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Research Article
Three new species of the Chinese planthopper genus Tetricodes Fennah (Hemiptera, Fulgoroidea, Issidae, Parahiraciini)
expand article infoZhi-Min Chang, Lin Yang, Jian-Kun Long, Xiang-Sheng Chen
‡ Guizhou University, Guiyang, China
Open Access

Abstract

The diagnostic characters of the Chinese planthopper genus Tetricodes Fennah, 1956 are redefined. Three new species of this genus, T. ansatus Chang & Chen, sp. n. (China: Guangxi), T. parvispinus Chang & Chen, sp. n. (China: Guizhou) and T. similis Chang & Chen, sp. n. (China: Guizhou) are described and illustrated. The female genitalia of the genus is described and illustrated for the first time. A checklist and key to the species of Tetricodes are given. The synonymy of Tetricodes songae Zhang & Chen, 2009 with Tetricodes polyphemus Fennah, 1956, syn. n., is proposed.

Keywords

Fulgoromorpha, issid, planthopper, morphology, taxonomy, distribution

Introduction

The planthopper tribe Parahiraciini Cheng & Yang, 1991 (Hemiptera: Issidae) is a relatively small tribe distributed mainly in the Oriental region but also occurring in the Palaearctic region (Bourgoin 2016). In recent years, some genera and species were added or moved into this tribe (Gnezdilov and Wilson 2005; Meng et al. 2015; Wang et al. 2015, 2016; Gnezdilov and Hayashi 2016; Gnezdilov and Bourgoin 2016); to date, the tribe contains 19 genera and 68 species. Parahiraciini can be distinguished from other issid tribes by: body elongate ovate; forewings with claval suture; hindwings well developed, bilobed, and with a network of veins.

The genus Tetricodes was established by Fennah (1956) for T. polyphemus Fennah, 1956 as the type species from China (Hubei), with generic characters mainly based on a female specimen. Zhang and Chen (2009) transferred Tetricodes from the tribe Thioniini Melichar, 1906 to Parahiraciini and were first to recorded the male genitalia of the type species and one other species, T. songae from Guizhou, China. Subsequently, Chen et al. (2014) described another new species T. anlongensis. According to the frons with or without a light median line, Gnezdilov (2015) proposed that T. polyphemus in Zhang and Chen (2009) was a new species, T. fennahi Gnezdilov, 2015; up to date, the genus includes four species.

The main aim of this paper is to describe three new species of Tetricodes (Hemiptera: Issidae) from China (Guangxi, Guizhou), and propose a new species synonym. Also, type specimens are discussed, generic characteristics are redefined and a checklist and key to species of Tetricodes are provided.

Materials and methods

The morphological terminology of the head and body follows Chan and Yang (1994), and the terminology of male and female genitalia follows Bourgoin (1993) and Gnezdilov (2002, 2003). Dry specimens were used for descriptions and illustrations. External morphology was observed under a stereoscopic microscope. All measurements are in millimeters (mm). The body measurements are from the apex of vertex to the tip of the forewing. The genital segments of the examined specimens were macerated in 10% KOH, washed in water and transferred to glycerine. Illustrations of the specimens were made with a Leica MZ 12.5 stereomicroscope. Photographs were taken with a KEYENCE VHX-1000C.

The type specimens of new and other species and other examined specimens are all deposited in the Institute of Entomology, Guizhou University, Guiyang, China (IEGU).

Taxonomy

Tetricodes Fennah, 1956

Tetricodes Fennah, 1956: 513; Zhang and Chen 2009: 17; Chen et al. 2014: 116.

Type species

Tetricodes polyphemus Fennah, 1956.

