiang, Anhui, Fujian, Guangxi, Guangdong, Hunan, Guizhou, Yunnan).

Yin, 1996

Yin, 1977

882EFFBD-5783-527D-ABCB-25EEC2A623E2

1 female, locality 5, 26-II-2003, coll. Y. Xiong.

China (Hainan, Shanghai, Jiangsu, Zhejiang, Anhui, Ningxia).

Berlese, 1909

Zhang, 1983

8A312847-ED66-526E-8B96-A2742CA7D017

8 females, 6 males, 2 mj, locality 1, XII-1984; 24 females, 24 males, 27 mj, locality 3, 23-I-1985, coll. G. T. Jin & Z. Y. Liu. 2 females, 3 males, locality 8, 22-II-2003, coll. Y. Xiong. 2 females, 3 mj, locality 11, 22-III-2017, coll. Y. Bu.

China (Hainan, Guangdong, Sichuan).

Yin, 1986

BF9B93BA-485A-5E00-B4F6-27E47728AEE5

40 females, 33 males, 2 mj, locality 1, 25-XI-1984, coll. G. T. Jin & Z. Y. Liu. 2 females, VI-1993, locality 1, coll. C. H. Liao. 1 female, 5 males, locality 8, 22-II-2003; 4 females, 1 male, locality 6, 2-III-2003; 1 female, 3 males, 4 mj, locality 5, 15-VI-2003; 19 female, 15 males, 8 mj, locality 1, 7-X-2003; 33 females, 31 males, 15 mj, locality 1, 14-I-2004; 1 male, locality 3, 960 m elev., 27-I-2004; 8 females, 8 males, 1 mj, locality 1, 15-VII-2004, coll. Y. Xiong.

China (Hainan, Yunnan).

Imadate, 1965

CDE43EFB-E528-58D3-A558-6A8D8C89DCAE

2 females, 2 males, locality 1, 27-XI-1984, coll. G. T. Jin & Z. Y. Liu.

China (Hainan); Malaysia, Brunei.

Yin, 1982

E06C531A-0D94-5E98-BFA5-7851EBFB78FD

2 females, 1 male, locality 2, 14-XI-1984, coll. G. T. Jin & Z. Y. Liu.

China (Hainan, Yunnan).

Yin & Zhang, 1982

E6A367D6-24B0-5D55-A550-8D1BE9B2EDB6

1 female, 2 males, 1 mj, locality 5, 26-II-2003; 1 female, 1 male, 2 mj, locality 5, 15-VI-2003; 1 female, 2 males, locality 1, 6-X-2003; 6 females, 3 males, 1 mj, locality 1, 15-I-2004; 2 females, 1 male, 1 mj, locality 2, 820 m elev., 27-I-2004; 3 females, 2 males, 2 mj, locality 3, 1000 m elev., 27-I-2004; 2 females, 1 male, locality 1, 14-IV-2004; 2 females, locality 1, 15-VII-2004, coll. Y. Xiong. 1 female, locality 10, 23-III-2017, coll. Y. Bu.

China (Hainan, Guangdong, Sichuan).

Yin, 1965

8AE595C2-7549-5F4F-A1E8-A80214AFDCD8

1 female, locality 11, 22-III-2017, coll. Y. Bu.

China (Hainan, Shanghai, Jiangsu, Anhui, Jiangxi, Liaoning, Shaanxi)

Yin, 1965

1694DE9C-605E-5ACD-B2F5-C520818FF429

1 female, locality 1, 25-XI-1984, coll. G. T. Jin & Z. Y. Liu.

Widely distributed in China (Hainan, Shanghai, Jiangsu, Zhejiang, Anhui, Jiangxi, Hubei, Hunan, Guangxi, Guangdong, Sichuan, Chongqing, Guizhou, Ningxia, Shaanxi, Liaoning).

