Sabahia polypodii gen. et sp. nov. (Hemiptera: Cicadellidae: Evacantihinae) and its Phylogenetic Position within the Nirvanini Tribe

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constitutes a relatively small tribe of a pan-tropical range, which is limited in its distribution to equatorial and subtropical humid forest zones (montane cloud forests) and other humid mountain habitats. It is associated with a variety of host plants belonging to the class Magnoliophyta. Its representatives feed on some trees (e.g. of the genus Aesculus, Quercus, Acacia), as well as shrubs (e.g. Citrus, Camellia, Vitis) and numerous perennials and grasses (SCHUMACHER 1915;AHMED & MAHMOOD 1970;MESHRAM & RAI 2017). Some species of the genus are regarded as pests among farmers growing sugarcane and horticultural crops (VIRAKTAMATH & WESLEY 1988), or even show features typical of an invasive species (WEBB & VIRAKTAMATH 2004;AGUIN-POMBO et al. 2007). The group was previously classified as the subfamily Nirvaninae Baker, 1923(OMAN et al. 1990, which is now considered to be a junior synonym of Evacanthinae (DIETRICH 2004). In the past, numerous genera of the current Nirvanini were often placed in the subfamily Typhlocybinae (MATSUMURA 1931). However, five tribes have recently been assigned to the subfamily Evacanthinae Metcalf, 1939: Nirvanini Baker, 1923, Balbillini Baker, 1923, Evacanthini Metcalf, 1939, Pagaroniini Anufriev, 1978, and Pentoffiini Wang, Dietrich & Zhang, 2017(WANG et al. 2017. The subfamily shows many similarities, and is most probably closely related, to the subfamily Cicadellinae (DIETRICH 2004). MESHRAM and RAI (2017) presented an updated review of the taxonomy of the Nirvanini tribe.
In spite of numerous morphological analyses and other comparative studies, no single and sufficiently unambiguous features could be identified which would distinguish the Nirvanini tribe from the remaining ones (especially from Evacanthini) (DIETRICH 2004;WANG et al. 2017).
The Nirvanini tribe includes mostly medium-sized species with an elongated, slender body which is flattened dorsoventrally. Light colouration is dominant, generally with a species-specific colour pattern in the form of red, orange and black stripes, streaks and spots (on both the dorsum and the wings). The head is wide, equal to the pronotum or wider. The vertex is predominantly elongated (or strongly elongated) and flattened (but not concave), with a distinct sutura coronalis, rarely extending beyond the ocelli. Vertex margins are arcuate, converging at the apex, rounded or pointed. Typical of Evacanthini, the double lateral carinae are absent. In some species (e.g. of the genus Chudania Distant), the lateral parts of the vertex are covered with delicate sculpture. Ocelli are located near the vertex margins above compound eyes. In the lateral view, the frontoclypeus is usually flat, forming an acute angle with the vertex line (e.g. Kana Distant, Nirvana Kirkaldy, Chudania Distant and some species within the genus Sophonia Walker), more rarely broadening below the apex (e.g. in some species of Sophonia Walker), or sporadically forming an almost right angle (Decursusnirvana Gao & Zhang). Median carina of the frontoclypeus are short, and only sporadically longer (e.g. Chudania delecta Distant) or absent. Clypeus is wide, trapezoidal in shape and lacking median carinae. Lora are very fine. Lower edges of the gena (maxillary plates) are equal to the clypeus or extending beyond its apex. This is the only characteristic to unequivocally separate the Nirvanini species from those of Evacanthini and Pagaronini. Antennae are long, reaching at least the end of the pronotum or wing apices. Antennal carinae are distinct. Pronotum is with lateral carinae. There are longwinged forms. Forewings are with the transverse venation reduced, limited to the wing apex. The wing appendix is considerably reduced. Four external cells are subequal (e.g. Kana Distant), or the fourth (inner) is twice as long as the remaining ones, while being rectangular or irregular in shape (e.g. Sophonia Walker and Nirvana Kirkaldy). Hindwing is always developed, with three or four apical cells. In the latter case, a marginal vein runs close to the outer wing margin and joins its costal margin near the nodular area (e.g. Kana Distant, Decursusnirvana Gao & Zhang, Sinonirvana Gao & Zhang). Fore and middle tibiae are cylindrical. The ventral row of setae on the protibia are