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Review of the leafhopper tribe Deltocephalini Dallas, 1870 (Hemiptera, Cicadellidae, Deltocephalinae) in Pakistan with description of a new species of Paramesodes
expand article infoHassan Naveed, Bismillah Shah§, Kamran Sohail|, Yalin Zhang, Keping Chen
‡ Jiangsu University, Zhenjiang, China
§ Zhejiang A&F University, Hangzhou, China
| The University of Agriculture, Peshawar, Pakistan
¶ Northwest A&F University, Yangling, China
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Abstract

A review of the leafhopper tribe Deltocephalini in Pakistan is provided, including comprehensive illustrated keys to all genera and species, along with their diagnosis and distribution. A new species of Paramesodes is described, P. dirensis sp. nov., which was discovered in the Upper Dir District of Khyber Pakhtunkhwa, Pakistan. A detailed description of this new species is provided together with photographs for visual reference. This tribe now has three genera and 13 species from Pakistan. The genus Deltocephalus Burmeister includes two species, and Maiestas Distant has nine species; Paramesodes Ishihara is now represented with two species.

Key words

Auchenorrhyncha, distribution, key, morphology

Introduction

Leafhoppers of the tribe Deltocephalini can be identified by their small to moderate size, tapering or parallel-sided clypellus, narrow lorum, linear connective with closely appressed anterior arms, connective fused to the aedeagus, and dorsal sculpturing of the first valvula imbricate. This tribe includes 74 genera and over 600 species distributed worldwide (Duan et al. 2020; Zhang et al. 2023). Until now, 12 species of Deltocephalini have been known from Pakistan (Naveed et al. 2021; Shah et al. 2021).

The genus Paramesodes Ishihara, comprising 16 Old World species, is poorly represented in Pakistan, with only a single previous species, P. lineaticollis (Distant), known. Wilson (1983) revised the genus and recognized one species from Europe and the Middle East, one species from Africa, and nine species from Asia. Five other species were subsequently included: P. montanus Rao, (1989) from India, P. wilsoni Rao & Ramakrishnan, (1990) from India, P. iraniensis Webb & Viraktamath, (2009) from Iran, and P. menghaiensis Li, Dai & Xing, (2011) and P. cangshanae Duan & Zhang, (2012) from China. Paramesodes is externally similar to Paramesus Fieber, Exitianus Ball, and Ctenurellina McKamey but they can be differentiated by having the forewing with the inner anteapical cell open basally (closed in Paramesus and Ctenurella), and the veins are always pale or white (usually brown in Exitianus). In the male genitalia the connective and aedeagus are fused (free in Paramesus and Exitianus), and the pygofer side has a dense covering of long, stout, pale macrosetae (2–6 dark or black macrosetae along apical margin in Exitianus) and the pygofer side has a large process (lower edge of pygofer pectinate in Ctenurella). This, together with the aedeagus, is the most important character for species identification (Duan and Zhang 2012).

In this paper, we provide a list of leafhoppers of the tribe Deltocephalini from Pakistan, illustrated keys to genera and species with their diagnostic characters, and a description and illustration of a new species, P. dirensis sp. nov.

Material and methods

All specimens were examined with a Leica ZOOM2000 stereomicroscope. Drawings were made using an Olympus drawing tube. Photos were taken by a ZEISS SteREO Discovery.V20 stereomicroscope equipped with a ZEISS AxiocamICc 5 camera that also provided measurements. Adobe Photoshop CS was used to compile photographs. Specimens from Pakistan are deposited in the various collections as indicated in the published records. Type specimens of the new species, examined and figured for this study, are deposited in the Entomological Museum, Northwest A&F University, Yangling, Shaanxi, China.