Imadate & Yosii, 1959

A9CD6AEA-1164-5FF4-BB40-7D12DF2E5C98

5 females, 3 males, locality 1, 25-XI-1984, coll. G. T. Jin & Z. Y. Liu. 13 females, 7 males, 9 mj, locality 8, 22-II-2003; 1 female, 1 mj, locality 5, 20-I-2003; 8 females, 13 males, 7 mj, locality 5, 26-II-2003; 1 female, 1 male, locality 6, 2-III-2003; 2 females, 5 males, 2 mj, locality 5, 15-VI-2003; 2 females, 1 mj, locality 1, 6-X-2003; 3 females, 1 male, 2 mj, locality 7, 13-VII-2003; 5 females, 2 males, 2 mj, locality 1, 15-I-2004; 46 females, 45 males, 29 mj, locality 4, 23-I-2004; 12 females, 20 males, 10 mj, locality 2, 820 m elev., 27-I-2004; 5 females, 5 males, 2 mj, locality 1, 500 m elev., 27-I-2004, coll. Y. Xiong. 1 male, locality 2, 20-III-2011, coll. Y. Bu & C. W. Huang. 8 females, 7 males, 3 mj, locality 11, 22-III-2017, coll. Y. Bu.

Widely distributed in China (Hainan, Shanghai, Jiangsu, Zhejiang, Anhui, Jiangxi, Hubei, Hunan, Guangxi, Guangdong, Yunnan, Sichuan, Fujian, Guizhou, Taiwan, Hong Kong, Shaanxi).

(Zhang, 1984)

1D0B51EB-5F67-5C3B-A074-D3F2CB089F16

2 females, 1 male, locality 4, 23-I-2004; 1 female, 2 males, locality 1, 14-IV-2004; 5 females, 1 mj, locality 1, 15-VII-2004, coll. Y. Xiong.

China (Hainan, Guangxi, Hunan, Jiangxi).

Yin, 1986

508839A1-4D50-5849-9BE9-1818C48F2BDC

1 female, 1 male, locality 1, 27-XI-1984, coll. G. T. Jin & Z. Y. Liu.

China (Hainan).

Yin, 1986

8DE2605A-639E-515E-882F-4E8A98B669C5

5 females, locality 1, 25-XI-1984, coll. G. T. Jin & Z. Y. Liu. 1 female, VII-1993, locality 1, coll. C. H. Liao. 1 female, 3 males, 3 mj, locality 8, 22-II-2003; 4 females, 3 males, 4 mj, locality 1, 6-X-2003; 2 mj, locality 1, 15-I-2004; 3 females, 1 male, locality 1, 15-VII-2004, coll. Y. Xiong. 2 females, locality 11, 22-III-2017, coll. Y. Bu.

China (Hainan).

Yin, 1988

8D963A9F-C583-5F19-9E45-7569CB835357

2 females, locality 1, 27-XI-1984, coll. G. T. Jin & Z. Y. Liu. 1 mj, VI-1993, locality 1, coll. C. H. Liao. 1 female, locality 5, 20-I-2003; 3 mj, locality 1, 14-IV-2004; 4 females, 2 males, locality 1, 15-VII-2004, coll. Y. Xiong. 1 female, locality 9, 20-III-2011, coll. Y. Bu & C. W. Huang. 2 females, 1 male, 1 mj, locality 11, 22-III-2017; 1 male, locality 10, 23-III-2017, coll. Y. Bu.

China (Hainan, Xisha Islands, Yongxing Island).

Yin & Zhang, 1982

AFFD70A7-FBA1-5D48-8985-47CBCD7068C6

2 females, locality 1, 25-XI-1984, coll. G. T. Jin & Z. Y. Liu. 2 females, 4 males, 7 mj, locality 6, 2-III-2003; 9 females, 3 males, 10 mj, locality 4, 23-I-2004; 2 females, 2 males, locality 1, 14-IV-2004; 2 females, 1 mj, locality 1, 15-VII-2004, coll. Y. Xiong.

China (Hainan, Guangxi, Guangdong, Fujian, Jiangxi, Hunan, Hubei, Yunnan).

Zhang, 1983

C3271BDF-683F-53FD-A4FF-13D10DECCC7B

2 females, locality 1, 19-I-1985, coll. G. T. Jin & Z. Y. Liu. 1 females, 2 males, locality 1, 14-IV-2004, coll. Y. Xiong.