with a single seta distinctly longer than others. Metafemur is with a macrosetal formula 2+1+1, sometimes with supernumerary setae. Pygofer is cylindrical or flattened, with more or less clearly delineated lobes and ventral processes. Apices of the pygofer are with a numerous macrosetae. Anal tube is with processes or processes absent. Genital valve is relatively minute. Genital plates are usually elongated, almost equally narrow, and sometimes tapering (e.g. Nirvana Kirkaldy) or with a broadened and upwardly bent apex (e.g. Decursusnirvana Gao & Zhang). Style is differing in shape, hooked or awl-like elongate (e.g. Nirvana Kirkaldy). Connective is Yshaped with sub-equal arms and T-shaped with an elongated basal arm. Aedeagus are with a single shaft and gonopore, often with long and numerous processes, rarely with processes absent. Some species characteristically have a strongly-reduced atrium at the shaft base (e.g. Decursusnirvana Gao & Zhang, Sinonirvana Gao & Zhang). Details of the ovipositors in Nirvanini have been described quite recently, while the sculpture, shape and distribution of teeth on the ovipositor valvules are of diagnostic value for some Nirvanini species (VIRAKTAMATH 1992;DIETRICH 2011aDIETRICH , 2011bWANG et al. 2017). The assignment of Sabachia polypodii gen. et sp. nov. to the tribe Nirvanini, and the subfamily Evacanthinae, is confirmed by a set of characteristics including well-developed ocelli located near the anterior vertex margin, cylindrical fore tibia, macrosetal formula on the hind femur apex 2+1+1, medial carina on the frontoclypeus absent, basal seta in the fore femur ventral row that is distinctly longer than others, and a lower margin of the maxillary plate slightly extended beyond the lower margin of the anteclypeus.

Materials and Methods
The research was carried out in 2011 on the research plots near D'Villa Rina Ria Lodge in Ranau District (Borneo, Sabah). The coordinates of the location are: 6°00N02ON, 116°32N45OE. The research was conducted in August at night (during a random night trip). The insects were found using a standard entomological sweep net (i35 cm), in accordance with the methodology commonly used for this type of research (GÊBICKI et al. 1977;KLIMASZEWSKI et al. 1980;STEWART 2002). The research with the use of an entomological sweeping net was supplemented by viewing the ferns (possibly the host plant of this species) with Sabahia polypodii gen. et sp. nov. specimens.
Dry-mounted specimens were studied under a Leica M205C microscope, Leica DM3000LED stereomicroscope and a Delta Optical IPOS-810 stereomicroscope. Measurements were made with an ocular micrometer. The SEM micrographs were obtained using a Phenom XL field emission scanning electron microscope with a BackScatter Detector (BSD). The panoramic image stitcher Image Composite Editor 2.0.3.0 and the graphic editor Adobe Photoshop CS6 were used to prepare the figures. No procedures that could damage the specimens, e.g. washing, dehydration and sputter-coating with a film of electrically conducting material, were used. The specimens were only cleaned with a brush and were then mounted on aluminium stubs (by inserting an insect pin into a fragment of a sponge that was glued to the stub). Next, the specimens were covered with an anti-static spray. The specimens were measured using the Leica Application Suite 4.9.0 software. The genitalia of the insects were dissected after boiling the abdomen three times (about 10 minutes) in a 10% solution of potassium hydroxide (KOH). This preparation was performed according to the KNIGHT (1965) procedure. Then, the specific parts of the genital structures were separated from the abdomen using thin forceps and a needle blade. The whole was then placed in glycerine. The genitalia were subsequently dissected and examined using a Delta Optical IPOS-810 stereomicroscope and a Leica DM3000LED stereomicroscope. Photographs of the habitus were taken with a Canon EOS 50D digital camera equipped with a Canon 100 mm f/2.8 USM Macro and a Sigma 150 mm f/2.8 USM APO Macro DG HSM lens. Photographs of the genitalia were taken using a Nikon D7500 camera and the Nikon M Plan10 lens. The images were stacked, aligned and combined using the Helicon Focus 7 (the first author is the licence holder). The map was generated with the QGIS program (GNU General Public Licence) and using a vector map downloaded from the website: https://www.naturalearthdata.com/.