Taxonomy

Family Cicadellidae Latreille, 1802

Subfamily Deltocephalinae Dallas, 1870

Tribe Deltocephalini Dallas, 1870

Key to genera of Deltocephalini from Pakistan

1 Crown with broad black transverse submarginal band between eyes (Figs 1, 9); pygofer side with large process (Figs 3, 10) Paramesodes
Crown without black transverse band between eyes (Figs 12, 15, 18, 19); pygofer side without process 2
2 Aedeagal shaft short, robust, strongly curved dorsally, gonopore apical (Figs 13, 16) Deltocephalus
Aedeagal shaft long, slightly curved dorsally, with gonopore indistinct (Figs 20–25, 27, 29) Maiestas

Genus Deltocephalus Burmeister

Key to species of Deltocephalus from Pakistan (male)

1 Crown with six brown spots on anterior margin (Fig. 12); aedeagal shaft with shallow apical notch (Fig. 13) D. vulgaris
Crown with single brown spot on anterior margin adjacent to eyes (Fig. 15); aedeagal shaft without apical notch (Fig. 16) D. infirmus

Deltocephalus vulgaris Dash & Viraktamath

Figs 12–14

Deltocephalus (Deltocephalus) vulgaris Dash & Viraktamath, 1998: 4, figs 1–11 (India); Zhang and Duan 2011: 3, fig. 3A–H (China); Naveed et al. 2019: 285, figs 1A, B, 3A–D (Pakistan).

Diagnosis

This species can easily be identified by the color pattern and the aedeagus with a shallow apical notch.

Distribution

China, India, Pakistan.

Deltocephalus infirmus Melichar

Figs 15–17

Deltocephalus infirmus Melichar, 1903: 203, pl. 5 fig. 11 (Sri Lanka); Jassargus infirmus: Ishihara 1961: 244, figs 53–58 (misidentification); Deltocephalus infirmus: Webb and Viraktamath 2009: 13, fig. 10; Naveed et al. 2019: 285, figs 1C, 3D–G (Pakistan).

Diagnosis

A similar species to D. vulgaris but differs by having the crown with only a single brown spot and the aedeagal shaft without an apical notch.

Distribution

India, Pakistan, Sri Lanka, Thailand.

Genus Maiestas Distant

Key to species of Maiestas from Pakistan (males)

1 Overall color dark brown; forewing with sub basal and subapical irregular white transverse band (Fig. 18) M. albomaculata
Color not as above (Fig. 19) 2
2 Crown, face, and thorax with black patches (Fig. 19) M. maculata
Crown, face, and thorax without black patches 3
3 Forewing with extra cross-veins, at least in clavus 4
Forewing without extra cross-veins 5
4 Aedeagus with a large subapical ventral process (Fig. 20) M. indica
Aedeagus with a short apical ventral process (Figs 21, 22) M. pruthii
5 Aedeagus with pair of short lateral processes (Fig. 23) M. trispinosa
Aedeagus without lateral processes 6
6 Aedeagus in lateral view similar in width in distal half (Fig. 24) M. subviridis
Aedeagus in lateral view evenly tapered from base to apex 7
7 Style apophysis broadest sub basally (Fig. 26); aedeagal shaft in lateral view not sinuate (Fig. 25) M. tareni
Style apophysis broadest at base (Fig. 28); aedeagal shaft in lateral view slightly sinuate (Fig. 27) M. sinuata

Maiestas albomaculata (Dash & Viraktamath)

Fig. 18

Deltocephalus (Recilia) albomaculatus Dash & Viraktamath, 1998: 12, figs 29–34 (India); Maiestas albomaculata: Webb and Viraktamath 2009: 21; Maiestas albomaculata: Naveed et al. 2019: 287, figs 1E–I, 3H, I (Pakistan); Shah et al. 2021: 403, fig. 1A–D (Pakistan).

Diagnosis

This species differs from other species of Maiestas in color and male genitalia, including the dorsolateral, laminate serrations of the aedeagal shaft.

Distribution

Pakistan, India.

Maiestas indica (Singh-Pruthi)

Fig. 20

Allophleps indica Singh-Pruthi, 1936: 120, fig. 132; pl. 9 fig. 3 (Pakistan); Allophleps delhiensis Rao & Ramakrishnan, 1990: 111, figs 1–9 (India), synonymized by Dash and Viraktamath 1998: 35; Deltocephalus (Recilia) indicus: Dash and Viraktamath 1998: 35–36, fig. 305 (India); Maiestas indica: Webb and Viraktamath 2009: 21; Naveed et al. 2019: 287; Shah et al. 2021: 403, fig. 1E (Pakistan).

