A taxonomic study of six species of the genus Junonia Hübner , [ 1819 ] ( Insecta : Lepidoptera : Nymphalidae ) from the northwestern Himalayan region in India

For Focus, Scope, Aims, Policies and Guidelines visit http://threatenedtaxa.org/index.php/JoTT/about/editorialPolicies#custom-0 For Article Submission Guidelines visit http://threatenedtaxa.org/index.php/JoTT/about/submissions#onlineSubmissions For Policies against Scientific Misconduct visit http://threatenedtaxa.org/index.php/JoTT/about/editorialPolicies#custom-2 For reprints contact <info@threatenedtaxa.org> Short Communication A taxonomic study of six species of the genus Junonia Hübner, [1819] (Insecta: Lepidoptera: Nymphalidae) from the northwestern Himalayan region in India

There is considerable geographic variation within Junonia species, and possible on-going inter-specific hybridization suggests that Junonia may be a ring species, making this a very difficult group to define taxonomically (Borchers & Marcus 2014).Numerous studies at the molecular level have been initiated but no emphasis has been laid down on the genitalic attributes of the species of this genus (Vanlalruati et al. 2011;Gemmell et al. 2014;Win et al. 2015).Recent taxonomic review of the tribe Junoniini from Myanmar given by Win et al. (2016) included only adult descriptions, whereas important taxonomic characters of the external genitalia and wing venations have not been discussed.
In many insects, genitalia often provide the only way to reliably distinguish species using morphology (Özgül-Siemund & Ahrens 2015).Hence in order to fill the taxonomic gaps, the male and female external genitalia and wing venation of the six species of the genus Junonia have been illustrated, described and discussed in detail.

Materials and Methods
The material examined for the present study has been obtained from the preserved entomological collections lying in the Department of Zoology and Environmental Studies, Punjabi University, Patiala and Zoological Survey of India, HARC, Solan.To study wing venation, the methodology given by Common (1970) is followed.For the study of external male and female genitalia, the methodology given by Robinson (1976) is followed.
Terminological review: The terminology given by Sibatani et al. (1954) and Klots (1956) for the male and female genitalia have been followed.
Male genitalia: Various authors like Bethune-Baker (1891), Petersen (1904), Pierce (1909), Eyer (1924), Sibatani et al. (1954), and Ogata (1957) had presented different terminological reviews on the male genitalia in Lepidoptera.The term 'tegumen' is used to denote the dorsal part of the ninth abdominal segment.Pierce (1909) established the term vinculum for the ninth abdominal sternite, but later on Pierce (1914) synonymised it with the term 'saccus' (Bethune-Baker 1891) which signified both saccus and vinculum.In the present study, the terms used are as follows: saccus, lateral-dorsal projections of the saccus, and lateralventral projections of the tegumen.
Nevertheless, the subdivision of valvae has always been the point of debate.The valvae terminology reviewed by Sibatani et al. (1954) and Klots (1956) are applied in the present work.The terms used are as follows: costa (dorso-proximal region of the valvae), sacculus (ventro-proximal region of the valvae), ampulla (medio-dorsal region), and harpe (posterior projection of the sacculus, on the innerside of the valvae).The terminology for the copulatory organ of male Lepidoptera is followed as given by Klots (1956).The terms used are as follows: aedeagus (sclerotized tubular structure), vesica (membranous tube), ductus ejaculatorius (a duct along which spermatozoa move from the testes).
Female genitalia: All the terms adopted by Klots (1956) are used, except the term genital plate used by Pierce (1909) is here replaced by sterigma as the latter denotes the fused ante-and post-vaginal lamellae.Moreover, it is more commonly found mentioned in the literature and widely used by specialists (Carneiro et al. 2013).

