Systematics of the new genus Spinosuncus Chen, Zhang & Li with descriptions of four new species (Lepidoptera, Crambidae, Pyraustinae)

Abstract The new genus Spinosuncusgen. n. is proposed for three known species, S.contractalis (Warren, 1896), comb. n., S.praepandalis (Snellen, 1890), comb. n., and S.aureolalis (Lederer, 1863), comb. n. and four new species, S.rectacutussp. n., S.brevacutussp. n., S.curvisetaceussp. n., and S.quadracutussp. n. from the Oriental Region. An identification key is provided for all species. The habiti and genitalia of all species are figured. The monophyly of the genus is well supported by a phylogenetic analysis based on sequence data of the COI, 16S rRNA, and EF-1α genes. The potential sister groups of the new genus, the interspecific relationships and some intraspecific variations within the genus are discussed.


Introduction
Pyraustinae is the third largest subfamily in the family Crambidae, containing 173 genera that include more than 1176 described species (Nuss et al. 2003(Nuss et al. -2018. The monophyly of the Pyraustinae is well supported by phylogenetic analyses based on both morphological characters and molecular data (Solis and Maes 2002, Regier et al. 2013, Yamanaka et al. 2013, Scholtens and Solis 2015 and type specimens deposited in the Natural History Museum, London, the Zoological Institute, Academy of Sciences of Russia, St. Petersburg, the Australian National Insect Collection and the National Museum of Natural History Grigore Antipa, Bucharest, Romania, our efforts of placing these species in a suitable genus were unsuccessful. Moreover, they can't be placed in any African pyraustine genus (Dr Koen VN Maes, pers. comm.). The seven species treated here, currently with no appropriate generic placement, could be easily separated from other pyraustine taxa by several genital traits in both males and females, especially the peculiar uncus, for which the erection of a new genus is considered warranted.
Thus, the aim of this study is to propose a new genus, provide several synapomorphic characters, present an identification key based on external features and genitalia, redescribe three known species, and describe four new ones. A preliminary phylogenetic analysis of the genus and of several potentially related genera, is also proposed based on molecular data.

Molecular material and methods
All species of the genus Spinosuncus, two species of the genus Pseudopagyda, and four species of other genera of Pyraustinae were included for molecular phylogenetic analysis (Table 1). Pseudebulea fentoni Butler, 1881 was chosen as outgroup because it was considered as a basal lineage of the Pyraustinae (Zhang 2003).
Total DNA was extracted from one hindleg and one midleg of 24 specimens using the TIANGEN DNA extraction kit following the manufacturer's instructions. The nucleotide sequences of two mitochondrial genes, cytochrome c oxidase subunit I (COI) and 16S ribosomal RNA (16S rRNA), and one nuclear gene, elongation factor-1 alpha (EF-1α), were selected for study. Primers used in this study were chosen according to Simon et al. (2006), Wahlberg and Wheat (2008) and Hundsdörfer et al. (2009). PCR cycle conditions were an initial denaturation of 5 min at 95 °C, 30 s at 94 °C, 30 s at 48 °C (COI and 16S rRNA) or 51 °C(EF-1α), and 1 min at 72 °C for 35 cycles, and a final extension at 72 °C for 10 min. All amplifications were confirmed by gel electrophoresis on a 1.5% W/V agarose gel in TAE buffer. PCR products were directsequenced at Majorbio Bio-pharm Technology Co., Ltd (Guangzhou), utilizing the same primers used for PCR amplification.
The sequences were aligned using Clustal W (Thompson et al. 1994) under default settings. Gaps were treated as missing data in all analyses. Phylogenetic analyses were conducted using Bayesian inference (BI) method and Maximum likelihood (ML). The BI analysis was run in MrBayes 3.2.6 (Ronquist et al. 2012) with independent parameters for the gene partitions for COI and 16S rRNA under the GTR+G model and for the EF-1α gene partition under the SYM+I+G model as suggested by jModelTest 0.1.1  (Posada 2008). Two independent runs, each with four Markov Chain Monte Carlo (MCMC) simulations, were performed for 3 million generations sampled every 1000 th step. The first 25% of the trees were discarded as burn-in, and posterior probabilities (PP) were determined from remaining trees. The ML analysis was executed in RAxML 8.2.10 (Stamatakis 2014) for all gene partitions under the GTR + G model proposed by jModelTest 0.1.1 (Posada 2008) and with 1000 iterations for bootstrap test. The pairwise Kimura 2-Parameter (K2P) distances between species were calculated from the COI gene using MEGA 6 (Tamura et al. 2013).