Diagnosis

This species can be identified by the aedeagus, which has a distinctive, large, subapical ventral process, the forewings which have accessory cross-veins, and the shorter head.

Distribution

Pakistan, India.

Maiestas maculata (Singh-Pruthi)

Figs 19, 29

Cicadula maculata Singh-Pruthi, 1930: 58–59, figs 80, 81, pl. 5 fig. 2 (India); Thamnotettix prabha Singh-Pruthi, 1930: 62, figs 85, 86, pl. 5 figs 6, 6a (India), synonymized by Webb and Viraktamath 2009: 41; Recilia prabha: Ghauri 1980: 166–169, figs 1, 3–11; Deltocephalus (Recilia) maculata: Dash and Viraktamath 1998: 32, figs 260–269 (India); Maiestas maculata: Webb and Viraktamath 2009: 22; Maiestas maculata: Zhang and Duan 2011: 37–39, figs 33–35, pl. 4 fig. E, pl. 5 fig. P, pl. 6 fig. P (China); Shah et al. 2021: 404, fig. 2A–I (Pakistan).

Diagnosis

This species can be distinguished from other Maiestas species by its habitus, which has variable black patches on the head and thorax, and the shape of its aedeagus and style.

Distribution

China, India, Pakistan.

Maiestas pruthii (Metcalf)

Figs 21, 22

Deltocephlaus notatus Singh-Pruthi, 1936: 128–129, fig. 139, pl. 9 fig. 10 (Pakistan) (primary homonym: Deltocephalus notatus Melichar, 1896); Deltocephalus pruthii Metcalf, 1967: 1173 (nom. nov. pro D. notatus Singh-Pruthi, 1936); Deltocephalus (Recilia) pruthii: Dash and Viraktamath 1998: 22, 23, figs 150–158 (India); Maiestas pruthii: Webb and Viraktamath 2009: 20, new combination; Khatri and Webb 2010: 11, pl. 2a fig. 13, misidentification; Naveed et al. 2019: 286, fig. 2A–C (incorrectly cited as M. subviridis; H. Naveed pers. comm.), Fig. 3i, misidentification; Shah et al. 2021: 406, fig. 4F–L (Pakistan).

Diagnosis

This species has a relatively long, acute head, with a pair of inverted U-shaped markings basally, and forewings with extra cross veins. The identity of this species is based on the figures of Dash and Viraktamath (1998).

Distribution

India, Pakistan.

Maiestas setosa (Ahmed, Murtaza & Malik)

Recilia setosa Ahmed et al., 1988: 412, fig. 2 (Pakistan); Maiestas setosa: Webb and Viraktamath 2009: 20; Naveed et al. 2019: 287; Shah et al. 2021: 406 (Pakistan).

Diagnosis

The identity of this species remains uncertain due to the limitations of the original description and the accompanying figures. Additionally, the type series from Karachi, which was indicated in the original account as deposited in the Zoological Museum of the University of Karachi (Ahmed et al. 1988), is unavailable (Khatri and Webb 2010: 11). Until the type material can be studied, pinpointing the precise classification of this species will be challenging.

Distribution

Pakistan.

Maiestas sinuata Shah & Duan

Figs 27, 28

Maiestas sinuata Shah & Duan in Shah et al., 2021: 406, fig. 3A–H (Pakistan).

Diagnosis

This species differs in appearance of its habitus, presence of fine, apical setae on the subgenital plate, the style having a thicker apical process than other species, and the aedeagus lacking a ventrobasal “heel”.

Distribution

Pakistan.

Maiestas subviridis (Metcalf)

Fig. 24

Stirellus subviridis Metcalf, 1946: 125; Deltocephalus (Recilia) subviridis: Dash and Viraktamath 1998: 24, figs 166–172 (India); Maiestas subviridis: Webb and Viraktamath 2009: 19, fig. 40; Maiestas subviridis: Khatri and Webb 2010: 11, pl. 2b, c, fig. 12 (Pakistan); Zhang and Duan 2011: 19, fig. 17, pl. 2 fig. H (China); Naveed et al. 2019: 287; Shah et al. 2021: 408, fig. 4A–E (Pakistan).

Diagnosis

This species can be distinguished by the rounded apex of the aedeagus which bears a very short apical spine.

