Two new species of Platensina Enderlein (Diptera, Tephritidae, Tephritinae, Dithrycini) from India.

Two new species of Platensina Enderlein, P.rabbanii David & Hancock, sp. nov., and P.flavistigma David & Hancock, sp. nov., are described from Meghalaya and southern India, respectively. Platensinarabbanii can be differentiated from P.alboapicalis Hering by the presence of a single hyaline indentation in cell r1 and the apical hyaline band in cell r2+3 restricted to the apex; P.flavistigma differs from P.quadrula Hardy by the presence of a yellow/fulvous pterostigma and shape of the epandrium. DNA barcode sequences of P.acrostacta (Wiedemann), P.flavistigma and P.platyptera Hendel were obtained and reported. Postabdominal descriptions and illustrations of P.acrostacta, P.platyptera and P.zodiacalis (Bezzi) are also provided along with keys to all 23 species and the 7 known from India. K. J. David, D. L. Hancock, K. Sachin, R. G. Gracy, S. Salini.

Introduction

Enderlein is predominantly an Oriental and Australasian genus with 24 species recognized by Norrbom et al. (1999) [some generically misplaced] and 21 by Hancock (2012). They are medium-sized flies with broad, dark brown wings with hyaline indentations and spots. Host plants are not recorded except for (Wiedemann), reared from stem galls of an undetermined species in southern India (Hardy 1973; Hancock 2012). Agarwal and Sueyoshi (2005) listed five species from India, while David and Ramani (2011) provided keys to four species from peninsular India and the Andaman and Nicobar Islands. Hancock (2012) recorded Hendel and Hardy from India, regarded records of (Walker) from India as misidentifications of and provided a key to species of . In this paper two new species, one collected from Meghalaya and one from southern India, are described, along with descriptions of postabdominal structures of other species recorded from India except Hering and Hering, as specimens of these two taxa were not available for study. A key to species of recorded from India is provided, together with a revised key to all known species.

Material and methods

Specimens studied are deposited in the National Insect Museum, ICAR – National Bureau of Agricultural Insect Resources, Bengaluru, India ().

Collections were done by sweep netting. Images of specimens were taken using a Leica DFC 420 camera mounted on a Leica M205A stereo zoom microscope; images of genitalia were taken using an 8 MP camera temporarily attached to a Leica DM 1000 compound research microscope; the images were stacked and combined to a single image using Combine ZP (Hadley 2011). Measurements of male and female genitalia were taken using Leica Automontage Software, LAS 3.4. Terminology adopted here follows White et al. (1999) and wing terminology by Cumming and Wood (2017).

One hind leg was removed from one specimen of each of three species and used for DNA extraction. The DNA extraction was performed using a DNeasy Blood and Tissue Kit (Qiagen India Pvt. Ltd.) following the manufacturers’ instruction. For the molecular study, the standard DNA barcoding region of the mitochondrial COI gene was sequenced and the PCR was performed using the Universal COI primers (LCO1490/HCO2198) (David et al. 2020). The sequences were annotated using NCBI Blast tools and submitted to the NCBI GenBank Database where accession numbers were obtained ( – MT019893; – MT019891; – MW448367).

The pairwise genetic distance between three species of viz., , and has been calculated using mitochondrial COI gene sequences. Analyses were conducted using the Maximum Composite Likelihood model (Tamura et al. 2004). This analysis involved 4 nucleotide sequences. Codon positions included were 1st+2nd+3rd. All positions with less than 95% site coverage were eliminated, i.e., fewer than 5% alignment gaps, missing data, and ambiguous bases were allowed at any position (partial deletion option). There were a total of 557 positions in the final dataset. Evolutionary analyses were conducted in MEGA11 (Tamura et al. 2021).

Results

Taxonomy

Enderlein, 1911

CD4538D0-ABB2-50A8-8FFD-67BE8F501E55

Enderlein, 1911: 454. Type species:

Bezzi, 1913: 153. Type species:

Diagnosis.

Medium-sized flies (4–5 mm long), with frons as wide as long, three frontal setae, two orbital setae, well developed ocellar setae. First flagellomere shorter than face, with short-pilose arista, face usually fulvous, black in males of a few species. Scutum grey pubescent with yellow-white reclinate setulae; scutellum flat with one or two pairs of setae, apical pair less than half length of basal setae or absent. Wing broad, often distinctly angled along posterior margin, dark brown with hyaline indentations and subhyaline spots. Abdomen predominantly black with fulvous lateral regions. Epandrium broad, without demarcation between epandrium and lateral surstylus, lateral surstylus broad, epandrium elongate-oval in posterior view; medial surstylus with well sclerotised prensisetae (lateral one broader than medial one), proctiger not higher than epandrium, glans of phallus stout, with single sclerotised acrophallus. Taeniae short (0.25 of eversible membrane); spicules on eversible membrane conical; aculeus dorsoventrally flattened, tip conical, with reduced preapical setae; spermathecae club-shaped, with numerous papillae.

Key to all known species of

Modified from Hancock (2012).

New species

David & Hancock sp. nov.