Diagnostic characters

Body slender beetle-like, oval. Width of head (Figs 1, 3, 5) including eyes narrower than pronotum. Vertex (Figs 7, 22, 31) with length in middle shorter than width at base, disc of vertex depressed, without median carina, with anterior margin obtuse-angled convex, posterior margin slightly concave. Frons (Figs 9, 24, 33) flat, median carina without or only present at base, big black hemispheroidal protuberance with light median line running from the upper margin, pale transverse band under the black protuberance. Clypeus triangular, broadly rounded, with median carina distinct or obscure. Rostrum surpassing mesotrochanters. Pronotum (Figs 1, 3, 5) short, with median carina obscure, lateral carinae not reaching to the posterior margin. Mesonotum (Figs 1, 3, 5) triangular, with median carina obscure, lateral carinae distinct. Hind tibiae each with 2–4 spines. Forewings (Figs 10, 26, 35) elongate, with length 2.3–2.6 times longer than maximum width, apical margin acutely rounded, ScP and Rp convergent near base, MP three or four branched, CuA simple, not forked; CuP present, Pcu and A1 uniting in clavus. Hindwings (Figs 11, 26, 35) well developed, as long as forewings, apical margin deeply incised into two big lobes, veins network-liked, anal lobe reduced.

Male genitalia

Anal tube (Figs 13, 28, 37) short, irregularly oval or pentagonal in dorsal view. Anal style (paraproct and epiproct) (Figs 13, 28, 37) not surpassing anal tube. Pygofer (Figs 12, 27, 36) narrow, irregularly subquadrate. Genital styles (Figs 12, 27, 36) relatively short, without triangular prominence near dorsal margin before capitulum. Phallobase (Figs 14, 29, 38) with stout rod-like processes near base, lateral lobe splitting into two branches, with ventral lobe various. Aedeagus (Figs 14, 29, 38) with pair of foliaceous processes in lateral view.

Female genitalia

Anal tube (Fig. 16) very long, truncated apically, anal style short, located at the base of anal tube. Hind margin of gonocoxa VIII with endogonocoxal lobe unobvious (Fig. 18), endogonocoxal process gradually narrowing (Fig. 18). Anterior connective lamina of gonapophyses VIII with 7 lateral teeth bearing 7 keels in lateral group and 3 apical teeth (Fig. 18). Gonoplacs (Fig. 21) without keels.

Checklist of species of Tetricodes Fennah, 1956

Tetricodes anlongensis Chen et al., 2014; China (Guizhou)

Tetricodes ansatus Chang & Chen, sp. n.; China (Guangxi)

Tetricodes fennahi Gnezdilov, 2015; China (Guizhou)

Tetricodes parvispinus Chang & Chen, sp. n.; China (Guizhou)

Tetricodes polyphemus Fennah, 1956; China (Guizhou, Hubei, Hunan, Yunnan)

Tetricodes similis Chang & Chen, sp. n.; China (Guizhou)

Key to species of Tetricodes (males)

1 Anal tube concaved in apical margin 2
– Anal tube convexed in apical margin 4
2 Aedeagus with pair of small sheet processes at apical 1/3 in lateral view 3
– Aedeagus with pair of big sheet processes at apical 1/3 in lateral view (Fig. 38e) T. similis sp. n.
3 Anal tube with apical margin “V”-shaped concavely (see Chen et al., 2014: 2–63, Fig. H) T. anlongensis
– Anal tube with apical margin “U”-shaped concavely (see Chen et al., 2014: 2–62, Fig. H) T. polyphemus
4 Phallobase with two pairs of long processes (Fig. 14b, c) T. ansatus sp. n.
– Phallobase with pair of long processes 5
5 Anal tube irregularly pentagonal in dorsal view, the apical margin truncated (Fig. 28) T. parvispinus sp. n.
– Anal tube irregularly round in dorsal view, the apical margin arched (see Chen et al., 2014: 2–61, Fig. H) T. fennahi

Tetricodes anlongensis Chen, Zhang & Chang, 2014

Tetricodes anlongensis Chen et al., 2014: 120.

Material examined

♂ (holotype), China: Guizhou, Anlong, Xianheping (24°58'N, 105°36'E), 28 Aug. 2012, S.-Y. Xu; 1♂1♀ (paratypes), same data as holotype.

Distribution

China (Guizhou).

Tetricodes ansatus Chang & Chen, sp. n.