Distribution

China, India, Pakistan, Pacific.

Maiestas tareni (Dash & Viraktamath)

Figs 25, 26

Deltocephalus (Recilia) tareni Dash & Viraktamath, 1995: 74–76, figs 1–15; Dash and Viraktamath 1998: 16, figs 78–84 (India); Maiestas tareni: Webb and Viraktamath 2009: 22; Khatri and Webb 2010: 11, pl. 2d, fig. 11 (Pakistan); Zhang and Duan 2011: 20 (China); Naveed et al. 2019: 290, figs 2G–I, 3N, O; Shah et al. 2021: 408, fig. 5A–Z (Pakistan).

Diagnosis

This species can be identified by its relatively straight and stout style, apophysis with a serrated inner margin, and the aedeagus in lateral view evenly tapered from base to apex and relatively straight.

Distribution

China, India, Pakistan.

Maiestas trispinosa (Dash & Viraktamath)

Fig. 23

Deltocephalus (Recilia) trispinosus Dash & Viraktamath, 1998: 35, figs 296–304 (India); Maiestas trispinosa: Webb and Viraktamath 2009: 38; Maiestas trispinosa Shah et al., 2021: 408, fig. 6A–I (Pakistan).

Diagnosis

This species can easily be distinguished from the others by the lateral, spine-like processes of the aedeagus.

Distribution

India, Pakistan.

Parasmesodes Ishihara

Paramesodes Ishihara, 1953: 45. Type species: Athysanus albinervosus Matsumura, 1902.

Distribution

Palearctic, Oriental, and Ethiopian regions.

Remarks

Previously, only 1 species was recorded from Pakistan. This study adds one more new species to the genus, bringing the total to two for the country.

Key to species of Paramesodes from Pakistan (males)

1 Pale yellowish species (Fig. 1); pygofer broadly rounded posteriorly, with process in lateral view straight distally, apex directed posteriorly (Fig. 3); aedeagal shaft recurved distally in lateral view (Fig. 7); constricted subapically in ventral view (Fig. 6) P. dirensis sp. nov.
Dark yellowish species (Fig. 9); pygofer oval posteriorly, with process in lateral view directed dorsally distally (Fig. 10); aedeagal shaft evenly curved dorsally in lateral view (Fig. 10), not constricted preapically in ventral view (Fig. 11) P. lineaticollis

Paramesodes dirensis sp. nov.

Figs 1–8

Description

Length : male 5.2–6.1 mm, female 6.1–6.4 mm. Coloration: pale, with brown markings (Fig. 1). Crown with broad, black, transverse submarginal band between eyes (Fig. 1). Face pale yellow, with brown, transverse striations on clypeus (Fig. 2). Pronotum with medial dark brown longitudinal marking as well as three brown longitudinal markings on each side (Fig. 1). Scutellum with median, longitudinal, dark brown markings and pale brown lateral markings (Fig. 1). Forewings with variable brown markings; veins prominent and white (Fig. 1). Legs pale, with brown markings.

Male genitalia : pygofer lobe broad basally, narrowing apically, forming a rounded-oval apex, with long, yellowish-brown spines extending beyond apical margin, a large process arising near medial dorsal margin and straight apically, surpassing pygofer lobe, without any bend (Figs 3, 4). Subgenital plates triangulate; macrosetae uniseriate laterally (Fig. 5). Valve triangular (Fig. 5). Style as in Figure 5. Connective fused to aedeagus, with arms closely appressed distally (Figs 6, 7). Aedeagus tubular, tapering apically, recurved in lateral view, constricted preapically in ventral view; gonopore apical (Figs 6, 7).

Female. Same in appearance as male. Seventh sternum with lateral margins not extended, posterior margin with median projection, rounded (Fig. 8).

Materials examined

Holotype ♂, Pakistan: Khyber Pakhtunkhwa: Upper Dir, 35°9'55.89"N, 72°2'48.54"E, 1840 m, 24.07.2019, Hassan Naveed leg., sweep net. Paratypes 8♂, 5♀, same data as holotype.

Etymology

This species is named after type locality, the Upper Dir in Khyber Pakhtunkhwa.