Genus Junonia Hübner, [1819]
Common name: Buckeyes, Pansies or Commodores Junonia Hübner, [1819] Verz.bek.Schmett.(3): 34 General Characteristics: Adult Diagnosis: Broad-winged butterflies with a relatively small body, ground-colour dull grey-brown, red brown or black; head broad; eyes naked; palpi long, projecting beak-like; antennae about half the length of the forewing; latter triangular, costal margin strongly curved, apex more or less obliquely truncate, termen concave; wing shape variable in seasonal forms, wing more strongly angulate in dry-season form than in wetseason form; upper discocellular obsolete, middle one deeply concave, lower discocellular absent, discal cell open; hind wing triangular, apex round, tornal angle often produced into a lobe; precostal erect and then almost rectangular bent outward, sometimes bearing a short spur on basal side.
Immature stages: Larva cylindrical, with branched thorns; head with short bristles; pupa suspended, with small wart-like tubercles on the back.

Distribution and Natural History
The genus is distributed over all major biogeographical regions of the world except the Palaearctic (Kodandaramaiah & Wahlberg 2007).
Commonly known as the pansies and buckeyes, the members of the genus Junonia are sun-loving butterflies and can be found flying during the hottest hours of the day.Although having a preference for the riverine habitat, these butterflies are generalist species and are found venturing into wide variety of habitats.The wings have characteristic eye spots (ocelli).Their flight is very fast, remaining close to the ground.The dull colouration on the underside of these butterflies makes them nearly invisible while they sit on bare ground to indulge in mud puddling under the sun.They have a definite preference for salts from the ground and are often seen puddling in the rocky terrains near the source of water.These butterflies are fast fliers, and are fond of sitting with spread wings on the ground.
Identification keys to species of the genus Junonia Hübner, [1819]  Diagnosis: Adult (male): Upper side of forewing dark brown from base to post discal area, an oblique white band from costa to termen, apical area pale brown, two eye spots in post discal area; upper side of hind wing with ground colour shining blue, black angular patch below discal cell curving towards inner margin, two eyespots in post discal area; underside from dull ocheraceous to dark brown depending upon to seasonal form.
Female: Similar as male, but basal half of hind wing entirely black, and eye spots more prominent.
Male genitalia (Fig. 2): Tegumen well developed, arched, moderately sclerotized; latero-ventral projections of tegumen broad; saccus distinct and moderately sclerotized; latero-dorsal projections V-shaped, narrow, extending dorsally; anterior projection of saccus tubular in lateral view and posterior elongation absent; uncus differentiated from tegumen by a light sclerotization, broad proximally with distal portion descending into a sharp narrow pointed tip, sinous and sparsely setose and approximately as long as tegument; gnathos attached to caudal margin of tegumen, more or less rhomboid in shape and moderately sclerotized with two halves remain separate from each other; subscaphium developed into a u-shaped pad, densely setosed; valvae broad proximally, well sclerotized, costa convex, broad and strongly sclerotized, ampulla well developed into an arm, sparsely setose, ventral margins serrated and slightly folded upon it, sacculus convex, sparsely setose and well sclerotized, harpe developed into an arm, well sclerotized, bears serrations, extends slightly beyond ampulla and partially overlaps latter laterally; aedeagus curved, approximately as long as valvae, more or less cylindrical and narrows into a tip posteriorly; ductus ejaculatorius enters from lateral side.
Female genitalia (Fig. 3): Sterigma composed of coupling of lamella antevaginalis and lamella postvaginalis (no sharp distinction between these two) which after fusion forms a sclerotized rectangular plate forming a u-shaped arc; ductus bursae very small, funnel like, membranous, leading to corpus bursae after a narrow constriction; corpus bursae very long, extends far cephalad into abdomen, membranous, balloon shaped, narrow at its base but dilating gradually towards apex, signa absent; ductus seminalis enters dorsally at base of ductus bursae and its entry marked Range country: Madagascar, tropical Africa (dry), Arabia, India, Sri Lanka, Myanmar, New Guinea, northern Australia.
Remarks: This is a widely spread species.The wet season forms have prominent eyespots while the dry season forms have these eyespots reduced in size.No  Diagnosis: Adult (male): Upper side of forewing dull brown, a broad brown colour band traversed from costa to inner margin, two apical white spots present near costal margin; hind wing upper side dull brown, broad brown colour band in continuation from forewing, sub-marginal series of eyespots along terminal margin; under side of forewing dark brown, basal area traversed by three greyish fasciae, a discal prominent dark brown line traversed across; underside of hindwing eye spots not as prominent as above.
