Review of the cicada genus Paharia Distant ( Hemiptera , Cicadidae ) , with the description of a new species and its allied species

The genus Paharia Distant, 1905 is reviewed based on the description of a new species, Paharia oorschoti sp. nov., and redescription of the allied Paharia putoni (Distant, 1892), both from Turkey. The relationships among Paharia, Subpsaltria Chen, 1943 and Tibicina Kolenati, 1857 of the tribe Tibicinini Distant, 1905 are discussed. The morphology of the exuviae of Pa. oorschoti sp. nov. and S. yangi Chen, 1943 is described and compared. Tibicina insidiosa Boulard, 1977 is transferred to Paharia to become Paharia insidiosa comb. nov. A key to all species of Paharia is provided.

In the present paper we review the genus Paharia based on the description of one new species (Paharia oorschoti sp.nov.) from Kurubas Geçidi, Turkey, and the redescription of the allied Pa.putoni from the same locality and from Iran.In addition, Tibicina insidiosa Boulard, 1977 is transferred to Paharia to become Paharia insidiosa comb.nov.The relationships among Paharia, Subpsaltria and Tibicina are discussed.The morphology of the exuviae of Pa.oorschoti sp.nov.and S. yangi Chen, 1943 is described and compared.

Material and methods
This study is based on specimens deposited in the following institutions: NWAFU = Entomological Museum, Northwest A&F University, Yangling, China ZMAN = Zoölogisch Museum, Amsterdam, The Netherlands The type specimens of the new species were deposited in ZMAN (collection now held in Naturalis Biodiversity Center (NBC), Leiden, The Netherlands) and NWAFU.Various other material of Paharia and the related Subpsaltria and Tibicina was also borrowed from the above three institutes.
External morphology was studied using an Olympus SZX10 stereo microscope (Olympus Corporation, Tokyo, Japan), and photographed with a Nikon Coolpix P100 digital camera (Nikon Corporation, Indonesia).Male genitalia were studied and photographed using a scientific digital micrography system equipped with an auto-montage imaging system and a highly sensitive QIMAGING Retiga 4000R digital camera (CCD) (QImaging, Surrey, BC, Canada).Photographs were modifi ed with Adobe Photoshop CS3.Terminology for morphological features and classifi cation follows that of Moulds (2005Moulds ( , 2012)).All measurements are in millimeters.

Diagnosis
Head moderately narrow, including eyes less than 0.80 times as wide as base of mesonotum; postclypeus much protruding, rostrum reaching mid coxae.Pronotum convex, much longer than head, laterally depressed; lateral pronotal collar not dentate.Abdomen about as long as forebody; timbal covers entirely absent, tympana completely exposed; abdominal sternites with lateral margins strongly recurved.Wings hyaline, with 8 and 6 apical cells on fore wing and hind wing respectively; fore wing with ulnar cell 3 more than twice as long as apical cell 5, basal cell about twice as long as broad and narrower at apex than at base.Aedeagus much elongated, curved anteriorly, with apex bifurcated.

Diagnosis
This species can be recognized by the following combination of characters: a yellow drop-shaped spot extending from median ocellus to posterior margin of head; mesonotum black with a pair of large, irregularly shaped fasciae on disc; male uncus black, with median uncal lobe pale brown and tapering to apex; aedeagal shaft elongated, curved anteriorly and then posteriorly, apex sickle-shaped; female opercula reaching posterior margin of abdominal sternite II but not overlapping each other centrally.

Etymology
The species is named after one of the collectors, H. v. Oorschot.

Male
ABDOMEN (Fig. 1A-B, E).Black with no distinct markings, about as long as head and thorax together.Timbal covers yellow, undeveloped dorsally and leaving dorso-lateral timbal cavity wide open.Opercula black with yellow posterior margin, not overlapping each other centrally, not reaching posterior margin of abdominal sternite II.Abdominal sternites black and covered with golden hairs; deeply depressed, with epipleurites strongly developed and very prominent; epipleurites black with lateral and posterior margins yellow; sternite VIII longer than combined length of sternites VI and VII.GENITALIA (Fig. 1F-G).Pygofer elliptical in ventral view, with long, silvery hairs.Basal lobes of pygofer curved inward distally.Anal styles yellow.Uncus black; median uncal lobe pale brown and tapering to apex; apex pointed and lower margin curved inwardly in lateral view.Aedeagal shaft elongated, curved anteriorly and then posteriorly, apex sickle-shaped.