Remarks

Paramesodes dirensis sp. nov. is similar to its congeners in general appearance, but it differs from those species in the combination of male genitalia features, i.e., the pygofer is oval posteriorly with a relatively straight process distally and the aedeagal shaft is distally recurved in lateral view and constricted preapically in ventral view. In the Wilson’s (1983) key, the new species runs to couplet 7 along with P. lineaticollis.

Figures 1–8. 

Paramesodes dirensis sp. nov. 1 habitus, dorsal view 2 face 3 male pygofer, lateral view 4 male pygofer, dorsal view 5 subgenital plate, valve and style, dorsal view 6 connective and aedeagus, dorsal view 7 connective and aedeagus, lateral view 8 female abdominal tip and 7th sternum, dorsal view.

Paramesodes lineaticollis (Distant)

Figs 9–11

Paramesodes lineaticollis (Distant, 1908: 294, Paramesus) (India); Wilson 1983: 21, 22, figs 23–29.

Paramesodes ishurdii Mahmood & Meher, 1973: 135 (Pakistan), synonymized by Wilson 1983: 21.

Materials examined

♂, Pakistan: Khyber Pakhtunkhwa: Shinkiari, 34°28'19.1064"N, 73°16'14.3004"E, 22.07.2018, Bismillah Shah leg., sweep net.

Distribution

Bangladesh, China, India, Indonesia, Pakistan, Philippines, Taiwan, Turkey.

Diagnosis

The male pygofer processes is distinct, directed ventrally or postero-ventrally over basal half, and turned abruptly posteriorly immediately distad of its midlength. Mahmood and Meher (1973) reported this species as P. ishurdii for the first time from Pakistan, but that species was later synonymized by Wilson (1983).

Figures 9–11. 

Paramesodes lineaticollis 9 habitus, dorsal view 10 male pygofer, lateral view 11 connective and aedeagus, dorsal view.

Figures 12–29. 

12–14 Deltocephalus vulgaris Dash & Viraktamath: 12 habitus, dorsal view 13 aedeagus and connective, dorsal view 14 aedeagus and connective, lateral view 15–17 D. infirmus (Melichar): 15 habitus, dorsal view 16 aedeagus and connective, dorsal view 17 aedeagus and connective, lateral view 18 Maiestas albomaculata (Dash & Viraktamath) habitus, dorsal view 19 M. maculata (Singh-Pruthi) habitus, dorsal view 20 M. indica (Singh-Pruthi) aedeagus and connective, lateral view (after Dash & Viraktamath, 1998) 21, 22 M. pruthii (Metcalf): 21 aedeagus and connective, dorsal view (after Dash & Viraktamath, 1998) 22 apex of aedeagus, lateral view 23 M. trispinosa (Dash & Viraktamath) aedeagus and connective, dorsal view 24 M. subviridis (Metcalf) aedeagus, lateral view (after Khatri & Webb, 2010) 25, 26 M. tareni (Dash & Viraktamath): 25 aedeagus and connective, lateral view 26 subgenital plate, valve and styles, dorsal view 27, 28 M. sinuata Shah & Duan: 27 aedeagus and connective, lateral view 28 subgenital plate, valve and styles, dorsal view 29 M. maculata (Singh-Pruthi) aedeagus, lateral view.

Figures 30, 31. 

Paramesodes lineaticollis 30 subgenital plates, valve and styles, dorsal view 31 aedeagus and connective, lateral view.

Acknowledgements

We gratefully acknowledge Prof. C.A. Viraktamath (University of Agricultural Science, Bangalore, India), and Dr M. Webb (Natural History Museum, London, UK) for checking figures and confirming the new species. We are very grateful for John Richard Schrock (Emporia State University, USA) for revising the manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was funded by the National Natural Science Foundation of China (31861143051, 31872425).

Author contributions

All authors have contributed equally.

Author ORCIDs

Hassan Naveed https://orcid.org/0000-0002-9232-6299

Bismillah Shah https://orcid.org/0000-0002-8407-8627

Kamran Sohail https://orcid.org/0000-0003-1625-1130

Yalin Zhang https://orcid.org/0000-0002-1204-9181

Keping Chen https://orcid.org/0000-0001-5254-2299

Data availability

All of the data that support the findings of this study are available in the main text.

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