Morphological materials and methods
The specimens studied, including the types of the newly described species, are all deposited at the Museum of Biology, Sun Yat-sen University, Guangzhou (SYSBM) except those specified as being in the Insect Collection of the College of Life Sciences, Nankai University (NKU), the Natural History Museum, London, United Kingdom (NHMUK) and the Forest Canopy Ecology Lab, Yunnan (FCEL). Slides of genitalic dissections were prepared according to Robinson (1976) and Li and Zheng (1996), with some modifica-tions. Genitalia terms follow Klots (1970), Munroe (1976a), Maes (1995), and Kristensen (2003). Specimen images at different focal levels were made using a Canon EOS 1DX camera (provided with a Canon 100 mm macro lens) in combination with Helicon Remote. Genitalia pictures were taken using a Zeiss Axio Scope.A1 in combination with a Zeiss AxioCam camera and the Axio Vision SE64 program on a Windows PC; source images were then aligned and stacked on Helicon Focus to obtain a fully sharpened composite image. All the pictures were edited using Adobe Photoshop CS5.
Since the monophyly of Spinosuncus is well-supported and species within the clade are morphologically and genetically distinct from the potential sister groups, a new genus is proposed. The taxonomic details are provided below.

Taxonomy
Spinosuncus gen. n. http://zoobank.org/AF399C02-2BDC-48D6-9A57-384D3DD6F5AD Type species. Paliga contractalis Warren, 1896 Diagnosis. Species of Spinosuncus can be recognized externally by the yellow to fulvous wing ground colour, the fulvous to brown lines, the distinct subterminal Table 2. Pairwise distances of the COI barcode region based on Kimura-2-parameter model (intraspecific distances are highlighted in bold). lines usually arched to CuA 2 then obviously angled or concave near the tornus. Diagnostic characters in the male genitalia are the short and stout, strongly sclerotized uncus distally with two spines or teeth, the lamellate, distally inflated sella set with fin-shaped setae forming editum, the dorsally inflated sacculus with the dorsal margin sclerotized and usually spinulose, the distally broad and usually spinulose phallus, and the spine-like cornuti appear funnel-shaped in the distal end of the vesica. The female genitalia are characterized by the strongly sclerotized lamella postvaginalis always extended dorsolaterally, and the sclerotized transverse band posteriorly in the cup-shaped antrum.
Spinosuncus moths are most similar in appearance to Pseudopagyda Slamka, 2013. Some species of Spinosuncus can be distinguished by the much smaller wingspan (usually less than 24 mm). However, some Spinosuncus species have a similar body size to Pseudopagyda, but they can still be differentiated by the wavy or dentate lines on the wings dorsally, especially the sinuate (rather than oblique, or slightly curved as in Pseudopagyda) anterior part of the postmedial line near the costa. In the male genitalia, the sclerotized uncus, the fin-shaped setae (editum) of the sella, and the inflated sacculus distinguish Spinosuncus from Pseudopagyda. In the female genitalia, the long and slender ductus bursae is distinct from the extremely short ductus bursae of Pseudopagyda. Description. Head. Frons oblique, yellowish brown, with white lateral bands. Vertex with moderately raised scales projecting between antennae. Labial palpus obliquely upturned, exceeding frons by 2/3 length of head or slightly less, third palpomere porrect, yellowish brown with base contrastingly white. Maxillary palpus small, yellowish brown, tips pale yellow, sometimes mixed with white. Proboscis well developed, with basal scaling white. Antenna pale yellow, with cilia as long as width of corresponding flagellomeres in male. Thorax. With appressed scales, yellow. Legs unmodified. Foreleg brown, tibia white with brown cross band medially, tarsus white; midleg pale brown, tibia and tarsus white ventrally; hind leg pale yellow, tinged with white, basal inner spur longer than apical inner spurs. Forewing subtriangular, termen gently arched; retinaculum a tuft of curved bristles from below base of discal cell. Hindwing fan-shaped, costal margin translucent whitish; frenulum simple in male, with two acanthae in female. Wing venation ( Figure 2) in forewing with cell about half length of wing; R 1 free, from 4/5 of anterior margin of cell, R 2 free but adjacent to stem of R 3 +R 4 in about basal half, R 3 and R 4 stalked to about 2/3, R 4 to just before apex, R 5 parallel to stalked R 3 +R 4 at base then diverging; M 1 moderately close to R 5 at base, M 2 widely separate from M 1 , closing vein concavely curved; M 2 , M 3 and CuA 1 from posterior angle of cell, M 3 closer to M 2 at base than to CuA 1 , then diverging; CuA 2 from 3/5 of posterior margin of cell; 1A faintly sinuate to tornus, 2A forming complete loop and distally recurved before joining 1A, sometimes disconnected. Hindwing with cell about 1/3 length of wing; Sc+R 1 and Rs anastomosing for 1/3 beyond end of discal cell, Rs and M 1 short-stalked, closing vein concave, angled medially; M 3 closer to M 2 at base than to CuA 1 , parallel with M 2 at base, then diverging; 1A complete but weak, 3A curved. Abdomen. Slender, usually yellowish, sometimes dark brown, apical margin of segments usually tinged with white. Male genitalia. Uncus short and stout, nearly quadrate, with wide base; usually strongly sclerotized; distal end with two or four sharp spines laterally or distally bifid forming two teeth; glabrous or ventrolaterally set with few setae, or densely setose at base of teeth. Tegumen quadrate. Vinculum U-shaped. Saccus short, near triangular, rounded at apex. Valva tongue-shaped, varying in width, tapering towards apex, set