The holotype and nine paratypes are preserved in the entomological collection of the University of Silesia in Katowice, Faculty of Natural Sciences, Institute of Biology, Biotechnology and Environmental Protection (Katowice, Poland), and one of the paratypes is in the collection of the author: Marcin Walczak.

Taxon sampling
The collected data on the specimens and their Gen-Bank accession numbers are provided in Table 1. DNA extraction, amplification and sequencing DNA was extracted from one leg of each specimen. The genomic DNA was isolated without modifying the protocol, using the Syngen DNA Mini kit (Syngen, Wroc³aw, Poland) under a laminar flow. To elute the purified DNA, we applied 50 ìl of an Elution Buffer onto the silica membrane. Two of the genes were amplified: one mitochondrial COI and one nuclear H3. To amplify a fragment of the mitochondrial cytochrome c oxidase I (COI) gene, the primer pair LCO1490 and HCO2198 (FOLMER et al. 1994) was used. For the H3 amplification, the primers H3AF and H3AR (COLGAN et al. 1998) ware used. A polymerase chain reaction (PCR) amplification for all the DNA fragments that were analysed was carried out under a laminar flow in a final volume of 20 ìl containing: 30 ng of DNA, 1.25 U Perpetual OptiTaq (EURx, Poland), 0.4 ìl of 20 ìM of each primer, 2 ìl of 10x Pol Buffer B and 0.8 ìl of 5 mM dNTPs in a Mastercycler ep system (Eppendorf, Hamburg, Germany). The cycling profile for the PCR was: 95°C for 2 min, 35 cycles of 95°C for 30 sec, Tm of oligos for 30 sec, 72°C for 1 min and a final extension period of 72°C for 7 min.
In order to assess the quality of the amplification, the PCR products were electrophoresed in 1% agarose gel, for 45 min at 85 V, with a DNA molecular weight marker (Mass Ruler Low Range DNA Ladder, Thermo-Scientific, Waltham, MA, USA). The PCR products were purified using the Syngen Gel/PCR Mini Kit (Syngen, Wroc³aw, Poland).
The samples were sequenced in both directions using the same primers as for the PCR reactions, combined with a BigDye Terminator 3.1 Cycle Sequencing Kit (Applied Biosystems, (ABI) Foster City, CA, USA) using the chain termination reaction method (SANGER et al.1977). The sequencing reaction was carried out with the PCR product at a total volume of 20 ìl containing: 2 ìl of BigDye Terminator Reaction Ready Mix v. 3.1 (ABI), 2 ìl 5× sequencing buffer (ABI), 3.2 mol/ìl of the primer solution and 6 ìl of the purified PCR product. The cyclesequencing profile was 3 min at 94°C, followed by 30 cycles of 10 s at 96°C, 5 s at 50°C and 2 min at 60°C.
The sequencing products were precipitated using the ExTerminator (A&A Biotechnology, Gdynia, Poland), and were separated on an ABI PRISM 3130xl DNA Sequencer (Applied Biosystems, Foster City, CA, USA).

Sequence edition and alignment
The raw chromatograms were evaluated and corrected in Geneious v. R10.2.6 (https://www.geneious.com). In order to identify the NUMTs (BENSASSON et al. 2001;SONG et al. 2008), the mitochondrial COI sequences were translated into amino acid sequences with Geneious v. R10.2.6 using the standard invertebrate mitochondrial genetic code. All of the nucleotide sequences were verified using BLAST searches of the NCBI (http://blast.ncbi.nlm.nih.gov/Blast.cgi). The alignment of the combined sequences that were studied was performed using the MAFFT (KATOH et al. 2002) plugin available within Geneious v. R10.2.6.