D3B6ED8C-E85E-571D-8A26-650BD863112E

http://zoobank.org/FF51328D-55A7-49B0-934F-48D1D61EF92D

Type locality.

India: Meghalaya, East Khasi Hills, Laitsopliah.

Type data.

male, pinned. Original label: “INDIA: Meghalaya, East Khasi Hills, Laitsopliah, 17.iii.2021, Rabbani M. K.” (NIM).

This species is similar to Hering from Burma in the presence of an apical hyaline band extending from cell r2+3 to cell m1 but can be differentiated by the presence of a single hyaline indentation in cell r2+3, versus two hyaline indentations and spot in ; the apical hyaline band restricted to the apical one-third of cell r2+3, versus the whole of apex of cell r2+3 in ; and the hyaline indentations in cell cua ending well before vein CuA, unlike in where they almost reach vein CuA.

Habitus (lateral) of male of David & Hancock, sp. nov.

Description.

Male. Medium-sized fly (4.03 mm long) with broad, dark brown wing with hyaline apex.

(Fig. 2): nearly as high as long; frons fulvous with three pairs of frontal setae and two pairs of orbital setae (posterior one white); ocellar triangle fuscous, with well developed ocellar setae; medial vertical seta black; lateral vertical seta, paravertical seta and postocellar seta lanceolate and white; postocular setae black interspersed with prominent, white lanceolate setae. Face fulvous without any markings. Scape and pedicel fulvous, first flagellomere less than half length of face, concolorous with frons, arista short pilose. Gena narrow, with prominent genal seta, subvibrissal setae present.

Figures 2–8.

David & Hancock, sp. nov. 2 head (lateral view) 3 thorax (dorsal view); 4 abdomen (dorsal view) 5 thorax (lateral view) 6 wing 7 epandrium (lateral view) 8 epandrium (posterior view).

(Figs 3, 5): Scutum uniformly grey microtrichose, with creamy-white setulae. Chaetotaxy well developed: 1 postpronotal, 1 presutural supra-alar, 1 anterior notopleural, 1 posterior notopleural, 1 postsutural supra-alar, 1 dorsocentral placed in line with postsutural supra-alar, 1 postalar, 1 intra-alar, and 1 prescutellar acrostichal seta. Scutellum brown with two pairs of scutellar setae; apical one less than half length of basal one. Anepisternum grey, with single seta near phragma; anepimeron grey, with single seta, katepisternum fulvous, with single seta; anatergite dark brown; katatergite and meron fulvous. Legs predominantly fulvous without any black/dark markings except basal, brown longitudinal streaks present on ventral surface of mid and hind femora; forefemur with single row of 4 ventral setae.

(Fig. 6) broad, with posterior margin evenly rounded, length 4.47 mm; length/width ratio-2.10, predominantly dark brown, with hyaline indentations and spots; cell bc hyaline, cell c predominantly moderate brown, with narrow basal and broader medial hyaline areas, pterostigma entirely moderate brown without any hyaline markings, cell r1 with one trapezoidal basal hyaline indentation not extending beyond vein R2+3, cell r4+5 with elongate subbasal hyaline spot, cell dm with large hyaline spot at basal third, cell cua with two short hyaline indentations not reaching vein CuA, and apical hyaline band extending from apex of cell r2+3 to cell m; anal lobe and alula dark brown.

(Fig. 4): Entirely black with yellowish orange patches laterally on tergites 1–2.

(Figs 7, 8). Epandrium broad, lateral surstylus as broad as epandrium and connected at acute angle to epandrium (Fig. 7); apex of lateral surstylus blunt (in lateral view); proctiger short, smaller than epandrium; epandrium elongate oval in posterior view, medial surstylus with well developed prensisetae (lateral one broader than medial one).

Female. Unknown

Distribution.

Meghalaya (Northeast India).

Habitat.

Marshy grasslands.

Etymology.

The species is named after the collector, Rabbani Mehaboob K. It is a noun in apposition.

David & Hancock sp. nov.

C6AB1448-B895-59B5-9D5F-B7124012BA6C

http://zoobank.org/7512FEB5-3888-4CFF-8229-80C146706273

:

India: Karnataka, Bangalore, Attur.

male, pinned. Original label: “INDIA: Karnataka, Bangalore, Attur, 05.ii.2020, Sachin K (NIM)”. : Periyakulam, 30.iii.2012, David, K. J. (1♂ NIM); India, Tamil Nadu, Thandikudi, C.R.S, 31.iii.2012, David, K. J. (1♂ NIM).

This species is similar to Hardy from Thailand, Cambodia and Vietnam in the presence of two broad quadrate areas in wing cell r1, an enlarged basal discal spot in cell r4+5 and large, broad hyaline indentations in cell cua. It can be differentiated by the lack of silvery facial spots and white rather than silvery parafacial stripes in males, as well as the angulate posterior wing margin, pterostigma predominantly fulvous/yellow and pale brown, and epandrium of uniform width throughout its length, whereas in the parafacial is silvery rather than white and facial silvery spots are present lateroventrally in males, the wing is evenly rounded posteriorly, the pterostigma is hyaline basally and dark brown apically, and the epandrium tapers apically. This species was mistakenly listed as from India by Hancock (2012).