Adult (female): Similar as male.
Male genitalia (Fig. 5): Tegumen U-shaped from dorsal, moderately sclerotized, more or less obliquely slanting towards uncus.Saccus well developed and distinct; anterior projection of saccus tubular, reaches cephalad up to mid of sixth sternite, slightly curved upward; laterodorsal projections of saccus narrow, prominent, bears a strong convex curvature and produced into an anteriorly directed crest at point where valvae articulate with latter; uncus stout, well sclerotized, sparsely setosed, angulate from mid dorsal portion, posteriorly descending into sharp pointed tip directed ventral; gnathos paired, roughly rectangular, glued to tegumen by a membrane; valvae well developed, sclerotized, broad proximally, costa well sclerotized, more or less triangular in shape, ampulla well developed into a laterally flattened arm, sparsely setose and margins spinous, sacculus also well developed, harpe sparsely setose, slightly sclerotized, produced into a spatulate arm with serrated margins and overlaps ampulla laterally; aedeagus long, slender, uniform width throughout its length following apex descending into a sharp pointed tip.
Female genitalia (Fig. 6): Eighth sternite reduced and well sclerotized; antevaginal lamella and postvaginal lamella collapse, form a sclerotized plate giving rise to tubular, dorso-ventrally compressed, well sclerotized sterigma; ostium bursae cresent shaped; ductus bursae long and membranous; ductus seminalis enters at base of ductus bursae (region marked with prominent sclerotization) from latero-dorsal side; corpus bursae large, membranous, balloon shaped, narrow at its base but dilating gradually towards apex, signa absent; a pair of well sclerotized and thin anterior apophysis arises from anterolateral margin of 8 th tegument.8 th tegument and sternite not fused; anal papillae not small, lightly sclerotized, densely setose with fine setae, pair of posterior apophysis present, well sclerotized, slightly longer than anterior apophyses; both pairs of apophyses are oriented horizontally along abdomen.
Range country: Sri Lanka, India, Myanmar, Sumatra, China Discussion: This species is quite common throughout its range.The males exhibit roosting behaviour.No major variation in the external genitalia could be found.Diagnosis: Adult (male): Upper side of forewing brownish, two pale ochreous bars with black margins traverse discal cell, a prominent blue centred orange ringed eyespot near tornus encircled by pale ochre spots which extend obliquely towards costal margin, two more ochre spots near apex, termen sinous; upper side of hind wing brownish with two blue centred orange eye spots, upper eyespot prominent, second eyespot indistinct, two dark sinous sub-marginal lines, termen sinous; underside very variable according to seasons, series of eye spots on hind wing well developed in wet season forms but very obscure in dry season forms.
Male genitalia (Fig. 8): Tegumen strongly built, broad, moderately sclerotized, obliquely slanting towards uncus; uncus broad distally, strongly sclerotized, as long as tegumen, sickle shape with pointed tip, deeply grooved in centre at point of origin of uncus giving a y-shape from dorsal view; gnathos lightly sclerotized, more or less triangular, separated from tegument by a membrane, pair remains separate; saccus well developed, anterior projection tubular, directed upward, tip blunt; latero-dorsal projections of saccus narrow, arced and short; valvae well developed, sclerotized, longer than broad, directed in a diagonal fashion and hides pointed tip of uncus, costa well developed and sclerotized, ampulla developed into a small arm bearing bifid claw, curved ventrally, saccullus well sclerotized, harpe simple and sparsely setose with long setae, bears minute spines; aedeagus not long, slender, descending into a pointed tip posteriorly.Female genitalia (Fig. 9): Eighth sternite reduced and strongly sclerotized; antevaginal lamella and postvaginal lamella fused together and form a funnel like heavily sclerotized sterigma; ostium bursae cresent shaped; dustus bursae membranous but lightly sclerotized at basal portion; ductus seminalis entering at basal part of ductus bursae from latero-dorsal side, entrance marked by sclerotization; corpus bursae oval, large and somewhat elongated, membranous and signa absent; anal papillae not small, moderately sclerotized and beset with dense setae, remain withdrawn under intersegmental membrane of 8 th and 9 th segment and protrude only during egg laying; a pair of thin but prominent and sclerotized posterior apophysis present; a pair of anterior apophysis, shorter than posterior apophysis, arise from anterio-lateral margin of 8 th tergum, heavily sclerotized and bear spatulate club shaped tip.