Molecular phylogenetic
Models for the Bayesian (BI) analyses were calculated in jModeltest (DARRIBA et al. 2012;GUINDON & GASCUEL 2003) using the Akaike information criterion (AIC). The BI analyses were performed in MrBayes v. 3.2.6 (RONQUIST et al. 2012) with four independent runs, each having three heated and one cold chain. Analyses were run for 2 million generations, with trees sampled every 1000 generations. The first 25% of each run was discarded as a burn-in. The convergence among the runs was assessed using Tracer (RAMBAUT et al. 2018). The ML analyses were performed using PhyML v. 3.0 (GUINDON et al. 2010). All the trees were visualised using FigTree (http://tree.bio.ed.ac.uk/software/figtree). The trees were edited and annotated in Corel Draw 17.1.0.572 (Corel Corporation, 2014).

Results
Phylogenetic inferences were obtained for the H3 and COI genes using the Bayesian inference (BI) and Maximum likelihood (ML). The GTR+G+I model with a gamma correction of 1.389 and invariable sites of 0.548 was selected by the AIC in jModelTest for the matrix. Heuristic searches resulted in one ML tree (-ln= -9591.77984). The phylogenies were resolved and the main splits were supported by high bootstrap values and posterior probabilities. Only four nodes had bootstrap scores <80, and eight nodes had posterior probabilities <90 (Fig. 1).

Tribe -Nirvanini Baker
Sabahia gen. nov. (Figs 1-8) Type species: S. polypodii sp. nov. D i a g n o s i s. Medium-sized Nirvaninae. Vertex is slightly shorter than the pronotum, with the anterior margin arcuate. In the lateral view, the lines of the vertex and frontoclypeus form an almost right angle. Frontoclypeus is lacking median carina, with the lateral margins subparallel. Antennae are almost reaching the end of the body. Clypeus (anteclypeus) is elevated medially. Wings are considerably longer than the body. Apical cell 4 of the forewing is longest; marginal vein of the hindwing running close to the wing margin (especially in the apical part), with 4 apical cells present. Male pygofer is with a narrow rounded apex, divided into a dorsal and a ventral part by a horizontal groove, with a pair of hook-like ventral processes. Apical part of the style (with a ventral lobe) is distinctly shorter than the basal one, and is connective typical of the tribe. Aedeagus is lacking an atrium and is without spines or processes on the shaft, which is divided into two fused parts: the dorsal and the pointed, strongly sclerotised ventral one. Massive sternite VII plate in the female is slightly longer medially. Apex of the first pair of valvulae are elongate and pointed, while ventral margins of the second pair of valvules are arcuate. D e s c r i p t i o n. A medium sized genus (females app. 4.9 mm, males app. 5.5-5.6 mm) (Tab. 2, Figs  2a-b, 3a-b).
H e a d. Vertex (crown) is relatively short, distinctly broader around than between the eyes (0.78 x) and between the antennae (0.66 x). Anterior margin of the vertex is gently arcuate, lateral margins undulated, edged by a weakly defined carina (more distinct in the apical part of the head). Vertex surface is almost flat, with small depressions on the sides apically, sutura coronalis is short, transforming into a low median carina extending somewhat above the line of the ocelli. Apical part of the vertex is slightly convex, covered with numerous cuticular wrinkles converging apically (Fig. 4a). Ocelli, relatively large, are located just above the upper margins of the compound eyes and are separated from the apical margin by a distance equal to 2x the ocellus width. The edge between the vertex and frons elevated, visible from above, is with numerous cuticular wrinkles (Fig. 4a). Lateral angle of the head formed between the line of the vertex and medial line of the frontoclypeus is almost a right angle (Fig. 2b). Facial part of the head with the labrum is evenly covered with cuticular nodules, and the surface of the gena is glabrous (Fig. 4b).