Male. Medium-sized fly (4.10–4.99 mm long) with broad, angulate wing with fulvous markings (Figs 9, 10).

Figures 9–10.

Habitus of David & Hancock, sp. nov. 9 dorsal view 10 lateral view.

(Figs 11, 12). as high as long; frons fulvous with three pairs of frontal setae and two pairs of orbital setae; ocellar triangle dark brown, with well-developed ocellar seta; medial vertical seta black; lateral vertical seta, paravertical seta and postocellar setae white; postocular setae black interspersed with prominent, white lanceolate setae. Face fulvous with broad orange-brown patches on ventral half of antennal groove. Scape and pedicel fulvous, first flagellomere shorter than face, concolorous with frons, arista short pilose. Parafacial alongside ventral half of face distinctly whitish. Gena narrow, with prominent genal seta, subvibrissal setae present.

Figures 11–16.

David & Hancock, sp. nov. 11 head (profile view) 12 head (frontal view) 13 abdomen (dorsal view) 14 thorax (lateral view) and legs 15 scutum 16 wing.

(Figs 14, 15). Scutum uniformly grey microtrichose with creamy-white setulae. Chaetotaxy well developed: 1 postpronotal, 1 presutural supra-alar, 1 anterior notopleural, 1 posterior notopleural, 1 postsutural supra-alar, 1 dorsocentral placed in line with postsutural supra-alar, 1 postalar, 1 intra-alar, and 1 prescutellar acrostichal seta. Scutellum grey with two pairs of scutellar setae; apical one less than half length of basal one. Anepisternum grey, with single seta near phragma; anepimeron grey, with single seta, katepisternum fulvous, with single seta; anatergite dark brown; katatergite and meron fulvous. Legs predominantly fulvous without any black/dark markings; forefemur with single row of 4 ventral setae.

(Fig. 16) broad, angulate, length 4.44–4.99 mm, length/width ratio, 1.92–2.05; predominantly dark brown with hyaline indentations and spots; cell bc hyaline, cell c predominantly hyaline with pale basal and medial infuscations, pterostigma fulvous basally, dark brown apically, cell r1 with two broad quadrate indentations and small posterior subapical hyaline spot, the proximal quadrate indentation extended to vein R4+5, cell r4+5 with large circular hyaline spot near base and large, semicircular hyaline apical spot, cell dm with two large hyaline spots, cell m with two marginal hyaline marks, subbasal one much larger than subapical one, cells r2+3 and r4+5 with indistinct and isolated pale brown subapical spots, cell cua with three large marginal hyaline indentations, basal two almost reaching vein CuA, and anal lobe with two broad subhyaline markings, proximal one crossing into cell cua.

(Fig. 13). Entirely black with yellowish orange patches laterally on tergites 1–3.

Epandrium broad, lateral surstylus as broad as epandrium (Fig. 17); apex of lateral surstylus blunt (in lateral view); proctiger short, smaller than epandrium; epandrium oval in posterior view (Fig. 18); medial surstylus shorter than lateral surstylus and with well developed prensisetae (lateral one broader than medial one). Phallus elongate (1.78 mm long), with sclerotised acrophallus (Fig. 19).

Figures 17–19.

David & Hancock, sp. nov. 17 epandrium (lateral view) 18 epandrium (posterior view) 19 glans of phallus.

Female. Unknown

The specific name is derived from two Latin words flavus (=yellow) and stigma (=ptersotigma).

Karnataka and Tamil Nadu (southern India).

DNA Barcode.

NCBI GenBank accession number – MT019893 (1♂, India: Karnataka, Bangalore, Attur, 17.v.2018, Prabhu, G.)

Taxonomic notes on other species of from India

(Wiedemann)

6D6AB33A-6606-5F68-A14E-1CDE148F00A8

Wiedemann, 1824: 54. Type locality: India-orientali [east India].

Macquart, 1843: 387. Type locality Coromandel coast, Tamil Nadu, India.

Walker, 1849: 1030. Type locality North Bengal, India.

Walker, 1849: 1028. Type locality ‘Bahia, Brazil’ [

Material examined.