with hair-like setae on inner side; transtilla sub-triangular, meeting in middle, usually with setae on dorsal margin; costa simple, costal sclerotized band narrow to broad, extended to beyond 2/3 of dorsal margin; sacculus broad, expanded except basal part, with dorsal margin strongly sclerotized and often spinose; sella slender to broad, lamellate, distally inflated, set with modified setae (editum), varying from fin-shaped to thick, needle-shaped. Juxta heart-shaped to nearly pentagonal. Phallus with distal part broad and moderately setose, usually spinulose; vesica in distal part with numerous spine-like cornuti appear funnel-shaped, sometimes with several large spicules. Female genitalia. Ovipositor lobes flat, densely setose. Sinus vaginalis well developed, membranous, usually with sclerotized, streak-like or hook-like notches anterolaterally (absent in S. praepandalis and S. curvisetaceus); lamella postvaginalis band-shaped, sclerotized (weakly sclerotized in S. contractalis, S. rectacutus and S. brevacutus), always extended dorsolaterally. Antrum membranous or sclerotized and granulated, cup-shaped, with sclerotized transverse band posteriorly. Ductus seminalis originating from anterior end of colliculum. Ductus bursae with base slightly rotated, as long as or longer than length or diameter of corpus bursae; colliculum ring-shaped, sclerotized. Corpus bursae dropshaped or globular; accessory bursa present, sometimes with second signum at base; main signum rhomboid.
Biology. All of the Chinese material has been collected during the night at light. Host information is currently unavailable. Spinosuncus aureolalis and S. contractalis occur sympatrically with species of Pseudopagyda in some places. According to Bänziger (1995), they are not lachryphagous.
Distribution. Spinosuncus occurs in South China ( Figure 28), India, and Thailand. Etymology. The generic name is a compound word that refers to the uncus distally with spines ("spinosus" in Latin). The resultant name is masculine in gender. (

Diagnosis.
Within the genus, S. contractalis resembles S. rectacutus and S. brevacutus in the relatively small wingspan, the almost indistinguishable wing pattern, the glabrous uncus, a row of dense setae on the transtilla dorsally, the two sclerotized notches anterolaterally on the sinus vaginalis and the short ductus bursae (approximately as long as the length of the corpus bursae). However, it can be differentiated from S. rectacutus by the somewhat more sinuate postmedial line of the forewing near costa, in the male genitalia by the shorter, excurved spines of the uncus and the acinaciform, densely spinous extension of the sacculus distally. In the female genitalia, it is characterized by the curved sclerotized notches anterolaterally on the sinus vaginalis. The differences between S. contractalis and S. brevacutus are given in the diagnosis of the latter species.
Redescription. Head. As for the genus. Thorax. Yellow. Legs as described for the genus. Wingspan 18-22 mm. Wings yellow, lines fulvous. Forewing broadly triangular with moderately arched termen; antemedial line weakly sinuate from about 1/4 of costa to 2/5 of posterior margin; orbicular stigma small, sometimes faint; reniform stigma a fulvous, slightly curved streak; posterior angle of cell outwardly followed by a fulvous mark; postmedial line from 3/5 of costa slightly sinuate to beyond basal half of CuA 1 , bent inwardly to 1/3 of CuA 2 , then to 2/3 of posterior margin; subterminal line from distal end of R 2 , arched to about 4/5 of CuA 2 , then concave to 4/5 of posterior margin; fringe yellowish brown. Hindwing with costa and posterior margin translucent whitish; posterior angle of cell outwardly followed by a fulvous mark; postmedial line straight from basal half of M 1 to distal third of CuA 2 , bent inwardly to basal third of CuA 2 , then straight to near end of 2A; subterminal line from distal third of R S , arched, tapering to CuA 2 , then concave to distal end of 1A; fringe as in forewing. Abdomen. Yellow dorsally, apical margin of segments tinged with white. Male genitalia ( Figure 10). Uncus with lateral margin strongly bulging near distal end, with a sharply widened base; without setae; with two outwardly curved, pointed spines, weakly dentate between the spines. Valva of medium width, slightly narrowing towards apex, length approximately 2× its maximal width; transtilla dorsally strongly sclerotized and set with dense setae; costal sclerotized band narrow, slightly expanded to 2/3 of dorsal margin; sacculus with distal half expanded, forming acinaciform sclerotized process, dorsally set with dense spines; sella long and slender, rod-like, distal end strongly inflated, set with several narrow, fin-shaped setae forming editum, each seta with apex evenly divided into several filaments. Juxta heart-shaped, deeply divided distally. Phallus with distal 1/3 expanded and spinulose; vesica in distal part with numerous spine-like cornuti appear funnel-shaped ( Figure 10C). Female genitalia (Figure 19). Posterior apophysis with small expansion at basal third; anterior apophysis with small expansion beyond basal half. Sinus vaginalis with two curved, sclerotized notches anterolaterally; lamella postvaginalis weakly sclerotized medially, most strongly sclerotized dorsolaterally. Antrum membranous. Ductus bursae moderately broad, nearly as long as length of corpus bursae; colliculum narrow medially. Corpus bursae approximatively drop-shaped; accessory bursa arising from posterior 1/3 of corpus bursae, with small, densely spinulose second signum beside its base; rhombic signum with two opposing angles bearing well developed carinae and closely separated medially, the other two angles bearing dense spines.
