﻿Four new species of Zeugodacus Hendel (Diptera, Tephritidae, Dacinae, Dacini) and new records of dacines from India

﻿Abstract Four new species of Zeugodacus Hendel are described from India viz., Zeugodacusmomordicae David & Ajaykumara, sp. nov. from Arunachal Pradesh infesting male flower buds of Momordicadioica, Zeugodacusnasivittatus David & Abhishek, sp. nov. from Meghalaya, Zeugodacus (Sinodacus) sinuvittatus David & Abhishek, sp. nov. from Himachal Pradesh and Zeugodacus (Zeugodacus) umiam David & Kennedy, sp. nov. from Meghalaya. An illustrated key to all species of Zeugodacus from India is also included. Bactrocera (Parazeugodacus) abbreviata (Hardy) and Dacus (Mellesis) vijaysegarani Drew & Hancock are recorded for the first time from India.


Introduction
Zeugodacus Hendel is a genus in tribe Dacini with 196 species recorded from the world (Doorenweerd et al. 2018) and thirty described species from India (David et al. 2017;David and Ramani 2019).They are characterised by the shallow emargination of sternite 5 in males, posterior lobe of surstylus 5-6× longer than anterior lobe, glans of phallus with patterned acrophallus.Fruit flies of genus Zeugodacus Hendel are economically important as several of them are pests of various horticultural crops.Zeugodacus was originally treated as a subgenus of Bactrocera Macquart, it was elevated to genus level by Virgilio et al. (2015) based on molecular markers which confirmed the findings of Krosch et al. (2012).It was further supported by works by San Jose et al. (2018), Dupuis et al. (2018) and Zhang et al. (2022).Hancock and Drew (2018) consider Zeugodacus as a subgenus of Bactrocera.David et al. (2017) described Bactrocera brevipunctata David and Hancock from Maharashtra which was later transferred to genus Zeugodacus by Doorenweerd et al. (2018).David and Ramani (2019) studied the postabdominal structures of 16 species of Zeugodacus from India and performed a morphology based phylogenetic analysis of tribe Dacini wherein Bactrocera and Dacus Fabricius were monophyletic and Zeugodacus was polyphyletic, which might be due to the reason that only Indian species were included in the phylogenetic analysis.In this paper, four new species of Zeugodacus are described with illustrations of postabdominal structures.Two species of dacines, Bactrocera (Parazeugodacus) abbreviata (Hardy) and Dacus (Mellesis) vijaysegarani Drew & Hancock are recorded for the first time from India.An illustrated key to 34 species of Zeugodacus from India is also included.

Materials and methods
Specimens deposited in the following museums have been studied: Natural History Museum, London, United Kingdom (NHM) and National Insect Museum, ICAR-National Bureau of Agricultural Insect Resources, Bengaluru, India (NIM).
Images of specimens, epandrium, and ovipositor were taken using a Leica DFC 420 camera mounted on a Leica M205A stereo zoom microscope; images of glans of phallus, aculeus tip and spicules on eversible membrane were taken using an 8 MP camera temporarily attached to a Leica DM 1000 compound research microscope, Olympus DP 23 attached to BX51 and Olympus SC 50 attached to BX 43; 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) except for wing terminology which follows Cumming and Wood (2017).
One hind leg was removed from one specimen of Z. momordicae 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) (Hebert et al. 2003).The sequence was annotated using NCBI Blast tools and submitted to the NCBI GenBank Database where an accession number was obtained (Z.momordicae-OQ353070).
Diagnosis.Abdominal tergites free, scutum with medial postsutural vitta except for few species in several subgenera including Parasinodacus Drew & Romig, Paradacus Perkins and some species of Sinodacus Zia, sternite 5 of male with shallow/flat posterior emargination.In males, epandrium distinctly bulb-shaped in posterior view, proctiger hyaline, triangular (when uninflated) smaller than epandrium, lateral surstylus longer than epandrium (profile view); posterior lobe of lateral surstylus 5-6× longer than anterior lobe.Phallus with well-developed acrophallus (single semi-tubular lobe) and patterned/granulated praeputium.Dorsal sclerite of glans without hexagonal pattern.Aculeus dorsoventrally flattened with four pairs of preapical setae (David and Ramani 2019).Zeugodacus is similar to Bactrocera and Dacus in general appearance as they are wasp mimics and are characterised by the presence of reddish-brown to black colour with yellow vittae and markings.It can be differentiated from Dacus by the presence of free abdominal tergites and by the presence of four pairs of preapical setae; from Bactrocera by the shallow/ flat emargination sternite 5 in males, posterior lobe of lateral surstylus 5-6× longer than anterior lobe and patterned acrophallus.Medial postsutural vitta narrowed at both ends (Fig. 18), pecten of cilia absent in male (Fig. 28), face fulvous in male (Fig. 2), with a transverse band in female (Fig. 3    Diagnosis.Zeugodacus momordicae resembles Z. incisus in possessing black scutum, two transverse bands on face, continuous costal band and extensive femoral markings, but can be differentiated by the absence of yellow spot anterior to lateral vittae along transverse suture and presence of prescutellar acrostichal setae.It can be differentiated from Bactrocera ablepharus (Bezzi) by the presence of prescutellar acrostichal setae and face with two transverse bands.It can be differentiated from Z. cilifer 69) by the aculeus shape and spicules on distal end of eversible membrane as discussed below.Aculeus tip is elongate, parallel sided and not tapering abruptly beyond the preapical conical flange (width of the conical projection-0.06mm) and length of aculeus after the preapical flange is 0.21 mm in Z. cilifer (Figs 66,67), whereas in Z. momordicae, aculeus is tapering abruptly beyond the preapical conical flange (width of the conical projection -0.08 mm) (Figs 56, 57) and length of aculeus after the preapical flange is 0.15-0.18mm.Spicules on Z. cilifer are conical with single projection with a shorter base (Fig. 68), whereas Z. momordicae (Fig. 54) possess broader conical spicules.