India: 1♂, Karnataka, Tumkur, Kunigal, 05.iv.2013, Prabhu G. “leg”; 1♂, Karnataka, Bengaluru, Hebbal, 28.xi.2014, Prabhu G. “leg”; 1♂, Karnataka, Bengaluru, Attur, 24.iii.2016, Prabhu G. “leg”; India. 1♀, Karnataka, Chikkaballapur, 12.iv.2016, Prabhu G. “leg”; 1♂,1♀ Karnataka, Bengaluru, Attur, 16.v.2017, Prabhu G. “leg”; 1♂,1♀, Karnataka, Bengaluru, Attur, 20.vi.2017, Prabhu G. “leg”; 1♀, Karnataka, Bengaluru, Attur, 04.vii.2017, Prabhu G. “leg”; 1♂, Karnataka, Bengaluru, Attur, 07.ii.2018, Prabhu G. “leg”; 1♀, Karnataka, Bengaluru, Attur, 21.iii.2018, Prabhu G. “leg”; 1♂,1♀, Karnataka, Bengaluru, Attur, 24.iv.2018, Prabhu G. “leg”; 1♂, Karnataka, Bengaluru, Hebbal, 03.v.2018, Prabhu G. “leg”; 1♂,1♀, Karnataka, Bengaluru, Attur, 17.v.2018, Prabhu G. “leg”; 2♀, Karnataka, Bengaluru, Attur, 24.v.2018, Prabhu G. “leg”; 1♂, Karnataka, Bengaluru, Attur, 14.vi.2018, Prabhu G. “leg”; 1♂, Karnataka, Bengaluru, Attur, 14.x.2019, Sachin K. “leg”; 1♂, Kerala, Kasargod, C.P.C.R.I, 17.ii.2015, Prabhu, G.”leg”, 1♀,1♂, Tamil Nadu, Periyakulam, 30.iii.2012, David, K.J.”leg”, 1♂, Tamil Nadu, Thandikudi, C.R.S, 01.iv.2012, David, K.J. “leg”(NIM).

Diagnosis

(Figs A medium-sized species separated from the similar Hering largely by the black face in males; length of male (4.29–4.47 mm), of female (5.45–5.49 mm). This species was adequately described by Hardy (1973) except for detailed structures of male and female postabdomen. Epandrium broad, not demarcated from surstylus (profile view); proctiger smaller than epandrium. Epandrium and surstyli elongate oval in posterior view; prensisetae well developed. Phallus elongate (2.71 mm); glans with a sclerotised rod. Oviscape (1.21 mm) dark brown to black, dorsoventrally flattened; eversible membrane shorter than oviscape (0.96 mm) with conical spicules in the proximal region and triangular spicules towards distal end. Aculeus (1.01 mm) as long as eversible membrane and with triangulate apex. Spermatheca elongate oval with numerous papillae.

(Wiedemann) 20 habitus (dorsal) 21 habitus (lateral).

(Wiedemann) 22 epandrium (lateral) 23 epandrium (posterior) 24 glans of phallus 25 ovipositor 26 spicules on proximal end of eversible membrane 27 spicules on distal end of eversible membrane 28 spermatheca 29 aculeus 30 aculeus tip (100×).

This species is known from India (Karnataka, Tamil Nadu and Kerala) and Sri Lanka to Cambodia (Hancock 2012).

NCBI GenBank accession number – MT019891 (1♂, India: Karnataka, Bangalore, Attur, 29.v.2019, Sachin, K.).

Hering

49D86386-123D-57D7-A417-73943FC0E09E

Hering, 1941: 71. Type locality Lonaula, Maharashtra, India.

This species is similar to but can be differentiated primarily by the yellow face in males and larger basal spot in cell dm. Specimens were not available for study but photographs of both sexes have been examined: 2♂, 2♀, India: Rajasthan, Jodhpur District, 10 km SW Jodhpur, Machia Safari Pk, Malaise in dry wash 29.II–5.III.2008, 300 m, (in California Academy of Sciences, San Francisco, California, USA).

This endemic Indian species is known only from Maharashtra and Rajasthan.

Hendel

13C1560B-0286-59E1-A532-CE0324B851AC

Hendel, 1915: 461. Type locality Taihorin, Taiwan.

Curran, 1936: 29. Type locality Tai Lagoon, Malaita, Solomon Is.

Malloch, 1939: 459. Type locality Gordonvale, Qld, Australia.

: authors,

India: 1♀, Karnataka, Bengaluru, G.K.V.K, 02.ii.2012, David K.J. “leg”; 1♂, A&N Islands, Middle Andamans, Kadamtala, 09.iii.2012, David, K.J. “leg”; 1♀, Karnataka, Mandya, Maddur, 09.i.2013, David, K.J. “leg”; 1♀, Karnataka, Uttara Kannada, Dandeli, 15.i.2015, Rajesh S. “leg”; 1♀, Meghalaya, Mawlynnong Road, 12.x.2019, David, K.J. “leg” (NIM).

(Figs This species was originally described by Hendel (1915) from Taiwan. It was recorded from India by Hancock (2012), based on photographs by the senior author. It is a medium-sized species with broad quadrate hyaline markings in cells c and r1, round hyaline spots along the wing margin and in almost all the cells. It is similar to (Bezzi) but can be differentiated by the presence of apical scutellar setae.

Hendel 31 habitus (dorsal) 32 habitus (lateral).

Male genitalia.

Epandrium (Fig. 33) dark brown, heavily sclerotised, with no demarcation from lateral surstylus (in profile view); epandrium and surstyli oval in posterior view, with well developed prensisetae (Fig. 34). Phallus elongate (1.54 mm long including glans); glans stout, with broad sclerotised rod (Fig. 35).

Figures 33–41.

Hendel 33 epandrium (lateral) 34 epandrium (posterior view) 35 glans of phallus 36 ovipositor 37 spicules on proximal end of eversible membrane 38 spicules on distal end of eversible membrane 39 spermatheca 40 aculeus 41 aculeus tip (100×).

Female genitalia.

