Taxonomic studies on the gaudy grasshoppers ( Orthoptera : Pyrgomorphoidea : Pyrgomorphidae ) from the northeastern states of India

A survey of the northeastern states of India recorded 10 species representing five genera belonging to four tribes of the family Pyrgomorphidae. For identification, in addition to conventional morphological characters, the detailed structures of male and female genitalia were also included. All the genera studied are described. Morphometry and distribution of each species are also given.


INTRODUCTION
Pyrgomorphidae is a family of the order Orthoptera under the suborder Caelifera.
Its members are commonly known as Gaudy Grasshoppers in the superfamily Pyrgomorphoidea.Some members (tribes Atractomorphini and Psednurini) are green in colour while others (tribe Monistrini) have bright warning colours.The present study was based on conventional morphological characters along with a detailed study of genital structures of Pyrgomorphidae for a better understanding of the significance of morphological structures.A comparative study was done on the genitalia with reference to supra-anal plate and cerci, subgenital plate, epiphallus, and aedeagus of the male and the supra-anal plate, subgenital plate, ovipositor, and spermatheca of the female.
The system of classifying grasshoppers by earlier workers was mainly based on easily recognizable externally visible characters.Slifer & King (1936), Slifer (1939Slifer ( , 1940Slifer ( , 1943)), Dirsh (1957a,b), and Meinodas et al. (1982) showed the taxonomic significance of spermatheca in Acrididae.Dirsh & Uvarov (1953) studied the apical valve of aedeagus in three species of Anacridium.Dirsh (1956a,b) showed the importance of aedeagus in classifying and grouping various families of Acridoidea.Herrera & Schnidrig (1983) described the male genitalia of 64 species of Orthoptera from Navarra.Usmani & Ajaili (1993) showed the taxonomic significance of aedeagus in some Libyan species of Acridoidea.Ajaili & Usmani (1994) made comparative studies on the male subgenital and supra-anal plates and cerci in some Libyan species of Acridoidea.Kumar et al. (2014) conducted a comprehensive study on the male supra-anal plate, cerci, and subgenital plate in 12 species of grasshoppers representing six genera under four tribes belonging to the family Pyrgomorphidae.

MATERIAL AND METHODS
A survey for collection of Pyrgomorphid specimens during 2008-2011 from grasslands and agricultural fields of northeastern states was carried out.Specimens were handpicked or collected using sweeping nets.All the collected specimens were preserved in 70% ethyl alcohol.Specimens were stretched and photographed.For genitalic studies, the apical tip of the abdomen was cut and boiled in 10% KOH solution and the genital structures were isolated.All drawings were prepared under camera lucida (Nikon SMZ 25).Descriptions of male genitalia follow the terminologies used in Dirsh (1956a).All observations on morphometry are given in millimetres.
Diagnosis: Body of variable shapes, head acutely conical, fastigial furrow present.Prosternal process present.Elytra and wings fully developed, reduced, or absent.Tympanam normally present.The lower basal lobe of hind femur longer than the upper one.Brunner's organ present except in a few genera, with thin, almost cursorial hind legs.External apical spine of hind tibia present or absent.Ectophallus differentiated; cingulum capsule-like; valves of penis paired, undivided; spermatophore sac in dorsal position.Epiphallus bridgeshaped, with dorso-lateral appendices; ancore absent; lophi hook-like.Oval sclerites absent.No stridulatory mechanism known.The Pyrgomorphidae is a very well defined family, with a peculiar phallic complex which is rather uniform through the family.The relationship with other families is rather obscure and no close affinities exist.They have some common features with Lentulidae, such as undivided paired valves of the penis and the dorsal position of the spermatophore sac, and others with Ommexechidae, such as the presence of a fastigial forrow and the paired undivided valves of the penis.All other characters, however, are so distinct that the relationship is a very remote one.Pyrgomorphidae is represented in all the tropical and subtropical parts of the world by a large number of genera, a list of which appears unnecessary.1. Body usually depressed and usually rather strongly rugose (sometimes with plicate, longitudinal tubercles); coloration mottled brown or grayish; fastigium of vertex usually (but not always) short, blunt, broad; terminal segments of antennae incrassate, fused or partly so, often pitted; tegmina (when present) usually with small nodules on main veins; hind wings hyaline (sometimes faintly bluish) or infumated only; prosternum with reflexed, collar-like anterior margin and "double" tubercle; [phallic structures rather unspecialized; epiphallus with lateral plates having wide, basal, externo-lateral expansions, very strong, laterally directed lophi, and widely divergent appendices; ectophallus with wide basal emargination, short, broad ventral process, and central membrane of cingulum rather extensive, subquadrate or sub-rectangular; aedeagal scterites rather slender and acute, gonopore basal or mesal] ….... 