Distribution.( Figure 28). China (Hainan, Yunnan, Tibet), India, Thailand. Diagnosis. Spinosuncus rectacutus resembles S. contractalis and S. brevacutus, for which details are provided in the diagnosis of S. contractalis. It can be best distinguished from S. brevacutus by the dorsally densely setose transtilla (moderately setose in S. brevacutus), and the saddle-shaped sacculus with sclerotized margin densely set with a row of spinules. The distal spines of the uncus are straight and longer than those of S. brevacutus, and the lateral margin near the distal end of the uncus is less bulging. In the female genitalia, the length of the colliculum is approximately 1.5× as long as its minimal width and the notches on the sinus vaginalis are strongly sclerotized whereas in S. brevacutus, the length of the colliculum is approximately as long as its minimal width and the notches on the sinus vaginalis are weakly sclerotized.
Description. Head. As for the genus. Thorax. Yellow. Legs as described for the genus. Wingspan 18-22.5 mm. Wing pattern as in S. contractalis. Abdomen. Yellow dorsally, apical margin of segments tinged with white. Male genitalia (Figure 11). Uncus with lateral margin slightly bulging near distal end, with base sharply widened; setae absent; distal two corners with straight, pointed spines, outer margin between spines dentate. Valva of medium width, length approximately 2.3× its maximal width; transtilla with dorsal margin strongly sclerotized and set with dense setae; costal sclerotized band wide, slightly expanded to 2/3 of dorsal margin; distal half of sacculus expanded to a saddle-shaped structure, with sclerotized margin, basal half of margin slightly twisted, set with dense spines; sella long and slender, rod-like, distal end slightly inflated, set with several narrow, fin-shaped setae forming editum, each seta with apex evenly divided into several filaments. Juxta heart-shaped. Phallus as in S. contractalis. Female genitalia (Figure 20). Posterior apophysis with small expansion at basal third; anterior apophysis with small expansion beyond basal half. Sinus vaginalis with two straight, sclerotized notches anterolaterally; lamella postvaginalis weakly sclerotized medially, most strongly sclerotized dorsolaterally. Antrum membranous. Ductus bursae slender, nearly as long as length of corpus bursae; colliculum narrow medially. Corpus bursae drop-shaped, slightly spinulose; accessory bursa arising from posterior 1/3 of corpus bursae, with small, densely spinulose second signum beside its base; rhombic signum with carinae almost connected.
Etymology. The specific name is derived from the Latin recti-for straight and acutus, pointed, referring to the straight, pointed spines of the uncus.
Distribution.( Figure 28). China (Hubei, Guangxi, Guizhou, Chongqing). Diagnosis. Spinosuncus brevacutus is similar to S. contractalis and S. rectacutus. Differences with S. rectacutus are given in the diagnosis of S. rectacutus. It can be distinguished from S. contractalis by the minute and weakly outwardly curved spines of the apical uncus, the concave margin between those spines, the moderately setose transtilla and the semicircular sacculus distally with sclerotized, sparsely toothed margin in the male genitalia, by the straight, weakly sclerotized notches of the sinus vaginalis (curved, strongly sclerotized in S. contractalis) and the relatively broad ductus bursae in the female genitalia.