Oviscape 0.99 mm long, dark brown, conical, dorsoventrally flattened (Fig. 36); eversible membrane 0.83 mm long, shorter than oviscape, with conical spicules along entire length except for a few rows of flattened ones distally (Figs 37, 38); taeniae short, dark brown, 0.25 of length of eversible membrane. Aculeus shorter than eversible membrane, not dorsoventrally flattened, curved at its proximal end; apex of aculeus pointed and conical (Figs 40, 41). Spermatheca brown, club-shaped, with numerous papillae (Fig. 39).

This species is widespread from India (Kerala, Karnataka, Meghalaya, Andaman and Nicobar Islands) to Japan and Australasia (Hancock 2012).

NCBI GenBank accession number – MW448367 (1♂, India: Kerala, Kannur, Aaralam, 13.i.2020, David, K. J.).

Hering

45293323-B7B5-512F-A945-8C6968C9C60D

Hering, 1939a: 179. Type locality Trichinopolis, Tamil Nadu, India.

Hering, 1939b: 146. Type locality Fujian, China.

This species was adequately described by Hering (1939a, b) and is characterised by the reduced and often indistinct hyaline markings. Indian specimens were not available for study but wings of both Hering and Hering, considered synonyms by Hancock (2012), were illustrated by Wang (1998).

This species is known in India only from the type locality in Tamil Nadu. Elsewhere, it is known from China, Taiwan, Vietnam and West Malaysia (Hancock 2012).

(Bezzi)

60D96A31-7909-5F43-A4A4-39751C22FC9D

Bezzi, 1913: 163. Type locality Calcutta [Kolkata], India.

:

:

India: 1♂, Karnataka, Madikeri, Chettalli, 05.xi.2012, David K.J. “leg”; 1♀, Assam, Dibrugarh, 07.xi.2014, Ramesh Kumar A. “leg”; 1♂, Karnataka, Kidu, CPCRI, 19.ii. 2015, David K.J. “leg”; 1♂, Karnataka, Chettalli, C.H.E.S, 03.ii.2021, David K.J. “leg”; 1♂, Assam, Barpeta, K.V.K, 09.iii.2021, Sachin K. “leg”; 1♂, Assam, Barpeta, K.V.K, 10.iii.2021, David. K.J. “leg”; 1♀, Assam, Chirang, K.V.K, 13.iii.2021, Sachin K. “leg”; 1♂, Assam, Golpara, 14.iii.2021, Rabbani M.K. “leg” (NIM).

This species was adequately described by Bezzi (1913) except for the postabdominal structures. It is almost inseparable from except for the presence of only basal scutellar setae instead of both apical and basal setae as in (Figs 42, 43).

Figures 42–43.

(Bezzi) 42 habitus (dorsal) 43 habitus (lateral).

Epandrium (Fig. 44) brown, sclerotised, with no demarcation from lateral surstylus (in profile view); epandrium and surstyli oval in posterior view, with well-developed prensisetae (Fig. 45). Phallus 1.39 mm long including glans; glans stout, with broad sclerotised rod (Fig. 46).

Figures 44–51.

(Bezzi) 44 epandrium (lateral) 45 epandrium (posterior view) 46 glans of phallus 47 ovipositor 48 spicules on eversible membrane 49 spermatheca 50 aculeus 51 aculeus tip (100×).

Oviscape 0.92 mm long, dark brown, conical, dorsoventrally flattened (Fig. 47); eversible membrane 0.72 mm long, shorter than oviscape, with conical spicules along entire length except for a few rows of flattened ones distally (Fig. 48); taeniae short, dark brown, 0.25 length of eversible membrane. Aculeus shorter than eversible membrane, 0.63 mm long, dorsoventrally flattened; apex of aculeus rounded (Figs 50, 51). Spermatheca brown, club-shaped, with numerous papillae (Fig. 49).

This species is widespread from India (Karnataka, Assam) and Sri Lanka to southern China and Australia (Hancock 2012).

Evolutionary divergence among the Indian species of

Table 1 shows the pair-wise evolutionary divergence/distance between four sequences available in NCBI database for three species namely , and . Evolutionary distance between David & Hancock, sp. nov. and is 0.0429 and between and is 0.0759 which reveals that is a distinct species in based on the available sequences. Among the species included in the analysis, evolutionary distance between and was the highest (0.0837), which is evident in the morphological differences between these two species with respect to facial markings and wing pattern.

Table 1.

Evolutionary divergence among the Indian species of .