Figs 1–2, 7–15, 16–21

Type material

Holotype: ♂, China: Guangxi, Nonggang (22°28'N, 106°57'E), 8 May 2012, H. Li; paratypes: 1♂, Guangxi, Nonggang, 8 May, 2012, Z.-H. Fan; 1 ♀, Guangxi, Nonggang, 6 May 2012, Z.-H. Fan.

Diagnosis

This species is similar to T. polyphemus, but differs from the latter by the anal tube (Fig. 13) irregularly pentagonal in dorsal view, phallobase with two processes near basal dorsal margin (Fig. 14b, c) and aedeagus with pair of small sheet processes (Fig. 14f) at apical 1/3 in lateral view .

Description

Body length: male 6.5–6.6 mm, female 6.7 mm; Forewing: male 5.3–5.5 mm, female 5.4 mm.

Coloration. General color brown, with irregular black mottling. Vertex with black speckles near base (Fig. 7). Eyes brown to black (Fig. 8). Frons (Fig. 9) with small black protuberance with light median line running from the upper margin, with narrow pale transverse clearer band under the black protuberance, to nearly the basal 1/3 of face, not reaching ventrally level of compound eyes, and pale spots near lateral margins. Clypeus brown with pale marks in middle line and rostrum brown. Forewings (Fig. 1) with irregular black mottling. Legs with tips of spines on hind tibiae and tarsi black.

Head and thorax. Head (Fig. 7) including eyes narrower than pronotum (0.76: 1.00). Vertex (Fig. 7) shorter in middle than the wide at base (0.57: 1.00), disc of vertex without median carina. Frons (Fig. 9) flat, with median carina only present at base, longer in middle than the widest breath (1.30: 1.00). Clypeus triangular, with distinct median carina and rostrum surpassing mesotrochanters. Pronotum (Fig. 7) with median carina obscure, lateral carina not reaching to the posterior margin. Mesonotum (Fig. 7) triangular, with median carina obscure, Hind tibiae each with 2 spines, spinal formula of hind leg 8–8–2 (male); 7–8–2 (female). Forewings (Fig. 10) elongate, 2.3 times as long as maximum breadth, ScP and Rp convergent near base, M four branched, MP1+2 bifurcates apically, MP3+4 bifurcates at middle part, CuA simple, not forked; CuP present, Pcu and A1 uniting in basal 2/3 of clavus. Hindwings (Fig. 11) with two big lobes, anal lobe reduced.

Male genitalia. Anal tube (Fig. 13) irregularly pentagonal in dorsal view, the widest breath in apical 1/3. Anal style (Fig. 13) short, located at the base 1/3 of anal tube. Pygofer (Fig. 12) narrow and curved in lateral view, irregular sub-quadrate. Genital styles (Fig. 12) short, without triangular prominence near dorsal margin before capitulum; capitulum of genital styles relatively short, subtrapezoidal on short neck. Phallobase with dorsal lobe cystiform at apical part, with irregular lobes in apical 1/4 in lateral view (Fig. 14a), bean-shaped process in basal dorsal margin (Fig. 14b), with big ansate process (Fig. 14c) in subbasal dorsal margin in lateral view; lateral lobe splitting into two stout branches, with stout sub-angular processes near apical (Figs 14d, 15d) ; ventral lobe short, not reaching to the tip of dorsal lobe, in ventral view ventral lobe with three lobes near apical part (Figs 14e, 15e). Aedeagus with pair of small sheet-like processes at apical 1/3 in lateral view (Figs 14f, 15f).

Female genitalia. Hind margin of sternum VII wide concavity, with curved process in middle in ventral view (Fig. 17). Anal tube (Fig. 16) sub-rectangular, truncated apically, longer in middle than the widest breadth (2.12: 1.00), anal style short, located at the basal 1/6 of anal tube. Anterior connective lamina of gonapophyses VIII with 7 lateral teeth bearing 7 keels in lateral group and 3 apical teeth (Fig. 18). Posterior connective lamina of gonapophyses IX (Figs 19, 20) long subtriangular, narrowing, median field with a slender shaft prominence (medial dorsal process) (Fig. 19); sublateral field with one triangular process on lateral margins; lateral field without process; ventroposterior lobes bent at obtuse angle (posterior ventral lobes) (Fig. 19). Gonoplacs (Fig. 21) without keels.