Lateral margins of the upper part of the frontoclypeus are almost parallel in 2/3 of their course, and are distinctly narrowed along the basal 1/3 (Fig. 4c). Median carina are completely missing (Fig. 4b-c). Lateral surfaces of the frontoclypeus are with 7 indistinct pairs of cuticular carinae, which obliquely converge to a slightly elevated medial part of the face (where the mouthpart muscles are attached) (Fig. 4c). Clypeal groove is deep and gently arcuate. Clypeus is wide, app. 1.3 x shorter than the frontoclypeus. Median carinae are absent. A triangular protruding part of the clypeus is terminating on the sides with flatter fragments (Figs 4b-c). Lower margin of the clypeus are only 0.6 x shorter   than the upper one, straight and are slightly incised medially. Lora are very short and narrow, expanding from the vicinity of the clypeal groove to the subapical part of the clypeus. Lateral margins of the large maxillary plates are straight, reaching beyond the level of the clypeus. Rostrum is short, extending to the middle coxa (Figs 4b-c). Compound eyes are large, almost globular (in the lateral view), with a minute antennal incision on the inner margin (Fig. 2b). Antennal pit is broad, just anterad of the anterior eye margin (Figs 4b-c). Antennae are very long, reaching the end of the body, more than 8x longer than the head length (Figs 2a, 3a). Antennal scapus is slightly flattened, twice as wide as the cylindrical pedicellus (Fig. 4b).
T h o r a x. Pronotum is shield-shaped, smooth and somewhat longer than the head (app.1.3x), with the anterior margin arcuate and the posterior margin almost straight (slightly incised medially). Lateral margins (at the wing base) are deeply incised, lacking distinct carinae. Anterior part of the head capsule is basally distinctly swollen (Fig. 4a). Scutum (together with the scutellum) is shorter than the pronotum (0.8x), scutellar groove is deep and complete. Basal part of the scutum is furrowed and distinctly separated from a glabrous scutellum (Fig. 4a).
W i n g s. In both sexes, the wings reach far beyond the tip of the abdomen (Figs 3a-b). Forewings are elongate (especially in males) and relatively narrow, 3.4x as long as wide. Transverse venation is limited to r-m and m-cu in the apical part. Cells between the longitudinal veins and apical parts of three apical cells are covered with cuticular granulation and are opaque. Longitudinal veins are covered with macrosetae. There are four apical cells, including the fourth -the longest are regular in shape, resembling rectangles (especially cells 2 and 3). Appendix is strongly reduced, with veins PCu and A1 on the clavus distinct (Fig. 5a). Hindwing is elongate, 2.6x longer than wide. Four apical cells are present. Marginal vein runs close to the wing margin (especially in the apical part) joining vein R at the nodal part of the anterior margin, and vein m-cu is very short but distinct, located just below the M bifurcation (Fig. 5b).
L e g s. Fore femur basally with a 1 macroseta in the interior row (AV 1) and numerous microsetae; fore tibia is with a row of macrosetae gradually diminishing in length (Fig. 6a). Middle femur is slightly arcuate with 1 apical macroseta above the pit of the knee joint. Hind coxa is very wide, distinctly extending beyond the lateral margins of the abdomen, with a macrosetal formula on the femur apex 2+1+1 (Fig. 6a), and basitarsus almost 3x as long as the remaining tarsal segments (Figs 6b-d), with the apical margin straight, provided on the ventral side with two longer bristles (limiting the segment mobility) and a characteristic row of 6 short spines and 2 platellae. The margin of the shortest segment II is with 2 bristles, 4 spines and 2 platellae (Fig. 6b).
A b d o m e n. Male pygofer are elongate and relatively narrow, divided by a longitudinal groove into a higher dorsal part and a broader ventral part with a pair of hooked processes. Apical and lateral surfaces are with long macrosetae (extending beyond the pygofer) with a characteristic longitudinal sculpture. Anal tube is short and strongly sclerotised (Fig. 7a). Broad genital plates are with apices tapering and strongly upward bent, basally distinctly broadened and equipped with a row of moveable margin macro-  setae. Genital valve is triangular in the outline (Figs  7a-b). Style is hook-like, with the apical part (including an indistinct internal lobe) shorter than the narrower basal part (Fig. 7c). Connective is Y-shaped with short divergent arms (Fig. 7d). Aedeagus is lacking an atrium, differentiated into S-shaped curved dorsal part and a strongly sclerotised, pointed ventral part. Shaft processes are absent, with apical gonopore (Figs 8a-b). Female pygofer are uniform, with a long row of macrosetae and a very small anal tube. Sternite VII plate is slightly narrower than that of sternite VI, elongated medially (Fig. 3b). Valvula 1 is regularly arcuate (Fig. 8c). Valvula 2 is long and pointed at the apex, slightly folded to the ventral side and with sparse minute teeth on the ventral margin (Fig. 8d).