Spinosuncus brevacutus
Description. Head. As for the genus. Thorax. Yellow. Legs as described for the genus. Wingspan 19-24 mm. Wing pattern as in S. contractalis. Abdomen. Yellow dorsally, apical margin of segments tinged with white. Male genitalia (Figure 12). Uncus with the lateral margin strongly bulging near distal end, with base sharply widened; setae absent; distal two corners slightly extended, forming minute spines. Valva of medium width, length approximately 2.5× its maximal width; transtilla with dorsal margin slightly sclerotized, set with few setae; costal sclerotized band rather wide, slightly expanded to 2/3 of dorsal margin; distal half of sacculus expanded, semicircular, with strongly sclerotized margin, sometimes set with few tiny teeth, distal third of margin twisted; sella long and slender, rod-like, distal end slightly inflated and upcurved, set with several narrow, fin-shaped setae forming editum, each seta with apex evenly divided into several filaments. Juxta heart-shaped, distal half divided. Phallus as in S. contractalis. Female genitalia (Figure 21). Posterior apophysis with small expansion at basal third; anterior apophysis with small expansion beyond basal half. Sinus vaginalis with two straight, weakly sclerotized notches anterolaterally; lamella postvaginalis weakly sclerotized medially, most strongly sclerotized dorsolaterally. Antrum membranous. Ductus bursae moderately broad, as long as length of corpus bursae; colliculum somewhat constricted medially. Corpus bursae drop-shaped; accessory bursa arising from posterior 1/3 of corpus bursae, with small, weakly spinulose second signum beside its base; rhombic signum with two opposing angles bearing weak, narrow carinae almost connected medially, the other two angles set with spines.
Etymology. The specific name is derived from the Latin brevi-, short, and acutus for pointed, referring to the short, pointed spines of the uncus.
Distribution. (Figure 28). China (Jiangxi, Hunan, Guizhou).  Diagnosis. Spinosuncus praepandalis has a larger wingspan (24-30 mm) than in the species described above. It has a wingspan similar to that of S. aureolalis, but can be differentiated by the dentate lines and the thickened anterior part of the postmedial line of the forewing near the costa. In the male genitalia, it is distinguished by the distally bifid uncus, forming two sclerotized, large outwardly curved teeth with a hairy basal margin (as in S. curvisetaceus), the two to three straight, thick needle-shaped setae dorsally set on each side of the transtilla and the semicircular sacculus distally with the margin scle-rotized and with a small process distally. In the female genitalia, it is distinguished by the sinus vaginalis without sclerotized, streak-like or hook-like notches (as in S. curvisetaceus) and the long and slender ductus bursae, which is more than twice as long as the diameter of the corpus bursae, differs from that of the species described above (the ductus bursae is almost as long as the length of the corpus bursae). The differences between S. praepandalis and S. curvisetaceus are given in the diagnosis of the latter species.
Redescription. Head. As for the genus. Thorax. Yellow. Legs as described for the genus. Wingspan 24-30 mm. Wing pattern as in S. contractalis, apart from: wings yellowish brown; lines brown and wavy; postmedial line of forewing thickened near costa, strongly sinuate to half of CuA 1 ; postmedial line of hindwing curved to distal third of CuA 2 . Abdomen. Yellowish to brown, apical margin of segments tinged with white. Male genitalia ( Figure 13). Uncus tapering towards apex; distal 3/4 bifid, forming two outwardly curved, strongly sclerotized teeth, medially set with dense setae, arranged in a curved line. Valva of medium width, ventral margin beyond sacculus slightly concave, length approximately 2.3× its maximal width; transtilla extended ventrally into a projection, each lobe set with two to three straight, thick needle-shaped setae at dorsal base (one seta occasionally falls off), with one much bigger than other(s); costal band moderately wide, slightly expanded to 2/3 of dorsal margin; distal half of sacculus expanded, semicircular, with dorsal margin sclerotized, apically with small, triangular process; sella long and slender, rod-like, upcurved (bent in Figs 13A-B), distal end slightly inflated, set with a few broad, fin-shaped setae forming editum, each seta with apex evenly divided into several filaments. Juxta pentagonal, weakly bifid distally. Phallus with distal 1/4 slightly expanded and spinulose; vesica in distal part with numerous spine-like cornuti appear funnel-shaped ( Figure 13C). Female genitalia (Figure 22). Posterior apophysis with distinct hook-like expansion at basal 2/5. Sinus vaginalis without sclerotized, streak-like or hook-like notches; lamella postvaginalis band-shaped, well developed, extended to cover entire eighth segment ventrally. Antrum membranous, with a narrow sclerotized transverse band posteriorly. Ductus bursae long and slender, more than three times as long as diameter of corpus bursae; colliculum almost evenly wide. Corpus bursae small, globular; accessory bursa arising from posterior 1/3 of corpus bursae; rhombic signum with well developed, moderately separated carinae, other two angles bearing spines medially, the anterior angle smaller than the posterior angle; second signum absent.