3. Body rather heavy, somewhat depressed (especially in female), antennae inserted only a short distance in front of ocelli; infero-posterior angle of lateral pronotal lobe strongly acute; hind tarsal segments not elongate; male cerci rather specialized; [epiphallus with broad, wing-like lateral plates and lophi bifid; ectophallus with ventral process very short and pointed; endopallic apodemes with very long, forwardly directed ventral processes, aedeagal valves and sclerites very long and strongly decurved]; Southeast Asia only .Diagnosis: Small to medium size; integument finely rugose; antennae slightly compressed, shorter than head and pronotum together, their bases located in front of lateral ocelli, fastigium of vertex elongated, flat, horizontal or slightly upcurved with parabolic or angular apex; apical fastigial areolae poorly developed; head narrow, acutely conical, with a row of postocular tubercles; frons strongly oblique; frontal ridge incurved, narrow and low, shallowly sulcate, with obtuse lateral carinulae.Pronotum elongated, sub-cylindrical, slightly widening backwards; dorsum slightly flattened, crossed by three fine sulci; median carina and lateral carinar weak; metazoan much shorter than prozona, its posterior margin widely obtuse-angular, almost rounded; lateral lobe with row of low marginal tubercles; prosternal process cuneiform; elytra and wings fully developed, apex of elytron acutely attenuate; tympanal organ well developed; hind femur narrow, with external lower marginal area narrow, displaced ventrally to external medial area; lower lobes of hind knee much shorter than upper one; hind tibiae in apical part with expanded margins; external apical spine present; arolium of moderate size; male supra anal plate elongate angular; cercus subconical, straight with subacute apex; Subgenital plate short, with rounded apex.
The genus is represented by five species from this region.Bolívar, 1905 (Image 1; Morphological Characters: Build rather short and moderately stout; head and pronotum relatively short; fastigium of vertex shorter; pronotal lobe fairly deep, without a membranous area in metazona; outer face of hind femur rather strongly convexed and keeled; hind wings relatively long, their apices not falling much short of those of the tegmina at rest, extensively tyrian pink to light mallow purple at base.

Atractomorpha burri
Male genitalia: Supra anal plate elongate angular, epiproct long and narrow with obtusely conical apex, cerci as long as or slightly shorter than the epiproct, and conical at apex, paraproct smaller.Subgenital plate triangular with conical apex.Epiphallus anchor shaped bridge small, anterior projections not prominent, lateral plates broad and fused medially, broader at the base, middle piece of epiphallus much wider, lophi hooked with conical apex, appendices rod shaped, slightly expanded at apex, reaching up to the apex of lophi.Apical valve long anad narrow, apical tip pointed, longer tanan basal valve, basal valve wide at base.
Female genitalia: Supra anal plate elongated, longer than wide, apex obtusely rounded.Cercus short, longer than wide and incurved.Subgeniatal plate: posterior margin without setae, egg-guide wide at base; narrowing apically, one and half time longer than wide.Spermatheca with apical and pre-apical diverticula, pre-apial diverticulum moderately long tubular, apical diverticulum long, tubular with protuberance.Ovipositor: dorsal valve long, wide with apex pointed, slightly longer than apodeme and more than twice as long as wide, ventral valve uniformly wide, apex elongated and obtusely rounded.head and pronotum relatively long; fastigium of vertex narrower and longer; lateral pronotal lobe shallow, sometimes with a small membranous area in the metazona; outer face of hind femur not strongly convex or keeled; hind wings normally comparatively long in relation to tegmina, not falling much short of them when at rest, usually dull magenta purple, often rather pale, at extreme base, often with dark veins and somewhat infumated, sometimes colourless.Male genitalia: Supra anal plate elongate angular, epiproct triangular, long and broad at base with obtusely conical apex, cerci shorter than the epiproct and conical at apex, paraproct smaller.Subgenital plate rounded at apex.Epihallus anchor shaped, bridge much narrow, anterior projections not prominent with rounded apex, lateral plates fused medially, middle piece of epiphallus narrow, lophi hooked with pointed apex, appendices rodshaped, expanded at apex, slightly far from the apex of lophi.Apical valve narrow as long as basal valve, apex tubular, basal valve wide at base.
