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Systematics of the new genus Spinosuncus Chen, Zhang & Li... 115
Systematics of the new genus Spinosuncus Chen,
Zhang & Li with descriptions of four new species
(Lepidoptera, Crambidae, Pyraustinae)
Kai Chen1, Dandan Zhang1,2, Houhun Li2
1 State Key Laboratory of Biocontrol/e Museum of Biology, School of Life Sciences, Sun Yat-sen University,
Guangzhou, Guangdong 510275, China 2 College of Life Sciences, Nankai University, Tianjin 300071, China
Corresponding authors: Dandan Zhang (zhangdd6@mail.sysu.edu.cn); Houhun Li (lihouhun@nankai.edu.cn)
Academic editor: B. Landry|Received 27 January 2018|Accepted 1 October 2018|Published 28 November 2018
http://zoobank.org/566B1801-1160-4470-9BF1-A6B7A88E802E
Citation: Chen K, Zhang D, Li H (2018) Systematics of the new genus Spinosuncus Chen, Zhang & Li with
descriptions of four new species (Lepidoptera, Crambidae, Pyraustinae). ZooKeys 799: 115–151. https://doi.
org/10.3897/zookeys.799.23925
Abstract
e new genus Spinosuncus gen. 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. rectacutus sp. n., S. brevacutus sp. n., S. curvisetaceus sp. n., and S. quadracutus sp. n. from
the Oriental Region. An identication key is provided for all species. e habiti and genitalia of all species
are gured. e monophyly of the genus is well supported by a phylogenetic analysis based on sequence
data of the COI, 16S rRNA, and EF-1α genes. e potential sister groups of the new genus, the interspe-
cic relationships and some intraspecic variations within the genus are discussed.
Keywords
Aglaops, China, molecular phylogeny, new combinations, Ostrinia, Paratalanta, Placosaris, Pseudebulea,
Pseudopagyda, liptoceras
Introduction
Pyraustinae is the third largest subfamily in the family Crambidae, containing 173
genera that include more than 1176 described species (Nuss et al. 2003–2018). e
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.
ZooKeys 799: 115–151 (2018)
doi: 10.3897/zookeys.799.23925
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Kai Chen et al. / ZooKeys 799: 115–151 (2018)
116
2012). Based on specialized genitalic characters, e.g. the valva bearing sella and editum
in the male genitalia and a rhomboidal signum in the female genitalia, pyraustine spe-
cies are easily distinguished from members of its sister group, Spilomelinae (Regier et
al. 2012). However, the taxa belonging to Pyraustinae sensu stricto still have not been
all associated on a worldwide basis (Solis and Maes 2002), partly because they were for
a long time placed into Pyraustinae sensu lato along with various Spilomelinae. is
group has been relatively well studied in Europe and North America. In other regions,
however, particularly the Oriental Region, this work is far from complete. One of the
major contributions to the knowledge of the Pyraustinae of East Asia is a series of
papers by Munroe and Mutuura (1968, 1969, 1970, 1971) treating many pyraustine
genera of temperate East Asia, a historical milestone in the study of the Pyraustinae of
this area. Both authors’ generic concepts, however, were narrow, so that many genera
recognized by these authors were united by Tränkner et al. (2009) in a wider concept
of the genus Anania Hübner.
In recent years, a series of similar yellowish specimens collected from the south
of China, all supercially resembling species of Pseudopagyda Slamka, 2013, attracted
our attention. By examining the genitalia, three described species, Paliga contractalis
Warren, 1896, Botys aureolalis Lederer, 1863, Botys praepandalis Snellen, 1890 and four
unknown species were recognized. According to characters of male and female genita-
lia, they are congeneric, but obviously do not match the genitalic morphology of Pseu-
dopagyda or Paliga Moore, 1886. Bänziger (1995) placed Paliga contractalis and Botys
aureolalis in genus Microstega Meyrick, 1890 along with Pionea acutangulata Swinhoe,
1901 and Microstega homoculorum Bänziger, 1995. Bänziger (1995) also pointed out
that M. homoculorum and M. acutangulata are congeneric, but that M. contractalis and
M. aureolalis probably each belong to a dierent genus, and that Pionea praepandalis
resembles M. aureolalis supercially, without giving any details. Microstega was syn-
onymized with Paratalanta Meyrick, 1890 by Kirpichnikova (1986) and Maes (1994)
based on “the characteristic sclerotized hook (spicula-shaped sella) on the valvae of
the male genitalia”. is taxonomic decision is commonly accepted, but Zhang et al.
(2014) excluded the above ve species from Paratalanta because they share no generic
synapomorphies with Paratalanta. Almost at the same time, Slamka (2013) proposed
genus Pseudopagyda for M. homoculorum. Subsequently, M. acutangulata was trans-
ferred to Pseudopagyda (Chen and Zhang 2017). Slamka (2013) also suggested that
Paliga contractalis and Botys aureolalis should belong to Pseudopagyda. However, in a re-
vision of Pseudopagyda (Chen and Zhang 2017), several putative synapomorphic char-
acters of the genus were summarized, and it was found that Paliga contractalis and Botys
aureolalis are not congeneric with species of Pseudopagyda based on genitalia characters.