(Figs 1-8, Tab. 2)
S p e c i e s d e s c r i p t i o n. Body is almost uniformly light in colouration, predominantly dark yellowish (Figs 2a-b). Ocelli are with a broad dark rim (Fig. 2a). Eyes are yellow, sometimes with a black, mottled, irregular colour pattern (Fig. 2b). Frontoclypeus is light with two longitudinal yellowish-brown streaks (Fig. 4c). Sides of the scutum are with blurry yellow patches and three characteristic longitudinal black stripes medially. Scutellum is yellow (Fig. 2a). Margins of the corium are yellowish. Along the forewing sutura coronalis (especially in 2/3 of its length) there is a dark yellow longitudinal stripe transforming into a dark, arcuate colour streak overlapping all anal cells (Figs 2b, 5a). Hindwing is colourless (Fig. 5b). Abdominal sternites are yellowish-brown with lighter posterior margins (Figs 3a-b). A relatively narrow pygofer with a rounded apex is endowed with 12 apicolateral macrosetae on each side. Macrosetae length and pygofer width are subequal -visible in the lateral view (Fig. 7a). Style apex is narrow and pointed, with a fine subapical tooth on the inner margin (Fig. 7c). S-bent shaft of the aedeagus is transforming into a strongly sclerotised and terminally pointed ventral part (Figs 8a-b). First valvula is with a pointed apex, second valvula is with the ventral margin regularly arcuate, marginal teeth are broad and equidistant . Detailed dimensions of the species described here are presented in the E t y m o l o g y. The genus name is derived from the name of Sabah Province (Borneo: Malaysia) where the specimens were collected, while the species name polypodii draws attention to its trophic association with ferns.

Discussion
Sabahia polypodii sp. nov. is most probably an endemic taxon limited in its distribution to north-eastern Borneo. It is trophically associated with ferns (Polypodiopsida Cronquist, Takht. & W. Zimm.). The representative specimens of the species were captured while sweeping ferns mostly by torchlight. The insects remained calm, sitting along the fern leaves.
When touched, they moved away, jumped or even flew away for a longer distance. Some seemed to be feeding in spite of the late hour. All the specimens were collected from a small area at about 3 a.m. The temperature was relatively low (about 20 o C), due to the influence of the Kinabalu Mts. Field studies have clearly indicated that the taxon is mono-or oligophagous and trophically associated with small ferns found on the cloud forest floor.
Sabahia polypodii gen. nov. and sp. nov. is most closely related to the species of the genus Decursusnirvana Gao & Zhang and to a monotypical genus Sinonirvana hirsuta Gao & Zhang, which come from central and southern China (GAO et al. 2014). This is reflected most clearly in the morphological characteristics of the head, wings and the structures of the male copulatory block, as well as by DNA sequences (the analyses did not include Sinonirvana Gao & Zhang).
Both of the previously described genera (GAO et al. 2014) are characteristically endowed with processes or spines on the shaft of the aedeagus, which is a feature that separates them from Sabahia sp. nov. In Sinonirvana Gao & Zhang, there is a pair of lateral processes at the base and a pair of apical lateral processes, whereas Decursusnirvana aedeagus possesses a triangular ventral process, as well as a pair of subapical and a pair of apical spines (GAO et al. 2014). The new taxon is also similar to some species of the genus Chudania Distant (especially to Ch. guangxiana Wu Dai & Yalin Zhang) in the general structure of the aedeagus, including the absence of an atrium and a division of the shaft into two angularly connected parts (DAI & ZHANG 2005). Other features of diagnostic value include the fact that the body is less dorsoventrally flattened and the median carina is completely missing on the frontoclypeus, which is more elevated than in most Nirvaninae (especially in the lateral view); also, lateral grooves on the frontoclypeus run parallelly. Within Nirvanini (particularly the Old World forms), two main groups of species are distinguished by their distinct morphological traits. This is at least partly confirmed by the analyses of DNA sequences from selected representatives of the tribe (Fig. 1). The first group comprises genera with four apical cells on the hindwing and a well-developed marginal vein, generally running close to the outer wing margin and merging with its nodal part. The forewing typically possesses an apical row of transverse veins forming four apical cells of an identical shape and size. Other less significant features include: short and apically hooked styles, small anal plates with rounded apices, relatively short Yshaped connectives (only sporadically with a slightly elongate basal arm), and a pygofer usually with a distinct ventral lobe, often additionally provided with a process. It is noteworthy that the aedeagus has a reduced atrium, is often irregularly shaped and is species specific. The degree of body and head flattening, as well as the length of the median carina on frontoclypeus are variable, even in the individual genera of these and other Nirvanini. The group comprises among others, species of the Kana Distant, Chudania Distant and Australnirvana Wang, Dietrich & Zhang genera, which on the diagram of genetic sequences form a separate subclade related to the species of the Pagaronini and Evacanthini tribes (Fig. 1).