Etymology

The name is derived from the Latin noun “ansatus”, meaning phallobase with big handle-shaped process in subbasal dorsal margin in lateral view.

Host plant

Unknown.

Distribution

China (Guangxi).

Tetricodes fennahi Gnezdilov, 2015

Tetricodes fennahi Gnezdilov, 2015: 29.

Tetricodes polyphemus Fennah: Zhang and Chen 2009: 18; Chen et al. 2014: 118.

Material examined

♂ (holotype), China: Guizhou, Leigongshan National Natural Reserve (26°28'N, 108°17'E), 2 Aug. 2004, F.-L. Xu.

Distribution

China (Guizhou).

Tetricodes parvispinus Chang & Chen, sp. n.

Figs 3–4, 22–30

Type material

Holotype; ♂, China: Guizhou, Anlong, Xianheping (24°58'N, 105°36'E), 28 Aug. 2012, W.-B. Zheng; paratypes: 2♂♂, China, Guizhou, Anlong, Xianheping (24°58'N, 105°36'E), 28 Aug. 2012, W.-B. Zheng and J.-K. Long.

Diagnosis

This species is similar to Tetricodes anlongensis Chen, Zhang & Chang, 2014, all from Guizhou: Anlong, Xianheping, but can be distinguished from the latter by the anal tube (Fig. 28) irregular pentagonal in dorsal view, the apical margin truncated, the widest breath in apical 1/3; phallobase with small process in dorsal margin of long lobe-shaped process at base (Fig. 29b).

Description

Body length: male 6.4–6.8 mm; Forewing: male 5.3–5.5 mm.

Coloration. General color yellowish green, with irregularly black mottling (Fig. 3). Eyes yellow to brown (Fig. 23). Frons (Fig. 24) with medium black protuberance with light median line running from the upper margin, with broad pale transverse clearer band under the black protuberance to nearly the basal 1/2 of face, reaching or surpassing ventrally level of compund eyes, and pale spots near lateral margins. Clypeus brown with yellow mark in base and rostrum brown. Pronotum (Fig. 22) with three pale vurrucae betwwen median carina and lateral carina. Forewings (Fig. 3) with irregularly black mottling near margins.

Head and thorax. Head (Fig. 22) including eyes narrower than pronotum (0.67: 1.00). Vertex (Fig. 22) shorter in middle than the wide at base (0.62: 1.00), disc of vertex without median carina. Frons (Fig. 24) flat, with median carina only present at base, longer in middle than the widest breath (1.34: 1.00). Clypeus triangular, with distinct median carina and rostrum surpassing mesotrochanters. Pronotum (Fig. 22) with median carina obscure, lateral carina not reaching to the posterior margin. Mesonotum (Fig. 22) triangular, with median carina obscure, Hind tibiae each with 4 spines, spinal formula of hind leg 8–10–2. Forewings (Fig. 25) elongate, 2.4 times as long as maximum breadth, ScP and Rp convergent near base, MP four branched, MP1+2 bifurcates apically, MP3+4 bifurcates at middle part, CuA simple, not forked; CuP present, Pcu and A1 uniting in basal 2/3 of clavus. Hindwings (Fig. 26) with two big lobes, anal lobe reduced.

Male genitalia. Anal tube (Fig. 28) irregular pentagonal in dorsal view, the apical margin truncated, the widest breath in apical 1/3. Anal style (Fig. 28) short, located in the middle of anal tube. Pygofer (Fig. 27) narrow and curved in lateral view, irregular subquadrate. Genital styles (Fig. 27) short, without triangular prominence near dorsal margin before capitulum; capitulum of genital styles relatively short, subtrapezoidal on short neck. Phallobase with dorsal lobe with irregular lobes (Fig. 29a) at apical part in lateral view, with small spine in dorsal margin of long lobe-shaped process in base (Fig. 29b); lateral lobe splitting into two stout branches, apical part subtriangular (Figs 29c, 30c); ventral lobe short, ventral view ventral lobe with three lobe near apical part, the apical lobe narrowed (Figs 29d, 30d). Aedeagus with pair of big sheet-like processes at apical 1/3 in lateral view (Figs 29e, 30e).