Spinosuncus curvisetaceus
Description. Head. As for the genus. Thorax. Yellowish brown. Legs as described for the genus. Wingspan 24-26 mm. Wing pattern as in S. praepandalis, ground colour paler than that of S. praepandalis. Abdomen. Yellowish to brown, apical margin of segments tinged with white. Male genitalia (Figure 14). Uncus sharply tapering towards apex; distal half bifid, forming two slightly outwardly curved and sclerotized teeth, basally set with dense setae, arranged in a curved line. Valva of medium width, ventral margin beyond sacculus slightly concave, length approximately 2.1× its maximal width; transtilla extended ventrally into long and curved projection, set with two thick and curved, needle-shaped setae at base dorsally; costal sclerotized band moderately wide, slightly expanded to 3/4 of dorsal margin; distal half of sacculus expanded, rectangular, with dorsal margin strongly sclerotized and densely spinulose, distally twisted; sella long and slender, rod-like, upcurved, distally set with few broad, fin-shaped setae, each seta with apex evenly divided into several filaments. Juxta shield-shaped, distal half divided medially. Phallus as in S. praepandalis. Female genitalia (Figure 23). Posterior apophysis with hook-like expansion at basal 2/5. Sinus vaginalis without sclerotized, streak-like or hook-like notches; lamella postvaginalis band-shaped, well developed, extended to cover entire eighth segment ventrally. Antrum membranous. Ductus bursae long and slender, more than two times as long as length of corpus bursae; colliculum narrower at anterior end. Corpus bursae small, ovoid; accessory bursa arising from posterior 1/3 of corpus bursae; rhombic signum as in S. praepandalis; second signum absent.
Etymology. The specific name is derived from the Latin curv-(curved) and setaceus (setaceous), referring to the curved setae set at the dorsal base of the transtilla.
Distribution. (Figure 28). China (Fujian, Jiangxi, Guangxi). (Lederer, 1863) Diagnosis. Spinosuncus aureolalis has a large wingspan (more than 26 mm). The ground colour of the wings is the darkest within the genus. Though S. aureolalis has a similar wingspan as S. praepandalis, it can be distinguished by the sinuate but not thickened anterior part of the postmedial line of the forewing near costa and the smooth, not dentate wing lines. In the male genitalia, it is characterized by the uncus distally with two large spines, the cheliform sacculus projections, and the finand needle-shaped setae forming editum on the sella distally (as in S. quadracutus). In the female genitalia, the two large, hook-like notches anterolaterally on the sinus vaginalis and the laterally broad, granulated antrum (as in S. quadracutus) are diagnostic. The appearance of S. aureolalis is most similar to that of S. quadracutus, both having the same wing pattern. The differences between these two species are given in the diagnosis of S. quadracutus.

Spinosuncus aureolalis
Redescription. Head. As for the genus. Thorax. Yellow. Legs as described for the genus. Wingspan 26-32 mm. Wings yellow, with fulvous tinge, lines fulvous to yellowish brown, venation somewhat darker than the ground colour, making wings impressively reticulated. Wing pattern as in S. contractalis, apart from: postmedial line of forewing more sinuate, of hindwing more curve. Abdomen. Fulvous dorsally, apical margin of segments tinged with white. Male genitalia (Figs 15, 16). Uncus gradually tapering from base to middle; laterally membranous and set with several se- tae ventrally, other areas strongly sclerotized; distal 1/3 divided into two sharp teeth, thick, straight or slightly curved (weakly folded in Figs 15A, 16A), between two teeth usually two small and short spines ( Figure 16A), sometimes invisible ( Figure 15A) (longish, distinct in S. quadracutus, Figs 17A, 18A); with two caniniform teeth medioventrally. Valva narrow, length approximately 2.7× its maximal width; transtilla extended ventrally into long and narrow projection, dorsal margin with sparse setae; costal sclerotized band rather narrow, extended to near distal end of valva; sacculus with median caniniform projection and distal cheliform projection, distal half set with dense setae ventrally, distal projection with dorsal margin strongly sclerotized, set with dense and flat-lying spines (except distal half, Figs 15C, 16C) and two moderately downcurved spines pointing towards juxta (sometimes the longer one absent, Figure 16C); sella short and broad, distally inflated, set with modified setae forming editum, varying from fin-shaped to thick needle-shaped, ventral margin upcurved, thickened and sclerotized, distally spinose, ended in long, curved spine. Juxta shieldshaped, pentagonal, distal margin sometimes slightly indented medially. Phallus with distal 1/4 slightly expanded, vesica distally with numerous spinules and several large spicules arranged into funnel-shaped bunch of cornuti (Figs 15D, 16D, rotated in Figure 16D). Female genitalia (Figs 24, 25). Anterior apophysis sclerotized, slightly sinuate at distal third; posterior apophysis oblong, slender, strongly sclerotized. Sinus vaginalis with two large, thick, hook-like notches anterolaterally; lamella postvaginalis sclerotized, band-shaped, extended dorsolaterally to about 1/4 width of sinus vaginalis. Antrum granulated and broad. Ductus bursae long and wide, about two times as long as diameter of corpus bursae; colliculum well-developed, with anterior end narrower. Corpus bursae globular; accessory bursa arising from posterior end of corpus bursae; rhombic signum with carinae weak and widely separated, other two angles bearing dense spines; second signum absent.
