Zootaxa, A new genus and species of ...

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ZOOTAXA

A new genus and species of Chresmodidae (Insecta: Gryllones) from Upper Jurassic-Lower Cretaceous of Yixian Formation, Inner Mongolia, China XIN-WEN ZHANG 1, DONG REN 2*, HONG PANG 1* & CHUNG-KUN SHIH 2 1 State Key Laboratory of Biocontrol and Institute of Entomology, Sun Yat-Sen University, Guangzhou 510275, China 2 Key Lab of Insect Evolution & Environment Change, Capital Normal University, Beijing 100037, China *Corresponding author: E-mail: [email protected] or [email protected]

Abstract A new genus with a new species of Chresmodidae (Insecta: Gryllones), Sinochresmoda magnicornia gen. et sp. nov., including specimens of male, female and nymph, is described and illustrated. These fossils were collected from the Upper Jurassic-Lower Cretaceous of Yixian Formation of Liutiaogou Village, Inner Mongolia (Nei Mongol Autonomous Region), China. Both male and female possess wings, and have sexual dimorphism in antenna. Antennae of male are highly specialized and horn-shaped, incurvated as horn with the scape strongly expanded, and the first segment of the flagella incurvated as a pair of brackets, while antennae of female are normally filiform. Wings of the new species are short, not exceeding the length of the abdomen. Fringe hairs along the wing margins, a unique feature for Chresmodidae, are dense, wavy and bundled together. Nymphs and adults probably lived in the same environment. In addition, one specimen demonstrates the extraordinary structure of tarsus with a high number of “tarsomeres”, which has been reported in other species of this family. Key words: fossil insect, Chresmodidae, sexual dimorphism, new taxa, Yixian Formation, Late Jurassic-Early Cretaceous, China

Introduction Chresmodidae, an extinct insect family, is quite rare. It currently contains 1 genus with 4 species: Chresmoda obscura Germar, 1839; Chresmoda orientalis Esaki, 1949; Chresmoda aquatica Martínez-Delclòs, 1989; and Chresmoda libanica Nel, Azar, Martínez-Delclòs & Makhoul 2004. Besides, Saurophthiroides mongolicus is also suspected to belong to this family (Rasnitsyn, 2002; Nel et al., 2004). The family has been found in Eurasian region and Brazil, ranging from the Upper Jurassic to the Upper Cretaceous (Grimaldi & Engel, 2005; Nel et al., 2005). The first species, C. obscura, was erected by Germar (1839). Debates on its taxonomic status have never ceased ever since its finding, ranging from Hemiptera, Mantodea, Phasmatodea and Polyneoptera (Zherikhin, 1978; Carpenter, 1992; Martínez-Delclòs, 1989; Rasnitsyn and Quicke, 2002; Grimaldi & Engel, 2005). It is known that nymphs and males are apterous, but females have four wings with possible flight capability (Nel et al., 2005). Chresmodids are supposed to be aquatic insects, probably carnivorous (Baudoin, 1980; Nel et al., 2004; Nel et al., 2005). In this family, tarsi possess an extraordinary structure, so called ultraarticulated tarsi, which are probably associated with their capability of skating on water-surface (Nel et al., 2004; Nel et al., 2005). This family is still an enigma to date. Recently, we collected several well-preserved chresmodids from Yixian Formation of Liutiaogou Village, Inner Mongolia (Nei Mongol Autonomous Region), China. Based on their different and unique morphological characters, we erect a new genus and species, Sinochresmoda magni-

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Accepted by H. Song: 28 Dec. 2007; published: 13 Feb. 2008

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cornia gen. et sp. nov. by describing both male and female of this species, which exhibit remarkable sexual dimorphism in antennae. Age of the Yixian Formation is in contention. Three opinions were proposed: the Late Jurassic (Ren et al. 1997, Zheng et al. 2003), the transition from the Late Jurassic to the Early Cretaceous (Cao. 1999, Jin. 1999, Chen et al. 2004, Wang et al. 2004, Wang et al. 2005), and the Early Cretaceous (Swisher et al. 1999, Li et al. 2001, Pang et al. 2002, Zhou et al. 2003). It is impossible to draw a conclusion according our new specimens. We consider the age to be the Late Jurassic-Early Cretaceous here.