Species name with accession numbers	P.acrostacta (MH748566)	P.platyptera (MW448367)	P.flavistigma (MT019893)	P.acrostacta (MT019891)
P.acrostacta (MH748566)
P.platyptera (MW448367)	0.0837
P.flavistigma (MT019893)	0.0429	0.0759
P.acrostacta (MT019891)	0.0036	0.0882	0.0470	0.0000

1	Apex of wing hyaline from middle of cell r2+3 to cell cua (Fig. 6); posterior wing margin not distinctly angled near apex of cell cua	P.rabbanii David & Hancock sp. nov.
–	Wing with a hyaline spot restricted to apex of cell r4+5 (e.g. Figs 16, 20, 21); posterior wing margin distinctly angled near apex of cell cua	2
2	Wing with discal spots small and often indistinct or subhyaline; cell r1 with 0–2 small hyaline indentations from costa in basal portion beyond stigma, often not crossing cell; cell cua with 3 small, isolated hyaline marginal spots and with or without additional small, isolated discal spots; holotype illustrated by Hering 1939a, fig. 14	P.tetrica Hering, 1939
–	Wing with distinct hyaline discal spots; hyaline indentations in basal portion of cell r1 with at least the basal one broad and crossing into cell r2+3; cell cua with 2–3 hyaline marginal indentations, the basal pair usually elongate but often divided medially into 2 separate spots	3
3	Wing (Figs 20, 21) with pterostigma entirely dark brown to black, 2 elongate marginal hyaline indentations in cell r1, both crossing vein R2+3 into cell r2+3, no marginal preapical hyaline spots in cell r2+3, large hyaline spots near base of cell r4+5 and near base and apex of cell dm, 1 marginal spot in cell m near apex of vein CuA, 2 indentations in cell cua and 1 or 2 spots along margin of anal lobe	4
–	Wing markings not as above (Figs 16, 31, 32, 42, 43); pterostigma usually with a subhyaline or fulvous basal patch or spot, marginal preapical hyaline spot in cell r2+3 usually present, and cell m usually with 2 or 3 hyaline marginal spots	5
4	Face black in male, yellow in female; wing (Figs 20, 21) with basal spot in cell dm not distinctly larger than apical spot and not crossing or almost crossing cell; hyaline indentations in cell cua of approximately equal length, almost crossing cell but basal spot sometimes narrowly divided medially	P.acrostacta (Wiedemann, 1824)
–	Face yellow in both sexes; wing with basal spot in cell dm distinctly larger than apical spot and crossing or almost crossing cell; basal hyaline indentation in cell cua much smaller than second indentation or broadly divided medially into 2 small spots; holotype illustrated by Hering 1941, fig. 4	P.fulvifacies Hering, 1941
5	Wing (Fig. 16) with base and pterostigma largely fulvous to pale brown, contrasting with rest of wing; cell m without an isolated anterobasal hyaline spot	P.flavistigma David & Hancock, sp. nov.
–	Wing (Figs 31, 32, 42, 43) with pattern variable but uniformly dark brown with hyaline spots and markings; cell m with an isolated anterobasal hyaline spot	6
6	Scutellum with apical and basal setae	P.platyptera Hendel, 1915
–	Scutellum with only basal setae	P.zodiacalis (Bezzi, 1913)