Description. Head. Frons brown, vertex with moderately raised scales projecting between antennae, labial palpus brown, white at base ventrally. Maxillary palpus brown, with apex pale yellow. Thorax. Yellow. Legs as described for the genus. Wingspan 26-30 mm. Wing pattern as in S. aureolalis. Abdomen. Fulvous dorsally, apical margin of segments tinged with white. Male genitalia (Figs 17, 18). Uncus tapering from base to middle; laterally membranous and set with several setae ventrally; otherwise strongly sclerotized; with two caniniform teeth medioventrally; distally with four sharp and slender spines, the lateral two longer, about two times as long as the median two. Valva narrow, as in S. aureolalis; transtilla extended ventrally into a long and narrow projection, dorsal margin with sparse setae; costal sclerotized band rather narrow, extended to near distal end of valva; sacculus with central caniniform projection and distal cheliform projection, distal half set with dense setae ventrally, distal projection strongly sclerotized, set with dense and slightly raised spines and one moderately downcurved spine pointing towards juxta; sella short and broad, distally inflated, set with modified setae forming editum, varying form fin-shaped to thick needle-shaped, ventral margin upcurved, thickened and sclerotized, distally spinose, ending in long, curved spine. Juxta shield-shaped, pentagonal, distal margin slightly bifid. Phallus as in S. aureolalis. Female genitalia (Figs 26, 27). Anterior apophysis sclerotized, slightly sin-  vaginalis. Antrum granulated and broad. Ductus bursae long and moderately wide, about two times as long as diameter of corpus bursae; colliculum well-developed, with anterior end narrower. Corpus bursae globular; accessory bursa arising from posterior end of corpus bursae; rhombic signum with carinae well-developed and connected ( Figure 26A) or weak and wide separated ( Figure 27A), other two angles densely bearing spines, sometimes smooth medially ( Figure 27A); second signum absent.
Etymology. The specific name is derived from the Latin quadri-(four) and acutus (pointed), referring to the distal uncus with four pointed spines.

Discussion
The results of the molecular analysis robustly support the monophyly of Spinosuncus. The monophyly of the genus is further supported morphologically by the following potential synapomorphies: the sclerotized uncus distally with two spines or teeth, the lamellate, distally inflated sella with fin-shaped setae forming editum, the dorsally expanded sacculus with the dorsal margin sclerotized, the distally expanded, spinulose phallus, the funnel-shaped bunch of cornuti, the sclerotized lamella postvaginalis always extended dorsolaterally and the sclerotized transverse band at the posterior end of the antrum. According to the tree topology (Figure 1), Paratalanta is not so closely related to Spinosuncus, and as already discussed by Zhang et al. (2014), species of Spinosuncus share no synapomorphy with Paratalanta. The relationship between Pseudopagyda + Aglaops and Spinosuncus is well supported (PP = 1.00, BS = 77), but the clade Pseudopagyda + Aglaops is only weakly supported (PP = 0.81, BS = 50). Morphologically, Aglaops species are similar to Pseudopagyda in the campanulate uncus and the distally inflated sella (which is also similar in Spinosuncus), but other genitalia traits differ. The wing pattern of Aglaops species lacks a subterminal line, whereas Pseudopagyda and Spinosuncus species have such a line. Pseudopagyda species resemble Spinosuncus species in the wing pattern. In the male genitalia, the distally inflated sella is similar to that of Spinosuncus; the needle-shaped setae forming editum and the presence of several large spines in the phallus distally are also found in S. aureolalis and S. quadracutus; the heart-shaped juxta is similar in S. contractalis, S. rectacutus, S. brevacutus, S. praepandalis and S. curvisetaceus. Based on the molecular phylogenetic results and morphological characters, Pseudopagyda could be the most closely related genus to Spinosuncus.