Etymology

This species epithet is derived from combination of Latin root prefix “parv-” and “spine”, referring to the small spine in the dorsal margin of basal long process of aedeagus.

Host plant

Unknown.

Distribution

China (Guizhou).

Tetricodes similis Chang & Chen, sp. n.

Figs 5–6, 31–39

Type material

Holotype: ♂, China: Guizhou, Anlong, Xianheping (24°58'N, 105°36'E), 28 Aug. 2012, W.-B. Zheng.

Diagnosis

This species can be distinguished from other Tetricodes species by the anal tube (Fig. 37) irregular circular, with sunken trilateral at apical margin in dorsal view; phallobase with dorsal lobe with irregular collar-shaped and long lobe-shaped process (Fig. 38a); lateral lobe with a stout lobe-like prominence and hook-shaped prominence in apical part in ventral view (Fig. 39c). Aedeagus with big sheet-like processes in lateral view (Fig. 38e).

Description

Body length: male 5.8 mm, Forewing: male 4.9 mm.

Coloration. As in T. parvispinus Chang & Chen, sp. n., but clypeus black with yellow mark at base, rostrum brown and pronotum (Fig. 31) with four pale vurrucae between median carina and lateral carina.

Head and thorax. Head (Fig. 31) including eyes narrower than pronotum (0.67: 1.00). Vertex (Fig. 31) shorter in middle than the wide at base (0.63: 1.00). Frons (Fig. 33) longer in middle than the widest breath (1.30: 1.00). Clypeus triangular, with obscure median carina and rostrum surpassing mesotrochanters. Pronotum (Fig. 31) with median carina. Mesonotum (Fig. 31) with lateral carinae. Hind tibiae each with 4 spines, one of small spines near base, spinal formula of hind leg 8–10–2. Forewings (Fig. 34) elongate, 2.5 times as long as maximum breadth, ScP and Rp convergent near base, MP three branched, MP1+2 bifurcates apically, CuA simple, not forked; CuP present, Pcu and A1 uniting in basal 2/3 of clavus. Hindwings (Fig. 35) with two big lobes, anal lobe reduced.

Male genitalia. Anal tube (Fig. 37) irregular circular, with pitted triangular in apical margin in dorsal view, the widest breath in apical 1/2. Anal style (Fig. 37) long, located in the middle of anal tube. Pygofer (Fig. 36) narrow and curved in lateral view, irregular subquadrate. Genital styles (Fig. 36) short, with obscure triangular prominence near dorsal margin before capitulum; capitulum of genital styles relatively short, subtrapezoidal on short neck. Phallobase with dorsal lobe with irregularly collar-shaped process near apical part in lateral view (Fig. 38a), with long lobe-shaped process in dorsal margin in base (Fig. 38b); lateral lobe splitting into two branches, with stout lobe-like prominence and hook-shaped prominence in apical part in ventral view (Figs 38c, 39c); ventral lobe short, in ventral view ventral lobe with three lobe near apical part, the apical margin acute (Figs 38d, 39d). Aedeagus with pair of big sheet processes in apical 1/3 in lateral view (Figs 38e, 39e).

Etymology

This new species is named for its similarity to the above new species.

Host plant

Unknown.

Distribution

China (Guizhou).

Tetricodes polyphemus Fennah, 1956

Tetricodes polyphemus Fennah, 1956: 514.

Tetricodes songae Zhang & Chen, 2009 :19, figs 11-19, 23-27 ; Chen et al. 2014: 120, figs 2–62.

Material examined

Tetricodes polyphemus: 1♀, China: Hubei Province, Lichuan, Maoba, Xingdoushan National Natural Reserve (30°01'N, 109°02'E), 4 Aug. 2010, X.- F. Yu; 2♂♂, 1♀, Hunan Province, Badagongshan National Natural Reserve (29°39'N, 109°49'E), 3 Aug. 2013, Z.-M. Chang and L. Qun. Tetricodes songae: ♂ (holotype), Guizhou Province, Leishan, Leigongshan National Natural Reserve (26°28'N, 108°17'E), 1 Aug. 2004, Q.-Z. Song; 2♂♂, 1♀ (paratypes), same data as holotype.