1	Wing broad and almost circular beyond basal third, apex evenly rounded and entirely dark, without hyaline discal or marginal spots or indentations except for pair of small costal spots at bases of pterostigma and cell r1 adjacent to apices of veins Sc and R1, respectively; illustrated by Hardy 1974, fig. 129 [Philippines (Luzon)]	P.bezzii Hardy, 1974
–	Wing often broad but distinctly longer than wide, apex at least slightly produced and with at least a hyaline apical spot in cell r4+5; usually with hyaline discal and marginal spots or indentations	2
2	Wing (Fig. 6) with hyaline apical band distinctly crossing veins R4+5 and M1 into cells r2+3 and m1	3
–	Wing (Figs 10, 16, 20, 21, 32, 43) with oval or quadrate hyaline apical spot confined to cell r4+5	7
3	Male wing without hyaline spots or indentations apart from small marginal indentation in cell r1 at apex of vein R1 and crescentic hyaline apex; female wing with crescentic hyaline apex plus hyaline marginal spots and indentations and subbasal hyaline spot in cell dm but no spot near base of cell r4+5; illustrated by Wang 1998, fig. 253–4 [China (Yunnan)]	P.nigripennis Wang, 1998
–	Male wing (where known) with one broad or 2 narrow marginal hyaline indentations in cell r1 near apex of vein R1 and often a spot near base of cell r4+5; female wing (where known) with hyaline medial spot close to line of crossvein r-m and often a spot near base of cell r4+5	4
4	Wing of both sexes without hyaline spot near base of cell r4+5 and hyaline apex in cell m separate from the 2 hyaline marginal spots; hyaline marginal indentation in cell r1 broad and rectangular in male, divided into elongate indentation and 2 small round spots in female; male illustrated by Hardy and Drew 1996, fig. 167 [Australia (Queensland)]	P.parvipuncta Malloch, 1939
–	Wing of both sexes (where known) with hyaline spot near base of cell r4+5; apex with hyaline band in cell m enclosing one or both marginal spots, leaving no more than a single separate marginal spot; hyaline indentation from costa in cell r1 not as above	5
5	Male unknown; female posterior to vein R4+5 with spots in cells r4+5 (1, at base), bm (1, near apex), dm (1, near middle), m (1 marginal near apex of vein CuA), cua (3, 2 basal and 1 near middle) and anal lobe (1); not illustrated [Taiwan]	P.apicalis Hendel, 1915
–	Female unknown; male with 1 or 2 hyaline indentations in cell r1 near apex of vein R1 and 1 or no hyaline marginal spots in cell m near apex of vein CuA separate from the apical hyaline area, large hyaline spots at base of cell r1 and at basal third of cell dm, and 2 elongate marginal indentations in cell cua [India and Burma]	6
6	Wing in cell r1 with 2 hyaline indentations near apex of vein R1; cell m with one hyaline marginal spot in addition to apical hyaline area; cell cua with hyaline marginal indentations reaching or almost reaching vein CuA; illustrated by Hering 1938, fig. 50 [NE Burma]	P.alboapicalis Hering, 1938
–	Wing (Fig. 6) with cell r1 with 1 hyaline indentation near apex of vein R1 and cell m without a hyaline marginal spot in addition to apical hyaline area; cell cua with hyaline marginal indentations ending at or before middle of cell	P.rabbanii David & Hancock, sp. nov.
7	Wing with cell c and basal two-thirds of pterostigma hyaline; hyaline marginal spots (including 2 in cell m1 and 3 in cell cua) present but pale discal spots absent; head with 1–2 pairs of frontal setae; illustrated by Hardy 1974, fig. 127 [Philippines (Luzon)]	P.amita Hardy, 1974
–	Wing with cell c not entirely hyaline and pterostigma with at most a hyaline basal spot; both hyaline marginal and pale discal spots usually present; head with 3 pairs of frontal setae	8
8	One pair of scutellar setae, apicals absent; illustrated by Bezzi 1913, fig. 65, Hardy 1973, pl. v, fig. 45, Hardy 1974, fig. 130, Wang 1998, fig. 256 and this paper, figs 42–43 [India and Sri Lanka to Philippines, Indonesia and northern Australia]	P.zodiacalis (Bezzi, 1913)
–	Two pairs of scutellar setae, apicals present	9
9	Wing cell r2+3 with 2 narrow marginal hyaline spots or indentations; marginal hyaline indentations in cell r1 close to apex of vein R1 normally reduced in cell r2+3 to isolated spots in yellowish field; illustrated by Hancock 2012, figs 3–4 [Indonesia to NE Australia and Solomon Islands]	P.ampla de Meijere, 1914
–	Wing cell r2+3 with 0–1 narrow marginal hyaline spots or indentations; marginal hyaline indentations in cell r1 close to apex of vein R1 present or reduced in cell r2+3 but not reduced to isolated spots in a yellowish field	10
10	Wing with single large marginal hyaline indentation in cell r1 close to apex of vein R1 that extends across cell r2+3 and is aligned with posterior indentation in cell cua that crosses cell; cell m with single hyaline marginal spot placed close to apex of vein CuA; illustrated by Hardy 1974, fig. 128 [Philippines (Luzon)]	P.aptata Hardy, 1974
–	Wing with 2 marginal hyaline indentations in cell r1 close to apex of vein R1, the distal one sometimes reduced and largely united with basal one, sometimes both reduced to small marginal spots or single indistinct pale area; cells m and cua not both with a single hyaline marginal spot or band	11
11	Wing cell m with 3 hyaline marginal spots or indentations	12
–	Wing cell m with at most 2 hyaline marginal spots or indentations	13
12	Wing cell dm with 3 rounded hyaline spots; cell m with 2 hyaline spots in anterobasal quarter; illustrated by Hardy and Drew 1996, fig. 168 [Australia (Queensland)]	P.trimaculata Hardy & Drew, 1996
–	Wing cell dm with 2 rounded hyaline spots; cell m with 1 hyaline spot in anterobasal quarter; illustrated by Wang 1998, fig. 251 [Japan (Ryukyu Is), Taiwan and China (Sichuan, Guangxi)]	P.assimilis (Shiraki, 1968)
13	Wing cell r4+5 with very small apical hyaline spot; illustrated by Hardy 1988, fig. 22 [Indonesia (Java, Sumba)]	P.sumbana Enderlein, 1911
–	Wing cell r4+5 with large apical hyaline spot filling all or most of cell apex	14