Female genitalia: Supra anal plate triangular, longer than wide, apex obtusely rounded, cercus short, wide at base, longer than wide, narrowing apically.Subgenital plate, posterior margin smooth without setae, notched in the middle, egg-guide narrowing apically, twice as long as wide, tip pointed.Spermatheca: apical and pre-apical diverticulum long, both are long, tubular and narrow.Ovipositor: dorsal valve broad, long, slightly longer than apodeme with apical tip pointed, ventral valve uniformly broad with tip pointed.
Material  Morphological Characters: Generally rather small; eyes generally shorter and rather convex; fastigium of vertex often comparatively short; inter-ocular space generally slightly convex; membranous area in metazona of lateral pronotal lobe usually very distinct in female and well indicated in male; hind wings normally tyrian pink to light mallow purple or pale magenta at base, but quite often rather heavily infumated.
Male genitalia: Supra anal plate elongate angular, epiproct triangular, long and slightly wider at base with obtusely conical apex, cerci shorter than the epiproct, and conical at apex, paraproct smaller.Subgenital plate oval at apex.Epihallus anchor shaped, bridge broader, anterior projections prominent with pointed apex, lateral plates fused medially, middle piece of epiphallus with subparallel margins, lophi hooked with pointed apex, appendices rod shaped, slightly expanded at apex, slightly far from the apex of lophi.Aedeagus: apical valve long, narrow, upcurved, distinctly longer than basal valve, apical tip pointed, basal valve narrow, broad at base.
Female genitalia: Supra anal plate elongated, slightly  Male genitalia: Supra anal plate broad basally, moderately narrowing apically, as long as wide, apex rounded, cercus elongated, broad basally, narrowing apically, more than one and half time as long as wide, apex obtusely rounded.Epiphallus anchor shaped, anchorae elongated, narrow basally, broad apically.Aedeagus, apical valve narrowing apically with apex pointed, twice as long as wide, basal valve short and wide at apex.
Female genitalia: Supra anal plate broad at base, slightly narrow at apex, as long as wide, apex obtusely rounded, cercus short, broad, narrow apex, one and half time as long as wide.Subgenital plate smooth without setae, notched in the middle, egg-guide short, broad at base, narrowing apically, twice as long as wide.Spermatheca, apical diverticulum long, tubular and narrow.Pre-apical diverticulum uniformly tubular, S-shaped.Ovipositor: dorsal valve elongated, broad,  Morphological Characters: Size generally a little larger; fastigium of vertex usually rather broad and generally very flat dorsally; pronotum with distinct carinae; lateral pronotal lobes without membranous area; hind wings rose red or rose, not frequently infumated.
Male genitalia: Supra anal plate elongate angular, epiproct triangular, long and wider at base with obtusely conical apex, cerci shorter than the epiproct and conical at apex, paraproct smaller.Subgenital plate flat and globular at apex.Epihallus anchor shaped, bridge wider, anterior projections prominent with hook like apex, lateral plates fused medially, broader at base, middle piece of epiphallus slightly narrow, lophi hooked with obtusely conical apex, appendices rod shaped, much expanded at apex, slightly far from the apex of lophi.Aedeagus apical valve elongated, uniformly tubular, upcurved, apex obtusely rounded, basal valve short and broad.