Diagnosis

Frons with big black protuberance with light median line running from the upper margin, with broad pale transverse band under black protuberance, to nearly basal 2/3 of face, reaching or surpassing ventrally level of eyes, and pale spots near lateral margins of frons.

Distribution

China (Guizhou, Hubei, Hunan, Yunnan).

Remarks

The above synonymy is based on examination of the type specimens (male and female) of Tetricodes songae (from Guizhou Province) and the specimens of T. polyphemus collected from the type locality (Hubei Province, 1 female) and Hunan Province (male and female). Although Hunan, to which the locality of collecting the male material of T. polyphemus belongs, is different from the type locality (Hubei), the distance between the above localities is very close. The male genitalia of T. songae and T. polyphemus are identical.

Figures 1–6. 

Dorsal habitus of Tetricodes species. 1–2 Tetricodes ansatus Chang & Chen, sp. n. 3–4 Tetricodes parvulus Chang & Chen, sp. n. 5–6 Tetricodes similis Chang & Chen, sp. n.. Scale bars = 0.5 mm.

Figures 7–15. 

Tetricodes ansatus Chang & Chen, sp. n. 7 Head and thorax, dorsal view 8 Head and thorax, lateral view 9 Head, ventral view 10 Forewing 11 Hindwing 12 Male genitalia, lateral view 13 Anal segment, dorsal view 14 Aedeagus and phallobase, lateral view 15 Aedeagus and phallobase, ventral view. a–irregular lobes, b–bean-shaped process, c–ansate process, d–lateral lobe, e–ventral lobe, f–sheet-like processes. Scale bars = 0.5 mm.

Figures 16–21. 

Female genitalia. Tetricodes ansatus Chang & Chen, sp. n. 16 anal segment, dorsal view 17 Sternum VII, ventral view 18 Anterior connective lamina of gonapophyses VIII, lateral view 19 Posterior connective lamina of gonapophyses IX, dorsal view 20 Posterior connective lamina of gonapophyses IX, lateral view 21 Gonoplacs, lateral view. Scale bars = 0.5 mm. Abbreviations. lf–lateral field of posterior connective lamina of gonapophyses IX; mdp–medial dorsal process; mf–medial field of posterior connective lamina of gonapophyses IX; pvd–posterior ventral lobes; slf–sublateral field of posterior connective lamina of gonapophyses IX.

Figures 22–30. 

Tetricodes parvulus Chang & Chen, sp. n. 22 Head and thorax, dorsal view 23 Head and thorax, lateral view 24 Head, ventral view 25 Forewing 26 Hindwing 27 Male genitalia, lateral view 28 Anal segment, dorsal view 29 Aedeagus and phallobase, lateral view 30 Aedeagus and phallobase, ventral view. a–irregular lobes, b–long lobe-shaped process, c–lateral lobe, d–ventral lobe, e–sheet-like processes. Scale bars = 0.5 mm.

Figures 31–39. 

Tetricodes similis Chang & Chen, sp. n. 31 Head and thorax, dorsal view 32 Head and thorax, lateral view 33 Head, ventral view 34 Forewing 35 Hindwing 36 Male genitalia, lateral view 37 Anal segment, dorsal view 38 Aedeagus and phallobase, lateral view 39 Aedeagus and phallobase, ventral view. a–collar-shaped process, b–long lobe-shaped process, c–lateral lobe, d–ventral lobe, e–sheet-like processes. Scale bars = 0.5 mm.

Discussion

The species of Tetricodes are very similar in colouration and external morphology. Based on black protuberance of frons without light median line, Gnezdilov (2015) considered that the species T. polyphemus treated by Zhang and Chen (2009) is another new species (T. fennahi). Actually, the light median line is variable in the examined specimens of all species. By contrast, in addition to male genitalic differences, the frons with black protuberance and pale transverse band, have certain value: small black protuberance and narrow pale transverse band (together in basal 1/3) in T. ansatus Chang & Chen; medium black protuberance and narrow pale transverse band in T. fennahi (together in basal 1/2); big black protuberance and broad pale transverse band (together in basal 2/3) in T. polyphemus; big black protuberance and broad pale transverse band in T. anlongensis, T. parvispinus Chang & Chen and T. similis Chang & Chen (together in basal 1/2–2/3).