14	Wing cell r1 with 2 large and quadrate hyaline indentations, distal one broader than wide, plus subapical posterior spot not reaching costa; cell m with 2 elongate hyaline indentations, the anterior one much smaller and narrower than the posterior one; cell cua with 3 broad hyaline indentations, the basal pair crossing or almost crossing cell, plus basally with extension of large hyaline mark in anal lobe; male with distinct white or silvery parafacial stripes (females unknown)	15
–	Wing markings not as above, distal hyaline indentation in cell r1 narrower than wide and/or one or both marks often reduced or absent, subapical spot in cell r1, when present, small and marginal at costa or crossing cell, and marginal marks in cell m usually subequal in size; male without white or silvery parafacial stripes	16
15	Wing (Figs 9, 10, 16) with dark markings anterior to vein R4+5 pale brown to yellow-brown in basal two-thirds and dark brown elsewhere; pterostigma with basal spot yellow; hyaline spot at base of cell r4+5 circular; head with parafacial stripes white [southern India (Karnataka, Tamil Nadu)]	P.flavistigma David & Hancock, sp. nov.
–	Wing with dark markings uniformly dark brown; pterostigma with basal spot hyaline; hyaline spot at base of cell r4+5 oval; head with parafacial stripes silvery; illustrated by Hardy 1973, fig. 149 [Thailand, Cambodia and Vietnam]	P.quadrula Hardy, 1973
16	Wing cell r2+3 without small and narrow marginal spot posterior to apex of vein R2+3 and pterostigma without hyaline basal spot; cell m with 1 small semicircular marginal hyaline spot near apex of vein CuA	17
–	Wing cell r2+3 with small and narrow marginal spot below apex of vein R2+3 and pterostigma often with hyaline basal spot; cell m usually with 2 small and often elongate marginal hyaline spots	18
17	Face largely black in male, yellow in female; basal spot in cell dm not distinctly larger than apical spot and not crossing or almost crossing cell; hyaline indentations in cell cua of approximately equal length, almost crossing cell but basal spot sometimes narrowly divided medially; illustrated by Bezzi 1913, fig. 57, Hancock 2012, fig. 1 and this paper, figs 20–21 [India and Sri Lanka to southern China (Yunnan) and Cambodia]	P.acrostacta (Wiedemann, 1824)
–	Face yellow in both sexes; basal spot in cell dm distinctly larger than apical spot and crossing or almost crossing cell; basal hyaline indentation in cell cua much smaller than second indentation or broadly divided medially into 2 small spots; illustrated by Hering 1941, fig. 4 [India (Maharashtra, Rajasthan)]	P.fulvifacies Hering, 1941
18	Wing with discal spots often subhyaline or indistinct and pale brown; cell r1 often with 0–2 small hyaline indentations or spots in basal portion (especially in males) or with 2 indentations often largely fused; cell cua with 2 or 3 small and isolated hyaline marginal spots and with or without additional small and isolated discal spot	19
–	Wing with discal spots normally hyaline and distinct; cell r1 with second hyaline indentation distinct and at most weakly joined to first; cell cua with 2–3 hyaline marginal indentations not all small and isolated, basal pair often almost crossing cell or medially divided into separate spots	20
19	Wing cell r1 with basal hyaline indentations often reduced to 0–2 small hyaline spots (especially in males, better developed in females); cell cua with 3 small and isolated hyaline marginal spots and with or without additional small and isolated discal spots; anal lobe with 2 distinct hyaline marginal spots; illustrated by Hardy 1973, pl. v, fig. 44 and Wang 1998, figs 252 & 255 [India (Tamil Nadu), Thailand and West Malaysia to China (Guangxi, Fujian) and Taiwan]	P.tetrica Hering, 1939
–	Wing cell r1 with second hyaline indentation narrow and strap-like or fused with first indentation leaving only small dark costal spot between them; cell cua with 2 small marginal spots; anal lobe with hyaline marginal spots vestigial or absent; illustrated by Bezzi 1913, fig. 64 and Hardy 1973, pl. v, fig. 42 [southern Burma to Vietnam and Indonesia (Sumba)]	P.euryptera (Bezzi, 1913)
20	Wing cell m with a small anterobasal hyaline spot and no marginal spots; cell cua with 2 undivided indentations almost crossing cell; cell r4+5 with basal spot large and ovate, much larger than the 2 distinct spots in cell dm; illustrated by Hardy 1973, pl. v, fig. 43 [Thailand, Cambodia and Vietnam]	P.intacta Hardy, 1973
–	Wing cell m with 2 small hyaline marginal spots; cell cua with 2 or 3 indentations, with at least the more distal of the 2 basal indentations divided medially and apical spot small or absent; cell r4+5 with basal spot small and circular, not much larger than the 2 distinct spots in cell dm	21
21	Wing evenly rounded posteriorly, not distinctly angled basal of end of vein CuA and with numerous distinct discal spots, 1–2 in cell br, 3 in cell r4+5, 1–2 in cell r4+5, 2 in cell dm and 1 anterobasally in cell m; posterior marginal spot in cell m distinctly larger and broader than anterior marginal spot; illustrated by Wang 1998, fig. 259–260 [Japan (Ryukyu Is), China (Sichuan) and possibly Indonesia (West Papua)]	P.shirouzui (Ito, 1984)
–	Wing distinctly angled posteriorly, broadest just basal of end of vein CuA and with only 3 distinct discal spots, 1 at base of cell r4+5 and 2 in cell dm; posterior marginal spot in cell m not distinctly larger and broader than anterior marginal spot	22
22	Apical scutellar setae distinct, about half length of basals; anal lobe of wing with hyaline marginal spots vestigial or absent; posterior hyaline marginal spot in cell m narrow, elongate and perpendicular; illustrated by Hancock 2012, fig. 2 [southern Thailand to Indonesia (Java, Sulawesi)]	P.amplipennis (Walker, 1860)
–	Apical scutellar setae weak, about quarter length of basals; anal lobe of wing with hyaline marginal spots round and distinct; posterior hyaline marginal spot in cell m often short and broad; illustrated by Hancock 2012, figs 5–6 and this paper, figs 31–32 [India to Japan (Ryukyu Is), Australia, Solomon Islands and Vanuatu]	P.platyptera Hendel, 1915