Material and methods The specimens were examined with a Leica MZ 7.5 dissecting microscope and illustrated with the aid of a drawing tube attachment. Partially magnified images of the specimens were taken under a Nikon SMZ1000. Line drawings were prepared with Adobe Illustrator CS 11.0.0 graphics software. All the type specimens studied in this paper are housed in the Key Lab of Insect Evolution & Environmental Changes, the College of Life Sciences, Capital Normal University, Beijing, China (CNU; Ren Dong, Curator). We follow the systematic arrangement of the Chremodidae proposed by Rasnitsyn (2002), who considered Chesmodidae as a separate family in Gryllones of uncertain position.

Systematic paleontology Gryllones of uncertain position (after Rasnitsyn, 2002) Family Chresmodidae Handlirsch, 1908 Genus Sinochresmoda gen. nov. Zhang, Ren, and Pang Type species. Sinochresmoda magnicornia sp. nov. Zhang, Ren, and Pang Etymology. The generic name is a combination of the Greek prefix sino- (China) and Chresmoda (the type genus of this family). Gender: feminine. Diagnosis. Head small. Both male and female have wings. Antennae exhibit sexual dimorphism. Scape expanded slightly in female, while expanded significantly in male. The first segment of the flagellum is elongated in both genders, but incurvated as a pair of brackets in male, while normally straight in female. Legs are long and narrow with similar shape; mid leg femora are the longest. Coxae on the same side of thorax are far apart from each other. Fringe hairs along the wing margins, a unique feature for Chresmodidae, are dense, wavy and bundled together. Comparison. Sinochresmoda n. gen. is closely related to Chresmoda Germar, 1839, but can be distinguished by the following characters: expansion of the first segment of antennae and male with wings. Fringe hairs along the wing margins are absent in Chresmoda. Saurophthiroides Ponomarenko, 1986, is represented only by one specimen (probably nymph) without wings. The new genus differs from it by larger body size and expansion of the first segment of antennae.

Sinochresmoda magnicornia sp. nov. Zhang, Ren, and Pang Etymology. A combination of the Latin prefixes Magni- (large) and cornus (meaning corn, referring the antenna’s horn-shape of the male). Material. Holotype: No. CNU-CH-NN2007004-1, -2, female, a well-preserved part and counterpart, with a visible ovipositor.

A NEW FOSSIL CHRESMODID FROM CHINA

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Paratype: Female specimens: No. CNU-CH-NN2007005-1, -2; Male specimens: No. CNU-CHNN2007001-1, -2; No. CNU-CH-NN2007002-1, -2 and No. CNU-CH-NN2007003; Nymph specimen: No. CNU-CH-NN2007006. No. CNU-CH-NN2007001, CNU-CH-NN2007002, CNU-CH-NN2007004, and CNU-CH-NN2007005 are part and counterpart specimens. No. CNU-CH-NN2007003 is smaller than other specimens with its abdomen missing. It is probably a male, because of its “male” antennae, strong and robust, with the first segment strongly expanded. Furthermore, No. CNU-CH-NN2007006 is a well-preserved nymph, with wing buds and an under-developed ovipositor. Type locality and horizon. Yixian Formation, the Late Jurassic-Early Cretaceous, Liutiaogou Village, Ningcheng City, Inner Mongolia, China.

FIGURE 1. S. magnicornia n. g. n. sp., holotype, female: A, photograph of CNU-CH-NN2007004-1; B, photograph of CNU-CH-NN2007004-2; C, line drawing of CNU-CH-NN2007004-1; D, line drawing of CNU-CH-NN2007004-2; scale 10mm.