China (Hainan, Guangdong).

Xiong, Bu & Yin, 2008

157D9716-E249-5B6E-B3D3-F4891A8D3716

1 female, 1 male, locality 1, 6-X-2003, coll. Y. Xiong.

China (Hainan).

Zhang & Yin, 1981

F3177C7D-A610-5DC4-AC41-62669FE5DE1D

1 female, 1 male, locality 1, 7-X-2003, 14 females; 7 males, locality 1, 14-I-2004, coll. Y. Xiong. 1 male, 1 mj, locality 11, 22-III-2017, coll. Y. Bu.

China (Hainan, Guangxi, Guangdong).

Zhang & Yin, 1984

74612738-A6E7-56F1-9E5E-B5C92570CAB1

1 female, locality 1, 14-IV-2004, coll. Y. Xiong. 1 female, 1 male, locality 10, 23-III-2017, coll. Y. Bu.

China (Hainan, Guangdong).

(Imadate & Yosii, 1959)

06343631-9405-5DCF-B99B-AE12D3DDF16B

6 females, 1 mj, locality 3, 1000 m elev., 27-I-2004, coll. Y. Xiong.

China (Hainan, Hubei, Hunan, Jiangxi, Anhui, Guizhou, Shaanxi).

Zhang & Yin, 1984

CF1EE964-6F9F-5B23-BBCA-62A1A810F6CA

1 female, 1 mj, locality 11, 22-III-2017, coll. Y. Bu.

China (Hainan, Guangxi).

Zhang & Yin, 1984

8037AC5C-72A5-578C-89B5-5D666247C7AB

5 females, 2 males, 1 mj, locality 1, 6-X-2003; 1 female, locality 1, 14-IV-2004, coll. Y. Xiong.

China (Hainan, Guangdong, Guangxi, Yunnan).

Zhang & Yin, 1984

3022B716-9730-512B-8167-F277C02FE66C

2 females, locality 1, 27-XI-1984, coll. G. T. Jin & Z. Y. Liu. 2 females, 4 males, locality 4, 23-I-2004; 1 male, locality 3, 600 m elev., 27-I-2004, coll. Y. Xiong.

China (Hainan, Guangxi, Hunan).

Discussion.

The 34 species of recorded from Hainan Island belong to 11 genera and five families (, , , and ) (Table 5). Most species (91%) belong to (21 species) and (10 species), while the other three families are represented by one species each. Both and are widely distributed in China and have high species richness. In contrast, , , and each has fewer species occurring in China. Undoubtedly, proturans found in Hainan Island are mainly related to the fauna of Oriental Region and are distinctly different from those from Russian Far East and Siberia (Bu et al. 2014; Shrubovych 2014), which are dominated by the family .

By comparing the species distribution, we found that the fauna of Hainan Island is closely related to those of neighboring mainland regions (Yin 1999; Szeptycki 2007), and there are 13, 10, and 11 species shared with Guangdong, Guangxi, and Yunnan provinces, respectively (Fig. 1; Table 5), which is consistent with the geological history of Hainan Island (Wang 1991; Zhang and Fang 2012). Until the Quaternary period (2.5 million years ago), Hainan Island was still connected with Leizhou Peninsula of Guangdong Province. In the Middle Pleistocene, fault depression led to the separation of Hainan Island from the mainland. With sea level fluctuations, Hainan Island was connected to or separated from the mainland for several times. Since the end of the Quaternary period, due to the drastically rise of sea level, Hainan Island has been separated from the mainland without interruption.

Among the mainland regions neighboring Hainan Island, the fauna of Yunnan Province has been systematically studied (Zhang et al. 1996; Yin et al. 2000), and nearly 80 species were reported from that province, with the and having fairly high diversity (Zhang et al. 1996; Yin et al. 2000). In this study, we found the diversity of fauna from Hainan Island is very similar to that from Yunnan Province. The only difference is the presence of family in Yunnan, which is absent in Hainan Island.

Sampling localities in Hainan Island are still sparse, and additional collection of proturans should be made in the future, so as to reveal the true diversity and provide a better understanding of the biogeography of on the Hainan Island.