Key to species of Zeugodacus Hendel from India
Description.Female.Medium sized species (wing length 4.37-5.45mm), face with two broad black bands.Scutum black with yellow lateral postsutural vitta ending beyond intra-alar seta, anepisternal stripe broad reaching anterior notopleural seta dorsally, continued as a small transverse marking on katepisternum.Wing hyaline with costal band continuous from cell sc to the apex of the wing and confluent with vein R 2+3 , anal streak well developed.Abdomen predominantly black with a narrow transverse fulvous band on tergites 1 and 2 (in few specimens all tergites black).Females with two spermatheca, aculeus pointed with preapical projection.
Male.Similar to female except for face (Fig. 40) which is nearly black in few males with a narrow longitudinal fulvous line separating the bands, costal band discontinuous in few male specimens examined, sternite 5 in males black with shallow concavity, pecten present on tergite 3.
Etymology.The species name is derived from Latin words nasi vitta which means nose-shaped vitta.
Diagnosis.Zeugodacus sinuvittatus is similar to Z. hochii (Zia), Z. infestus (Enderlein) and Z. brevipunctatus David & Hancock in possessing reddish brown scutum, club shaped abdomen and wing with broad apical black spot.It can be differentiated from Z. hochii by the absence of medial postsutural vitta, face with separate black spots unlike transverse band, presence of discontinuous costal band slightly overlapping vein R 2+3 ; from Z. infestus and Z. brevipunctatus by the absence of lateral and medial postsutural vitta, absence of postsutural supra-alar seta, narrow costal band interrupted in cell r 1 and by the broad apical spot.Unlike Z. brevipunctatus, acrophallus of Z. sinuvittatus is fully patterned.
Description.Male.Large sized species (wing length 7.05 mm); face fulvous with two elongate black markings in the antennal furrow and a medial longitudinal line; scutum reddish brown in ground colour without lateral and medial vitta, with broad quadrate black patches in presutural and postsutural areas, notopleuron and postpronotal lobe yellow, inconspicuous yellow spot anterior to notopleural suture; anepisternal stripe reaching midway between anterior notopleural seta and notopleuron; scutellum yellow; wing predominantly hyaline with costal band slightly overlapping vein R 2+3 , discontinuous towards apex of cell r 1 , with a broad apical spot covering the apex of cell r 2+3 , r 4+5 and upper one-fourth of cell m, anal streak narrow, no dense aggregation of microtrichia around A 1 +Cu 2 ; abdomen club shaped, tergite 2 with a prominent black semicircular spot, tergites 3-5 with dark fuscous lateral markings and a narrow medial longitudinal band.
Head (Fig. 85).Height 1.60 mm.Frons length 1.85× breadth; fulvous with fuscous marking on anteriomedial hump and around bases of frontal and orbital setae, all setae black: two pairs of frontal setae and one pair of orbital setae; lunule black.Ocellar triangle black, vertex yellow.Face fulvous with two separate elongate black markings in antennal furrows and a medial longitudinal black line.Scape (0.23 mm long) and pedicel (0.22 mm long) fulvous, first flagellomere (0.74 mm long) dark fuscous on outer side and apex, arista non plumose, combined length of pedicel and flagellomere longer than the vertical length of face.Gena fulvous with a black marking and a seta.Occiput fulvous; lateral and medial vertical setae present, occipital row with three or four stout black setae.Thorax (Figs 86,87).2.18 mm long, 2.03 mm wide; scutum red brown with two black quadrate markings one each in presutural and postsutural area.Pleura red-brown in ground colour with black markings anterior to anepisternal stripe, katepisternum and anepimeron.Yellow markings as follows: postpronotal lobe, notopleuron, anepisternal stripe reaching midway between notopleuron and anterior notopleural seta and continuing to katepisternum as a transverse spot; anatergite (posterior apex black); anterior 3/5 of katatergite (remainder black).Scutellum yellow without narrow black basal band, subscutellum red-brown with black lateral margins.Chaetotaxy: scutellar seta, 1; intra-alar seta, 1; postalar seta, 1; anepisternal seta, 1; anterior notopleural seta, 1; posterior notopleural seta, 1; scapular setae, 1. Leg (Fig. 87): Coxa, trochanter dark fuscous, all femora with extensive fuscous markings; fore femur wholly fuscous, 0.80 of mid femur and 0.60 of hind femur fuscous; fore and hind tibiae fuscous, mid tibia fulvous, tarsal segments slight fuscous.Wing (Fig. 89).Length, 7.05 mm, cells bc and c hyaline; microtrichia in outer corner of cell c only; remainder of wing hyaline except dark fuscous cell sc, costal band overlapping vein R 2+3 , interrupted towards apical one-fourth of cell r 1 and with a broad apical spot covering apex of cell r 2+3 , r 4+5 and anterior one fourth of cell m, extension of cell cua longer than cell cua, base of cell br with microtrichia, anal streak narrow, confined to cell cua, lacks dense aggregation of microtrichia around A 1 +Cu 2 ; supernumerary lobe weak.Abdomen (Fig. 88).3.69 mm long, 1.92 mm wide, club shaped, tergites free, tergites 1 fulvous, tergite 2 reddish brown with a black semicircular marking, tergite 3 with broad, black basal band and pecten, tergites 4 and 5 with dark lateral margins and a narrow medial longitudinal band.Tergite 5 without prominent shining spots (ceromata).
Etymology.The species name is derived from Latin words sine (= without) and vitta (= band), as the species lacks lateral and medial poststurural vitta on scutum.
Host plant.Not known, collected by sweep netting on grapevine Male parapheromone.Not known.
Remarks.This species is placed in Zeugodacus due to shallow/flat posterior emargination of sternite 5 in males, posterior lobe of lateral surstylus much longer than anterior lobe and patterned acrophallus.It is placed in subgenus Sinodacus as it lacks prescutellar acrostichal seta, basal scutellar seta and due the club-shaped abdomen.Diagnosis.Zeugodacus umiam is similar to Z. nigrifacies (Shiraki) in possessing black face, fore femur entirely black, scutellum with broad black basal band and an apical spot but can be differentiated by the absence of subapical band, band on crossvein r-m and two scutellar setae.It can be distinguished from Z. menglanus (Yu, Liu & Yang) by the facial markings (wholly black in male; dorsal half black in female), two scutellar setae and lack of apical expansion in costal band.It is similar to Z. diaphorus in possessing apical scutellar spot, two pairs of scutellar setae and black face, but can be separated by the presence of broad black basal band on scutellum, narrow anepisternal stripe not reaching anterior notopleural seta dorsally.