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Diagnosis. Fore tibia is approximately 40% of the femur length. Wings, short and not exceeding the length of abdomen, representing approximately 60% of the body length. Other diagnostic characters are the same as those of generic diagnosis. Description Female Mainly based on holotype specimen, CNU-CH-2007004-1, -2. Size medium, wings not exceeding the length of abdomen, antenna filiform (Fig. 1). Head: Short, length 3.2 mm. Antennae are filiform, 9.6 mm long for the holotype, and 9.1 mm for CNUCH-NN2007005 (Fig. 2). The scapes, slightly expanded, are larger than the other segments. Pedicel is short and less than half as long as scape. The first segment of the flagellum is elongated; the rest seem to become narrower toward apex. Compound eyes are inconspicuous. Most of mouthparts were covered, without an exact form. Two strong palpi with at least three segments are visible, probably maxillary palpus. The last segments of palpus are smaller than basal segments.

FIGURE 2. S. magnicornia n. g. n. sp., holotype, female, CNU-CH-NN2007004-1: A, line drawing of antenna; B, photograph of antennae; scale 3mm.

Thorax: Longer than width, with visible wing sheaths. The prothorax is somewhat trapezoidal, of cleared sides and vertices, the sides with smooth margin. Mesothorax and metathorax are tightly connected. The sterna of the thorax are poorly preserved. FIGURE 3. S. magnicornia n. g. n. sp., A, line drawing of fringe hairs along the wing margins, paratype, male, CNUCH-NN2007001-1; photograph of fringe hairs along the wing margins: B, paratype, male, CNU-CH-NN2007001-1; C, paratype, male, CNU-CH-NN2007002-2; D, holotype, female, CNU-CH-NN2007004-1; all scale 3mm



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Legs: Extended, long and narrow, covered with dense short hairs. The mid legs are the longest (70.1 mm), followed by the hind (59.2 mm) and the fore (56.4 mm). The pairs of coxae are widely separated, all of similar shape, those on the same side of thorax being apart from each other. The fore femur, straight and elongate, bears dense hairs and a series of setae. Fore tibia, short, 42% of the femur length in the holotype, 41% in CNU-CH-NN2007005. Fore tarsi are partially missing, the tarsomeres are indistinct. A structure, probably the joints of tarsomeres were found in the basal part of left fore leg. Mid leg has the longest femur, representing 37% of the total leg length. Mid tibia is 5.2 mm longer than fore tibia, all of similar shape. Tarsi are indistinct. The hind leg is slightly longer than the fore leg. Hind tibia represents 52% of the femur length. The entire tarsus is 24.6 mm long, tarsomeres are poorly preserved. Wings: Short, not exceeding the length of abdomen, 15.5 mm long, representing 61 of the body length. Due to folding and overlapping, most venation is invisible. Fringe hairs along the wing margins are dense, wavy and bundled together (Fig. 3 D). Abdomen: Eight visible sternites, 13.4 mm long approximately. Two cerci, 4.0 mm long. The ovipositor is long, sword-like, valvulae V1 bearing a row of inner indentations at the tail end. The ovipositor valvulae V2 are slightly longer than the V1. The valvulae V3 are at base of V1, somewhat difficult to discern (Fig. 4).

FIGURE 4. S. magnicornia n. g. n. sp., holotype, female, CNU-CH-NN2007004-2: A, photograph of ovipositor; B, line drawing of ovipositor; scale 2mm.

The dimensions of these and all other specimens are summarized in Tables 1 and 2. Male Mainly based on the paratype specimens, CNU-CH-2007001-1, -2. Similar to female, but remarkably different for its antennae, of which the scape is strongly expanded, the first flagellomere is incurvated. Antennae are longer than the female, but with less number of segments (Fig. 5). A NEW FOSSIL CHRESMODID FROM CHINA


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FIGURE 5. S. magnicornia n. g. n. sp., paratype, male: A, photograph of CNU-CH-NN2007001-1; B, line drawing of CNU-CH-NN2007001-1; C, photograph of CNU-CH-NN2007001-2; D, line drawing of CNU-CH-NN2007001-2; scale 10mm.