The other morphologically distinct group of Nirvanini includes the genera with three apical cells on the hindwing and a marginal vein somewhat shifted from the outer margin and merged with the radial vein, which results in the formation of a large membranous lobe in the apical part. On the forewing, four apical cells are usually varied in shape and size, and the fourth cell (the inner one) is distinctly longer than the remaining ones. Other characteristics are more diverse: styles with a considerably elongate apex, longer and often tapering genital plates (e.g. Nirvana Kirkaldy), T-shaped connectives with an elongate basal arm, and a pygofer lacking clearly emarginated lobes but endowed with ventral processes. The aedeagus is typical of most Evacanthinae, with a fully developed atrium and often long processes. This group contains the species of the Sophonia Walker and Nirvana Kirkaldy genera. Genetic research has confirmed the uniformity of the group forming a single separate clade. Accordingly, it seems interesting to define the phylogenetic relationships between the genera Sabahia gen. nov., Decursusnirvana Gao & Zhang and probably Sinonirvana Gao & Zhang with those of other Nirvanini. They form a distinctive and separate group, which is morphologically similar to the genera Kana Distant, Chudania Distant and Australnirvana Wang, Dietrich & Zhang, but is genetically it is closer to the genera Sophonia Walker and Nirvana Kirkaldy.
The analysis of the genetic relationships of selected species within Nirvanini, Evacanthini and Pagaronini in the subfamily Evacanthinae confirmed most of the previous results (DIETRICH 2004;WANG et al. 2017). Close relationships between the Evacanthini, Nirvanini and Pagarononi tribes, and especially close genetic relations between Evacanthini and Pagaronini were indicated (Fig. 1). Also, the paraphyletic character of the genus Sophonia Walker (Dietrich 2004) was proven to be valid. Similarly to previous reports (WANG et al. 2017), a group of relationships was revealed which includes the genera Kasunga Linnavuori, Kosasia Distant, Sophonia Walker (some species) and Ophiuchus Distant, with Sophonia Walker (some species), Oniella Matsumura and Convexfronta Li being most closely related. On the other hand, a close relationship between the genera Nirvana Kirkaldy and Tortor Kirkaldy was not confirmed (Fig. 1). The results attest to the phyletic distinctiveness of the species within the Narecho Jacobi genus, which is closer to the Evacanthini tribe (WANG et al. 2017 (Fig.1).
However, the presented results must be further supplemented before a decision on the mono-or paraphyletic character of the subfamily Evacanthinae within the Nirvanini tribe is made (DIETRICH 2004;WANG et al. 2017).
Finally, we would like to add some comments regarding the biotope in which the insects were found. Figure 9 shows a photo of the biotope. The specimens were collected from ferns, which are also visible in the photo (Fig. 9). Professor Adam ROSTAÑSKI (Botany Team, Institute of Biology, Biotechnology and Environmental Protection, University of Silesia in Katowice) identified these ferns covering the bottom of the rainforest as belonging to the Nephrolepidaceae family, most likely Nephrolepis brownii (Desv.) Hovenkamp & Miyam or Nephrolepis biserrata (Sw.) Schott. based on the photos. The paper by JAMAN et al. (1999) was helpful in determining the fern species. Fig. 9. The biotope of Sabahia polypodii gen. et sp. nov