Junonia almana (Linnaeus, 1758) (Fig. 10) Papilio almana Linnaeus, 1758 Syst.Nat.(Edn 10) 1: 472, TL: Canton, China Wing span: 60-65 mm Diagnosis: Adult (male): Upper side tawny, costal margin of forewing brown, two transverse brown lines across cell base, three dark bands across costal margin of forewing, two prominent eyespots in post discal area; hind wing suffused with dull brown scales, tawny beyond middle, a large prominent eyespot in discal area near costal, another discal eyespot small and obscure, two black submarginal lines across termen of forewing and hind wing; under side of wings seasonally variable with eyespots well developed in wet season form and eyespots obscure in dry season form.
Female genitalia (Fig. 12): Eight sternum reduced, surrounded and sustained by strong sclerotization of vaginal lamella (antero and post vaginal lamella) which in this case forms tubular sterigma; eighth tergum well developed; sterigma well developed and strongly sclerotized; ostium bursae crescent shaped; ductus bursae long, funnel shaped and membranous, lead to long and membranous corpus busae after a narrow membranous constriction; corpus bursae long, balloon shaped, membranous and narrow at base but gradually dilating towards apex; signa absent; anal papillae not large, moderately sclerotized, densely setose with short setae, kept retracted under intersegment membrane of 8 th and 9 th segment and emerge only during egg-laying; anterior and posterior apophyses present, thin but well sclerotized tips spatulate; posterior apophysis slightly longer than anterior apophysis.
Female: Forewing similar as male except a black band covers end cell and a prominent eyespot in lower discal area; hind wing similar as male but two small eyespots present in discal area and two subterminal black lines across termen.
Female genitalia (Fig. 15): Eighth sternum not reduced and moderately sclerotized, latter surrounded and sustained by strong sclerotization of vaginal lamella, which in this case from a U-shaped groove like sterigma; fusion of antevaginal lamella and postvaginal lamella forms a heavily sclerotized sterigma; ostium bursae crescent shaped; ductus bursae membranous and very long; ductus seminalis enters at base of ductus bursae from dorso-lateral side; corpus bursae slender, narrow at base but gradually dilating towards apex, signa absent; anal papillae moderately sclerotized and densely setose with short setae, kept retracted under intersegmental membrane of 8 th and 9 th segment and emerge only during egg-laying, equipped with a pair of posterior apophyses; anterior apophysis present, arise from anterior lateral margin of 8th tegument, shorter than posterior apophysis, crimped, both pair of apophyses have spatulate tips.
Remarks: This species prefers hottest temperatures  and is a quick flier.No prominent variation in genitalia is observed.
Junonia atlites (Linnaeus, 1763) (Fig. 16) Papilio atlites Linnaeus, 1763; Amoenitates Acad.6: 407; TL: Asia Wing span: 55-65 mm.Diagnosis: Adult (male): Upperside greyish in colour, a pair black transverse lines across mid cell, another pair of black traverse lines across end cell, a wide irregular transverse band across discal area interrupted by a series of six oval ocellated spots, of which first, second and third from costal margin are larger and brighter; hind wing upperside grey with numerous tranverse black lines around basal area, transverse band as in forewing except only five ocellated spots are present.Underside numerous transverse line around basal area in fore wing and hind wing, a prominent dark brown line across discal area, only three eyespots on fore wing and two eyespots in hind wing are distinct.
Range country: India, Sri Lanka, Myanmar Discussion: This species is not so common in the