Head: Short, 4.2 mm long, with width of 4.5 mm. Antennae, 14 segments, 13.3 mm long, slightly longer than the head and thorax combined. Antennae bent towards each other and extended forward with the hornshape (Fig. 6). The scape is strongly expanded, 2.7 mm long, 1.5 mm wide. Second segment is short and small, only 1/5 of the scape. The third one is the longest segment, incurvated as a pair of brackets, 5.0 mm long approximately. Following 11 segments are all similar, becoming narrower toward the apex. The whole antennae are covered with dense short hairs. Compound eyes are somehow prominent, located at both sides of the head. Most of mouthparts were covered dorsally, without an exact form. Two palpi are visible with at least three segments, probably maxillary palpus. The last segments of palpus are smaller than basal segments. Thorax: Slightly longer than wide, with visible wing sheaths. Prothorax is wider than head, somewhat trapezoidal, the sides with smooth margin. Pronotum is somehow prominent. Metanotum was covered by the wings. Mesonotum and coxal cavity are somewhat distinct. Mesothorax and metathorax are of nearly the same size, closely connected. Legs: Similar to the female, extended, long and narrow, covered with dense and short hairs. The small hairs are arranged in rows, dense in edge, sparse in the middle. Tibiae and tarsi preserved poorly, some parts are obscure. The pairs of coxae are widely separated, those on the same side of thorax being apart from each


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other. Femora of mid legs are the longest, 25.5 mm. Fore leg tibia, short, representing 40% of the femur length for CNU-CH-NN2007001; 42% for CNU-CH-NN2007002; 41% for CNU-CH-NN2007003. Tibiae of hind legs are slightly longer than the fore tibiae. Tarsi are partially preserved, segmentation of tarsomeres invisible. A structure has been found in the tarsus of left mid leg of CNU-CH-NN2007003, which probably also exists in the right. It seems to be the so-called ultra-articulated tarsi (Nel et al. 2004, Nel et al., 2005), which are quite different from normal segments or tarsomeres. The middle of the tarsus seems to be divided into many parts (>25). This division is not found in the apex, probably due to poor preservation. Wings: Short, arises at the mesothorax, folded on the back, represented about 60% of the body length. Veins are indistinct, few longitudinal veins can be observed faintly, but they seem to remain parallel to the margin. Cross-veins are invisible. Fringe hairs along the wing margins are dense, wavy and bundled together (Fig. 3 A-C). Wings are 17.2 mm long, representing 63% of the body length (CNU-CH-NN2007001). Abdomen: With eight visible segments, possibly 10 or 11 segments in total. Two cerci, 2.8 mm long, 0.4 mm wide at the base. Genitalia are poorly preserved. Epiproct covered the genitalia and larger than the sternum. Details of genitalia are blurry (Fig. 7).

FIGURE 6. S. magnicornia n. g. n. sp., paratype, male CNU-CH-NN2007001-2: A, photograph of head and antennae, B, line drawing of antenna; scale 3mm.

FIGURE 7. S. magnicornia n. g. n. sp., paratype, male CNU-CH-NN2007001-1: A, photograph of genitalia; B, line drawing of genitalia; scale 2mm.



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Nymph Based on paratype specimen of CNU-CH-NN2007006. Female, appears similar to adult, with wing buds and under-developed ovipositor (Fig. 8). Head short, pressed dorsally open. Antennae are poorly preserved, the right one was broken, only a few basal segments left. Compound eyes, prominent, located at both sides of the head. Prothorax is trapezoidal, similar to the adult. Two pairs of wing buds are well-preserved, arises at the mesothorax. Abdomen was ruptured into two parts, ten visible segments are well-preserved. Ovipositor and cerci are prominent. Three valvulae of the ovipositor are discernable, with almost the same shape. Compared to the adult, the ovipositor of the nymph seems to be under-developed.

FIGURE 8. S. magnicornia n. g. n. sp., paratype, female, nymph, CNU-CH-NN2007006: A, photograph of general habitus, scale 10mm; B, line drawing of antenna, scale 3mm; C, photograph of antenna, scale 3mm; D, photograph of wing buds, scale 3mm; E, photograph of ovipositor, scale 3mm.