Material Diagnosis : Body usually depressed and usually rather strongly rugose (sometimes with plicate, longitudinal tubercles); coloration mottled brown or greyish; fastigium of vertex usually (but not always) short, blunt, broad; terminal segments of antennae incrassate, fused or partly so, often pitted; tegmina (when present) usually with small nodules on main veins; hind wings hyaline (sometimes faintly bluish) or infumated only; prosternum with reflexed, collar like anterior margin and double tubercle; epiphallus with lateral plates having wide, basal, externo-lateral expansions, very strong, laterally directed lophi, and widely divergent appendices; ectophallus with wide basal emargination, short, broad ventral process, and central membrane of cingulum rather extensive subquadrate or sub-rectangular; aedeagal sclerites rather slender and acute, gonopore basal or medial.to the tip of abdomen.Hind tibiae nearly equal to hind femur with eight outer and nine internal spines.Male genitalia: Supra anal plate angular with broad and short epiproct, cerci as long as or slightly longer than the epiproct, obtusely conical at apex, paraproct broad and slightly longer than epiproct.Subgenital plate triangular and conical at the end.Epiphallus bridge shaped, bridge moderately slender, slightly wider and long; anterior projection prominent with angular tip; lateral plates separated medially, lophi with curved apices, apex acute; lateral appendices rod shaped and hooked at the end, apex pointed and reaching the tip of lophi.Aedeagus, apical valve slightly upcurved, apex pointed, longer than basal vave.Morphological Characters: Body yellowish brown with metazona of pronotum and mid of hind femur outer and upper surface white.Antennae eleven segmented, shorter than head and pronotum together.Lateral carina of pronotum represented by weak lines only in metazona.Tegmina slightly reaching up to the apex of hind femur.Hind tibiae with seven outer and eight inner spines.
Male genitalia: Supra anal plate angular with broad and long epiproct, cerci shorter than the epiproct, broad and obtusely rounded at apex, paraproct is also broader and as long as or slightly longer than the epiproct.Subgenital plate broad and rounded at the apex.Epihallus bridge shaped, bridge more slender, slightly narrow and elongated; anterior projection less prominent; lateral plates separated medially, lophi with curved apices, apex acute; lateral appendices rod shaped and hooked at the end, apex expanded and crossing the tip of lophi.Aeadegus apical valve narrow, tubular, as long as basal valve, basal valve broad, wide.Diagnosis: Small and slender; body slightly depressed, integument rugose and hairy; antennae in apical half thickened, with fused segments, shorter than head and pronotum together; fastigium of vertex short, sloping forwards, slightly concave, with obtuse angular apex and deep; large apical fastigial areolae and sharp marginal carinulae; weak occipital carina present; frontal ridge between antennae compressed and protruding forwards, with slit-like sulcus, below almost obliterated.Pronotum subcylindrical, widening backwards, tuberculate, with weak linear median carina, crossed by three sulci, lateral carinae absent, metazona about as long as prozona, its posterior margin rounded; anterior margin of prosternum strongly expanded, collar like, covering lower part of mouth; elytra and wings fully developed; tympanum absent; middle femur and tibiae elongated, thin; hind femur slender, with lower basal lobe slightly shorter than upper one; hind tibiae slightly expanded in apicl half; external apical spine absent; spurs of hind tibiae strongly elongated, longer than basal tarsal segment, thin, internal pair longer than external; all tarsi thin, slightly elongated; arolium very small; male supra anal plate angular; cercus short, obtusely conical; subgenital plate short, widely subconical; epiphallus with large lophi, moderately curved at apices.
The genus is represented by a single species from this region.
Morphological Characters: Body slightly depressed, integument rugose and hairy; antennae shorter than head and pronotum together; fastigium of vertex short, slightly concave; large apical fastigial areolae and sharp marginal carinulae.Pronotum subcylindrical, widening backwards, tuberculate, with weak linear median carina, crossed by three sulci, lateral carinae absent, metazona about as long as prozona, its posterior margin rounded; elytra and wings fully developed; middle femur and tibiae elongated, thin; hind femur slender, with lower basal lobe slightly shorter than upper one; hind tibiae slightly expanded in apical half.

Taxonomic studies on gaudy grasshoppers Khan et al. 12956 Tribe Atractomorphini Bolivar, 1905 Dignosis:
Body slightly robust, not depressed, or, if so slightly (some females), hind tarsal segments elongate; head elongated, acutely conical, fastigium of vertex usually long, acute, a row of fine, distinct granular tubercles extending from behind the eye across the inferior margin of the lateral pronotal lobe; inferoposterior angle of lateral pronotal lobe not strongly acute; antennae usually inserted well in advance of ocelli; tegmina usually fully developed, usually very tapered and pointed.Brachypterous forms, somewhat depressed (especially in females), strongly micropterous or apterous forms unknown; hind femur trigonal in cross section; posterior tibial spines rather long and sharp; male cerci unspecialized; ventral process of ectophellus moderately long, very broad and subtruncated apically.