Acknowledgments

We are grateful to Mick Webb for proofreading and advices. This work was supported by the Program of Science and Technology Innovation Talents Team of Guizhou Province (No. 20144001), the Program of Excellent Innovation Talents, Guizhou Province (No. 20154021), the China Postdoctoral Science Foundation funded project (No. 180127) and the Program of Scientific Research Foundation for Introduced Talent of Guizhou University (Grant No. 201674).

References

  • Bourgoin T (1993) Female genitalia in Hemiptera Fulgoromorpha, morphological and phylogenetic data. Annales de la Société Entomologique France 93: 225–244.
  • Bourgoin T, Wang RR, Asche M, Hoch H, Soulier-Perkins A, Stroiński A, Yap S, Szwedo J (2015) From micropterism to hyperpterism: recognition strategy and standardized homology-driven terminology of the forewing venation patterns in planthoppers (Hemiptera: Fulgoromorpha). Zoomorphology 134(1): 63–77. https://doi.org/10.1007/s00435-014-0243-6
  • Chan ML, Yang CT (1994) Issidae of Taiwan (Homoptera: Fulgoroidea). Chen Chung Book, Taichung, 188 pp.
  • Che YL, Zhang YL, Wang YL (2013) A new genus of the tribe Parahiraciini from China, with notes on the tribe (Hemiptera: Fulgoroidea: Issidae). Zootaxa 3701(1): 76–82. https://doi.org/10.11646/zootaxa.3701.1.6
  • Chen XS, Zhang ZG, Chang ZM (2014) Issidae and Caliscelidae (Hemiptera: Fulgoroidea) from China. Guizhou Science and Technology PubIishing House, Guiyang, China, 242 pp.
  • Fennah RG (1956) Fulgoroidea from southern China. Proceedings of the California Academy of Sciences. San Francisco 28(4): 441–527.
  • Gnezdilov VM (2002) Morphology of the ovipositor in members of the subfamily Issinae (Homoptera, Cicadina, Issidae). Entomologicheskoe obozrenie 81(3): 605–626.
  • Gnezdilov VM (2003) Review of the family Issidae (Homoptera, Cicadina) of the European fauna, with notes on the structure of ovipositor in planthoppers. Chteniya pamyati N.A. Kholodkovskogo (Meetings in memory of N.A. Cholodkovsky) 56(1): 1–145.
  • Gnezdilov VM (2015) A new species of the genus Tetricodes Fennah (Hemiptera, Fulgoroidea, Issidae) from southern China. Entomotaxonomia 37(1): 27–30.
  • Gnezdilov VM (2016) On the taxonomic position of Issus reticulatus Bervoets, 1910 (Hemiptera: Fulgoroidea: Issidae) from Baltic Amber. Entomological Review 96(5): 631–633. https://doi.org/10.1134/S0013873816050092
  • Gnezdilov VM, Hayashi M (2016) New genus of the family Issidae (Hemiptera: Fulgoroidea) from Japan. Japanese Journal of Systematic Entomology 22(1): 47–49.
  • Gnezdilov VM, Wilson MR (2005) New genera and species of the tribe Parahiraciini (Hemiptera, Fulgoroidea, Issidae). Acta Entomologica Slovenica 13(1): 21–28.
  • Wang ML, Zhang YL, Bourgoin T (2016) Planthopper family Issidae (Insecta: Hemiptera: Fulgoromorpha): linking molecular phylogeny with classification. Molecular Phylogenetics and Evolution 105: 224–234. https://doi.org/10.1016/j.ympev.2016.08.012
  • Zhang ZG, Chen XS (2009) Review of the oriental Issid genus Tetricodes Fennah (Hemiptera: Fulgoroidea: Issidae) with the description of one new species. Zootaxa 2094: 16–22.