Other genera included in the molecular analysis, represented by Ostrinia furnacalis (Guenée, 1854), Placosaris rubellalis (Caradja, 1925), Thliptoceras sinense (Caradja, 1925) and Aglaops youboialis (Munroe & Mutuura, 1968), all lack a forewing subterminal line. Placosaris rubellalis and Thliptoceras sinense have a rod-like sella similar to that of Spinosuncus in the male genitalia, but the editum are different. Ostrinia furnacalis has a weakly sclerotized uncus, distally divided into three small, laterally setose processes, which is somewhat similar in some Spinosuncus species. However, other traits of the male genitalia of O. furnacalis are quite different from those of Spinosuncus species. At present, it is impossible to confirm the generic position of Spinosuncus within the subfamily since only few genera of Pyraustinae were included in this study.
Taxonomically, Spinosuncus can be divided into three species groups: the contractalis group, the praepandalis group and the aureolalis group. The monophyly of these three species groups is well supported by the phylogenetic analysis (Figure 1). The aureolalis group, comprising S. aureolalis and S. quadracutus, is well characterized by the laterally setose uncus, distally with two or four spines, ventrally with two large teeth; the lamellate, distally inflated sella with fin-and needle-shaped setae forming editum; the cheliform sacculus with a long spine pointing towards juxta; and the two thick, hooklike notches anterolaterally on the sinus vaginalis. The contractalis group comprises S. contractalis, S. rectacutus and S. brevacutus. This species group is well defined by several characters: the glabrous, sclerotized uncus distally with two spines; a row of setae on the transtilla dorsally; and two streak-like sclerotized notches anterolaterally on the sinus vaginalis. Within the contractalis group, S. contractalis is closer to S. rectacutus than to S. brevacutus based on the relatively long spines on the uncus distally and the densely setose transtilla dorsally. The praepandalis group, comprising S. praepandalis and S. curvisetaceus, can be recognized by the following characters: the bifid uncus with two basally setose teeth; the two needle-shaped setae on the transtilla dorsally; a long, narrowly triangular lobe projecting from the transtilla ventrally; and the long and slender ductus bursae which is about twice as long as the diameter of the corpus bursae. Within the genus, the praepandalis group is closer to the contractalis group than to the aureolalis group.
In this study, four new species are described based on morphological and genetic differences from related species. The morphological differences are given above in the diagnoses of the new species. The genetic distance between species in Lepidoptera are ordinarily greater than 3% (Hebert et al. 2003) in the COI barcode. Among the new species, S. quadracutus, S. curvisetaceus and S. brevacutus are well recognized by distance values greater than 3% from their most closely related species (Table 2). Another new species, S. rectacutus showed relatively low genetic distance (2.5%-2.7%) to its most closely related species S. contractalis. However, S. rectacutus can be distinguished from S. contractalis as mentioned above under the diagnosis of S. rectacutus and by the key. Moreover, such cases of low genetic divergence are also observed in some other studies in Lepidoptera (Hebert et al. 2003, 2010, Yang et al. 2016. The low interspecific divergence of congeneric species pairs may indicate their recent origin or introgression (Hebert et al. 2003, Zahiri et al. 2014. Based on the covariation between barcodes and morphological traits, S. rectacutus is treated as a distinct species. A relatively high intraspecific divergence was observed in S. aureolalis (2.7%). The two specimens concerned, a male and a female, were collected in two localities in Yunnan that are distant by approximately 550 km ( Figure 28A). According to the genitalia (Figure 24), the female specimen belongs to the aureolalis species group and it can be distinguished from S. quadracutus by the two hook-like notches more widely separated from each other. Moreover, no obvious genital variation could be found in the males found in these two localities. Consequently, they are here treated as conspecific. Genital variation is observed in two male specimens collected in Daxichang, Yunnan and Nonggang, Guangxi (genitalia slides no. SYSU0173, SYSU 0909, respectively), both places which are near the north of Vietnam. The distal projection of the sacculus has only one large spine, as in those of S. quadracutus, whereas those specimens collected in other places of Yunnan, Thailand and India have two spines. However, other genital traits, as given in the redescription, are all uniform, suggesting their recognition as the same species, S. aureolalis.
In the present study, four new species are discovered which are superficially similar to the three described species. Considering the lack of sufficient generic revisions, especially in Oriental region, there is little doubt that many described species have been misplaced and more cryptic species will be revealed within the subfamily. As Munroe (1976a) pointed out, inclusion of genitalia structures and careful analyses of the interspecific and intraspecific differences will certainly help to move ahead to natural classifications as opposed to artificial arrangements. However, the understanding of the phylogenetic relationships between most genera of Pyraustinae is still very imperfect. Phylogenetic systematics based on morphology helps little as pyraustine genera are separated in most cases only by minute morphological differences which are difficult to interpret as apo-or plesiomorphic. The use of genetic data will facilitate species identification and help to understand the interspecific and intergeneric relationships. It calls for more comprehensive investigations on Pyraustinae in the future in order to understand this species-rich subfamily better.