Zeugodacus
Description.Female.Medium sized, black species (wing length 5.65 mm); face posterior half black; scutum black with narrow yellow lateral postsutural vitta and medial vitta, lateral vitta ending before postalar seta, notopleuron and postpronotal lobe yellow, small yellow spot anterior to notopleural suture, anepisternal stripe not reaching anterior notopleural seta dorsally, scutellum yellow with a broad black basal band, with an apical black spot; wing predominantly hyaline with costal band confluent with vein R 2+3 , expanded slightly towards apex of cell r 2+3 and r 4+5 , anal streak prominent; abdomen oval, all tergites black except tergite 2 with a broad fulvous band posteriorly, narrow fulvous bands in tergites 3-5.
Male (Figs 102-106).Similar to female except for entirely black face, broad basal band on the scutellum, dense aggregation of microtrichia around A 1 +Cu 2 , black extensive markings on all femora and pecten on tergite 3.
Etymology.The species name is type locality of the species and is a noun in apposition.
Remarks.This species is placed in Zeugodacus due to shallow/flat posterior emargination of sternite 5 in males, posterior lobe of lateral surstylus much longer than anterior lobe and patterned acrophallus.It is placed in subgenus Zeugodacus as it possesses medial postsutural vitta, postsutural supra-alar and prescutellar acrostichal seta.out costal band, short yellow lateral vitta ending at postsutural supra-alar seta, but can be differentiated by the presence of separate black spots on face, all femora fulvous without fuscous/black markings.Male of the species has been examined for genitalia characters.Epandrium quadrate with proctiger smaller than epandrium; posterior lobe of surstylus as long as anterior lobe, epandrium (oval) in posterior view.Glans of phallus with acrophallus patterned; phallus 1.82 mm.This species was originally described from Philippines, distributed across China and Thailand, is being recorded from Meghalaya, India for the first time.
Male attractant.Zingerone.Diagnosis.This species has been adequately described by Drew et al. (1998) and Drew and Romig (2013) except for postabdominal structures.A male of the fly collected from Meghalaya was dissected to study the postabdominal structures.Epandrium deeply sclerotised, black, lateral surstylus with posterior lobe slightly longer (2-3×) than anterior lobe; epandrium bulbous (in posterior view), glans of phallus elongate (0.7 mm) with patterned aculeus.