Female
ABDOMEN.Dorsally black without distinct marking, abdomen about as long as head and thorax together.Opercula black-brown with yellow posterior margin, reaching posterior margin of abdominal sternite II but not overlapping.Abdominal sternites dark brown to black and covered with golden hairs; epipleurites black, strongly developed and very prominent, with lateral corners and posterior margins orange-yellow.Sternite VIII with deep median incision, black with orange-yellow coloration along incision and on broad distal lobes.Lower part of segment IX orange-yellow to yellowish brown.Exuviae BODY (Fig. 2A-B).Curved in lateral view, with sparse setae mainly on venter.Pronotum yellow with fuscous transverse fascia on posterior margin.Mesonotum yellow with two fuscous transverse fasciae on posterior area.Abdomen yellow with fuscous transverse fasciae on anterior margin of each tergite.LEGS (Fig. 2C-E).Generally yellow with black markings on apices of fore femur.Fore femur with posterior tooth long and sharp, accessory tooth short and robust, intermediate tooth curved upward in lateral view; femoral comb with fi ve teeth, the fi rst shorter than the second.Fore tibia arched, with apical tooth long and pointed.Apex of mid tibia with three spines; apex of hind tibia with two spines.

Remarks
This species is similar to Pa. putoni, but can be distinguished by the following characters: mesonotum with a pair of large, irregularly shaped markings (mesonotum with pair of somewhat quadrangled markings in Pa.putoni); male median uncal lobe pale brown, with apex pointed and lower margin curved inwardly in lateral view (male median uncal lobe black in Pa.putoni, with apex blunt and lower margin nearly straight in lateral view); female opercula reaching posterior margin of abdominal sternite II (female opercula not reaching posterior margin of abdominal sternite II in Pa.putoni).
HEAD (Fig. 3A-B).Head including eyes much narrower than base of mesonotum; mostly black with a thin yellow spot extending from median ocellus to posterior margin of head.Distance between lateral ocellus and corresponding compound eye about equal to distance between lateral ocelli.Eyes green, THORAX (Fig. 3A, C-D).Inner area of pronotum black and distinctly longer than head, covered with dense golden hairs; a faint central longitudinal fascia yellow.Pronotal collar black without markings, covered with golden hairs.Mesonotum black with somewhat quadrangled marking around corresponding scutal depression; pair of yellow fasciae along lateral margins of mesonotum.Cruciform elevation yellow.Thoracic sternites black, covered with dense golden hairs.
LEGS (Fig. 3B, H).Mostly yellow.Fore femur with black marking near posterior margin in lateral view; primary spine pointed apically and slightly oblique to femur; secondary spine large, curved downward; subapical spine undeveloped.Trochanter with black marking.Tibiae fuscous to black basally.

Male
ABDOMEN (Fig. 3A-B, E).About as long as head and thorax together; black with white pruinosity on lateral areas and golden hairs on median areas of each tergite.

Distribution
Iran, Turkmenistan, Afghanistan, Kazakhstan and Tajikistan.

Remarks
This species is morphologically very close to Pa. oorschoti sp.nov.(for detail, see Remarks for Pa.oorschoti sp.nov.).

Distribution
Afghanistan and Tajikistan.