Remarks Sinochresmoda magnicornia n. gen., n. sp. can be distinguished from C. obscura by characters that the latter has all femora of equal length, with the wings exceeding the length of abdomen. C. aquatica most closely resembles S. magnicornia, but no male C. aquatica has been reported, and thus, the comparisons of antennae of male are impossible. Female antennae are similar in both species, but more segments of flagellum in the new species (>= 17 segments in S. magnicornia, 16 segments in C. aquatica); the first flagellomeres are elongated in S. magnicornia. Wing margins of S. magnicornia possess a series of fringe hairs. Saurophthiroides mongolicus is based on a specimen without wings, probably a nymph (Ponomarenko, 1986; Rasnitsyn, 2002; Nel et al., 2004). S. magnicornia can be distinguished by characters that the new species has a larger body size, with wings, and longer antennae.


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C. libanica are nymphs, different from nymph of S. magnicornia in their smaller size, shorter fore tibia and absence of visible wing buds. C. orientalis were also found in China, based on a unique adult specimen. At the first sight, C. orientalis seems to be similar to female of S. magnicornia. But, we do not have accurate knowledge about its structure. As it was not mentioned by Esaki (1949), it is impossible for us to confirm the gender of the holotype just by looking at the photograph of holotype specimen (Esaki, 1949: pi 1). Differed from S. magnicornia, C. orientalis seems to have a shorter fore tibia, compared to the fore femora (35%~36% in C. orientalis after Esaki, 1949: pi 1; and 40%~42% in S. magnicornia). Furthermore, C. orientalis possesses longer wings, whereas S. magnicornia has shorter wings; however, comparisons of details of wings are impossible. We could not eliminate the possibility that C. orientalis is included in Sinochresmoda. But due to lack of firm published data and detailed examination of the original holotype, it is more reasonable to keep its status unchanged for now. Discussion Due to similar characteristics among these new specimens and C. aquatica, C. libanica, C. obscura and Saurophthiroides mongolicus, we assigned them to Chresmodidae. Similar to most species of this family, wings of S. magnicornia were folded on the abdomen, causing a great loss of venation characters. Although the mouthparts are preserved incompletely, we think they should be non-sucking, same as Nel et al. (2004). Males of this genus certainly have unique antennae, never found in other species of Chresmodidae, but somewhat similar to the antennae of female Palophus phillips, which is an extant species, assigned to Phasmatodea (Brock, 1999). The morphological characters of nymph indicate that both nymphs and adults might have lived in similar environmental conditions. The systematic position of this family remains questionable. We could not draw a more definite conclusion based on our new materials. Ultra-articulated tarsi A structure was found in the left tarsus of mid leg of CNU-CH-NN2007003. The tarsus seems to possess numerous segments (Fig. 9B). Nel et al. (2004, 2005) once described this structure in Chresmoda and named it ultra-articulated tarsi, and considered it widely exists in Chresmodidae. However, this structure is somehow different from normal segments or tarsomeres. Another structure, probably joints of tarsomeres were found in the left fore leg of CNU-CH-NN2007004-1 (Fig. 9A). The length between the two joints is 3.8 mm. It appears similar to the normal segment, and should be the two basal segments of tarsus, the second is slender than the first one. Nel et al. (2004) also found this phenomena in Chresmoda libanica and described the two basal tarsomeres are of “normal size”. However, the basal segments are much longer in fore leg tarsus of CNU-CHNN2007004, while the mid leg tarsus of CNU-CH-NN2007003 has a greater full-length. Unfortunately, we could not find good evidence about these two structures among our other specimens, and could not draw a conclusion. Only future discovery of well-preserved specimens would shed more light on it. Sexual dimorphism in Sinochresmoda magnicornia Except for the difference in antennae and genitalia between males and females, they possess many similar characters. For example, they have same shape, short head, long legs, short wings (16.1 mm average length) (Table 1), and ratio of wing length/body length at about 60%. All adult specimens have fringe hairs along the wing margins are dense, wavy and bundled together, which are clearly visible (Fig. 3). Furthermore, thorax notum and propleuron are of similar shape, coxae on the same side of thorax are far apart from each other in every specimen. Legs all long and narrow, fore tibia are approximately 40% of the femur (Table 2). In addition, all specimens were collected from the Liutiaogou Village in the same deposit from nearby locations. We believe that they should have lived in a same area at about the same geological time.