Discussion
In the present paper we review the genus Paharia based on the description of one new species (Pa.oorschoti sp.nov.), the redescription of Pa.putoni and a comparative morphological study of this genus with the closely related Tibicina and Subpsaltria.Chen (1943) established Subpsaltria with the description of two species (i.e., S. yangi and S. sienyangensis) and placed it in the subfamily Tettigadinae, remarking how "this genus appears to be similar to Paharia, except for the presence of a pair of striated areas situated on each of the sides of the mesonotum, and a projecting scraper formed by the base of the claval area of each fore wing".However, Liu (1978) did not include Subpsaltria in his "Monograph of Chinese Cicadidae".Chou et al. (1997) synonymised S. sienyangensis with S. yangi and followed the division of Chen (1943), i.e., placing it in the subfamily Tettigadinae.Amyot (1847) established the genus Tibicina for Cicada haematodes Scopoli, 1763.Kolenati (1857) revised Tibicina and treated it as a subgenus of the genus Cicada Linnaeus, 1758.Fieber (1876) re-elevated Tibicina to generic rank, which was followed by later authors (cf.Metcalf 1963;Nast 1972;Duffels & van der Laan 1985).However, most denominations proposed by Amyot from 1845 to 1847 were rejected by the ICZN (Opinion 686 1963), and his denomination of Tibicina (Amyot, 1847) was also rejected by the ICZN later (Opinion 2165(Opinion 2006)).Thus, the name Tibicina was assigned to Kolenati (1857).Moulds (2005) put Subpsaltria in the tribe Tettigadini of the subfamily Tettigadinae and Tibicina in the tribe Tibicinini of the subfamily Tibicininae.Lee (2012) included Paharia, Tibicina and Subpsaltria, as well as six other genera (i.e., Tibicinoides Distant, 1914, Clidophleps Van Duzee, 1915, Okanagodes Davis, 1919, Okanagana Distant, 1905, Ahomana Distant, 1905and Subtibicina Lee, 2012), in the tribe Tibicinini of the subfamily Tibicininae (= Tettigadinae auct.).Sanborn (2013) listed all these nine genera in the sole subtribe Tibicinina of the tribe Tibicinini.Sanborn (2014) later transferred Ahomana to the tribe Carinetini.The tribe Tibicinini is defi ned by the following common characters: hind wing veins RP and M not fused at base; hind wing fi rst cubital cell width at distal end about equal to second cubital cell; male opercula not reaching posterior margins of sternite II; pygofer upper lobes absent; uncus very long (Lee 2012).However, phylogenetic relationships among the taxa of the Tibicinini remain unclarifi ed.
Paharia is more closely related to Subpsaltria and Tibicina among the eight currently known genera of Tibicinini in having broader fore wings, a black body with yellow markings (some species have a yellow body with black markings; e.g., Fig. 4A-B), and arched pygofer uncal lobes.However, some signifi cant characteristics of adults can be applied to distinguish these three genera.Paharia is distinguished from Tibicina by its longer fore wing ulnar cell 3 (more than twice as long as apical cell 5 in Paharia; about 1.5-2 times as long as apical cell 5 in Tibicina) and the structure of the timbals (upper part of timbal well sclerotized, with longest rib less than twice the length of shortest in Paharia; upper part of timbal weakly sclerotized, with longest rib greater than twice the length of the shortest in Tibicina) (Lee 2012;Ahmed et al. 2015).Paharia can be distinguished from Subpsaltria by the fore wing veins, which are not or very slightly incrassated, and by the absence of the projecting scraper at the base of the fore wing.
In addition to the dissimilarity of adults, the morphology of exuviae may be informative to the taxonomy and phylogeny of Tibicinini.For example, when we compare the exuviae of Pa.oorschoti sp.nov.and S. yangi, the following distinct differentiations, in addition to the difference of fasciae on thorax and abdomen, were found to be reliable characteristics to distinguish these two genera: i) the shape of femoral teeth (posterior tooth sharp apically, accessory tooth short and robust, femoral comb with fi ve teeth in Pa.oorschoti sp.nov.(Fig. 2C); posterior tooth rounded apically, accessory tooth absent, femoral comb with four teeth in S. yangi (Fig. 4C)); and ii) the number of spines at the apex of mid and Currently, all six species of Paharia are known from West Asia to Central and South Asia (Fig. 5).The distribution pattern of Paharia species indicates that this genus occurs mainly in the Palearctic Region, with Pa.lacteipennis expanding its distribution to Pakistan and India.P. lacteipennis has the widest distributional range, i.e., from Turkey to Kazakhstan and India.The remaining species of Paharia are all endemic to related regions.The diversity of Paharia is greatest in Afghanistan, where four species are distributed, i.e., Pa.lacteipennis, Pa.putoni, Pa.zevara and Pa.insidiosa comb.nov.Accordingly, we infer that Paharia likely originated from Palearctic Central Asia.In comparison with the distribution of Paharia, Tibicina is distributed in Europe and North Africa; Subpsaltria is endemic to central China; Tibicinoides, Clidophleps, Okanagodes and Okanagana are all endemic to North America; and Subtibicina is endemic to India.These distribution patterns of Tibicinini indicate that the evolution of this tribe has been closely related to the drift of continental plates and historical climate changes.The phylogeny and phylogeography of Tibicinini await further investigations that draw on multiple sources such as the morphology of both adults and nymphs, biogeography, acoustics, molecular data, etc.  (Walker, 1850); green triangle = Pa.putoni (Distant, 1892); purple circle = Pa.semenovi (Oshanin, 1906); brown pentagon = Pa.zevara (Kusnezov, 1931); yellow hexagon = Pa.insidiosa (Boulard, 1977) comb.nov.; black circle = Pa.oorschoti sp.nov.
About as long as head and thorax together, dorsally black with white hairs on lateral and median areas of tergites but without distinct marking.Opercula black with narrow yellow posterior margin, not overlapping and not reaching posterior margin of abdominal sternite II.Abdominal sternites black and covered with silvery hairs; epipleurites prominent, black with yellow lateral margins.Sternite VIII with deep median incision, black with distal lobes orange-yellow.Lower part of segment IX orangeyellow to yellowish brown.
Timbal covers yellow with black markings, undeveloped dorsally and leaving dorso-lateral timbal cavity wide open.Opercula black with yellow posterior margin, not overlapping each other centrally, not reaching posterior margin of abdominal sternite II.Abdominal sternites black and covered with golden hairs; deeply depressed, with epipleurites strongly developed and very prominent; epipleurites black with lateral margins yellow.Sternite VIII longer than length of two preceding sternites together.GENITALIA (Fig.3F-G).Pygofer elliptical in ventral view, with long, silvery hairs.Basal lobes of pygofer curved inward distally.Anal styles yellow to fuscous.Uncus black; medial uncal lobe black and tapering to apex, with apex blunt and lower margin nearly straight in lateral view.Aedeagal shaft elongated, curved anteriorly and then posteriorly, apex sickle-shaped.FemaleABDOMEN.