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FIGURE 9. S. magnicornia n. g. n. sp., A, holotype, female CNU-CH-NN2007004-1, photograph left tarsus of fore leg; B, paratype, male CNU-CH-NN2007003, photograph left tarsus of mid leg; scale 1mm.

TABLE 1. Length of body parts (mm). NO.

Head

Thorax

Wings

Total body

Wing length/ body length

CNU-CH-NN2007001(m#)

4.2

9.0

17.2

27.2

63%


3.3

9.3

16.5

27.7

60%


2.7

9.2

15.2

CNU-CH-NN2007004(f#)

3.2

9.6

15.5

25.6

61%

CNU-CH-NN2007005(f#)

3.6

7.8

27.1

CNU-CH-NN2007006(f#, Nymph)

3.1

7.3

24.8


ZHANG ET AL.

TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. TABLE 2. Length of the legs (mm). NO.

Fore legs femur

tibia

tarsus

Mid legs tibia/

femur

tibia

Hind legs tarsus

femur

tibia

tarsus

22.7

11.2

24.3?

femur CNU-CH-

23.0

9.3

40%

25.5

23.1

9.7

42%

25.4

15.7?

21.9

9.0

41%

23.2

13.3

25.2

21.5

11.5

23.9

10.0

42%

25.9

15.2

29.0

22.6

11.8

24.6

20.8

8.6

41%

22.9

14.3

18.9

7.6

40%

19.6

10.5

8.8

18.7

NN2007001(m#) CNU-CH-

23.3

NN2007002(m#) CNU-CHNN2007003(m#) CNU-CH-

22.5

NN2007004(f#) CNU-CH-

20.6

NN2007005(f#) CNU-CH-NN2007006

17.7

21.1

17.4

(f#, Nymph)

FIGURE 10. S. magnicornia n. g. n. sp., photograph of paratype specimen, female: A, CNU-CH-NN2007005-1; B, CNU-CH-NN2007005-2; scale 10mm.



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FIGURE 11. S. magnicornia n. g. n. sp., photograph of paratype specimen: A, male, CNU-CH-NN2007002-1; B, male, CNU-CH-NN2007002-2; C, CNU-CH-NN2007003; scale 10mm.


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Based on evidences given above, we believe it reasonable to classify these specimens as a single species. The differences of antennae morphology between males and females is proposed as a sexual dimorphism. Primarily, antenna is a sensing organ, which can be used to smell, touch, and/or perceive sounds (Chapman, 1998). Antennae of male S. magnicornia are horn-shaped, while those of female are normal filiform. We suppose that the male antenna may have some other special function besides sensing. More studies are needed to know its function accurately. All specimens of S. magnicornia were found with short wings, except for nymph and the CUN-CH2007003 of which abdomen is missing. We could not eliminate the possibility of that these specimens are brachypterous individuals of this species, or they are at the last ontogenetic stage before adults. However, due to four specimens showing the similar wing length (representing approximately 60% of the body length), we are inclined to consider that these specimens are adults with short wings here, unless future discovery of specimens of S. magnicornia to prove otherwise.

Acknowledgements We sincerely thank Dr. Martínez-Delclós X. (Dept. Estratigrafia, Paleontologia i Ceociències Marines, Univ. Barcelona, Barcelona), and two anonymous reviewers for revision of the manuscript. We sincerely thank Mr. Fang Liang for his excellent fossil specimens. We are very grateful to E. Baum (Entomology Group, Institut für Spezielle Zoologie und Evolutionsbiologie, Friedrich-Schiller-Universität Jena, Jena, Germany) for his referral of important reference papers. We also wish to thank Prof. Liang Geqiu and Dr. Jia Fenglong (State Key Laboratory of Biocontrol and Institute of Entomology, Sun Yat-sen University, Guangzhou, China), for their valuable consultations and practical help. The research has been supported by grants from the National Nature Science Foundation of China (No. 30430100), PHR Project of Beijing Municipal Commission of Education and the Beijing National Science Foundation (Grant No. 5082002), and the Funding Project for Academic Human Resources Under the Jurisdiction of Beijing Municipality.

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