After comparing these species with taxonomic treatments, faunal surveys, and
checklists of Spilomelinae and Pyraustinae (Hampson 1893, 1896, 1898, 1899, Car-
adja 1925, Shibuya 1928, 1929, Munroe and Mutuura 1968, 1969, 1970, 1971,
Munroe 1976a, 1976b, Wang 1980, Inoue 1982, Heppner and Inoue 1992, Munroe
1995, Speidel 1996, Shaer et al. 1996, Kirpichnikova 1999, 2009, Wang and Speidel
2000, Mathew 2006, Shaer and Munroe 2007, Bae et al. 2008, Leraut 2012, Slamka
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 117
2013, Yamanaka et al. 2013, Scholtens and Solis 2015) and type specimens depos-
ited 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
eorts 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.).
e 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.
us, the aim of this study is to propose a new genus, provide several synapomor-
phic characters, present an identication key based on external features and genitalia,
redescribe three known species, and describe four new ones. A preliminary phyloge-
netic analysis of the genus and of several potentially related genera, is also proposed
based on molecular data.
Materials and methods
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 analy-
sis (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. e nu-
cleotide 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 nal extension at 72 °C for 10 min. All amplications were conrmed by gel
electrophoresis on a 1.5% W/V agarose gel in TAE buer. PCR products were direct-
sequenced at Majorbio Bio-pharm Technology Co., Ltd (Guangzhou), utilizing the
same primers used for PCR amplication.
e sequences were aligned using Clustal W (ompson 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). e
BI analysis was run in MrBayes 3.2.6 (Ronquist et al. 2012) with independent param-
eters 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
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
118
Table 1. Species sampled for the molecular phylogenetic analysis.
GenusSpecies Voucher number Locality GenBank accession number Reference
COI 16S EF-1α
Pseudebulea fentoni SYSU-LEP0074 Hunan Prov. MG739570 MG739582 MG739594 Chen et al. 2018
Paratalanta ussurialis SYSU-LEP0158 Hunan Prov. MK000093 MK000070 MK000116 present study
Ostrinia furnacalis SYSU-LEP0157 Jiangxi Prov. MK000094 MK000071 MK000117 present study
Placosaris rubellalis SYSU-LEP0087 Jiangxi Prov. MK000095 MK000072 MK000118 present study
liptoceras sinense SYSU-LEP0080 Jiangxi Prov. MK000096 MK000073 MK000119 present study
Aglaops youboialis SYSU-LEP0068 Jiangxi Prov. MK000097 MK000074 MK000120 present study
Pseudopagyda
homoculorum SYSU-LEP0116 Yunnan Prov. MK000098 MK000075 MK000121 present study
acutangulata SYSU-LEP0011 Jiangxi Prov. MK000099 MK000076 MK000122 present study
acutangulata SYSU-LEP0126 Jiangxi Prov. MK000100 MK000077 MK000123 present study
Spinosuncus
aureolalis SYSU-LEP0132 Yunnan Prov. MK000101 MK000078 MK000124 present study
aureolalis SYSU-LEP0146 Yunnan Prov. MK000102 MK000079 MK000125 present study
quadracutus SYSU-LEP0001 Hainan Prov. MK000103 MK000080 MK000126 present study
quadracutus SYSU-LEP0002 Hainan Prov. MK000104 MK000081 MK000127 present study
curvisetaceus SYSU-LEP0129 Jiangxi Prov. MK000105 MK000082 MK000128 present study
praepandalis SYSU-LEP0006 Guizhou Prov. MK000106 MK000083 MK000129 present study
praepandalis SYSU-LEP0131 Yunnan Prov. MK000107 MK000084 MK000130 present study
brevacutus SYSU-LEP0009 Guizhou Prov. MK000108 MK000085 MK000131 present study
brevacutus SYSU-LEP0010 Guizhou Prov. MK000109 MK000086 MK000132 present study
brevacutus SYSU-LEP0156 Jiangxi Prov. MK000110 MK000087 MK000133 present study
rectacutus SYSU-LEP0134 Guizhou Prov. MK000111 MK000088 MK000134 present study
rectacutus SYSU-LEP0155 Guizhou Prov. MK000112 MK000089 MK000135 present study
contractalis SYSU-LEP0133 Yunnan Prov. MK000113 MK000090 MK000136 present study
contractalis SYSU-LEP0135 Yunnan Prov. MK000114 MK000091 MK000137 present study
contractalis SYSU-LEP0153 Yunnan Prov. MK000115 MK000092 MK000138 present study
(Posada 2008). Two independent runs, each with four Markov Chain Monte Carlo
(MCMC) simulations, were performed for 3 million generations sampled every 1000th
step. e rst 25% of the trees were discarded as burn-in, and posterior probabilities
(PP) were determined from remaining trees. e 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. e
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
e specimens studied, including the types of the newly described species, are all deposit-
ed at the Museum of Biology, Sun Yat-sen University, Guangzhou (SYSBM) except those
specied as being in the Insect Collection of the College of Life Sciences, Nankai Uni-
versity (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 modica-
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 119
tions. Genitalia terms follow Klots (1970), Munroe (1976a), Maes (1995), and Kris-
tensen (2003). Specimen images at dierent 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 com-
posite image. All the pictures were edited using Adobe Photoshop CS5.
Results
Phylogenetic relationships
e concatenated dataset of three genes consisted of 1863 nucleotide positions (658
for COI, 434 for 16S rRNA and 771 for EF-1α, respectively). Pairwise distances of the
barcode region (COI) are given in Table 2. e genetic distances between the genus Spi-
nosuncus (described below) and the other genera range from 9.0% (Aglaops) to 17.0%
(Pseudebulea). Interspecic genetic distances within Spinosuncus range from 2.5% (S.
contractalis to S. rectacutus) to 13.8% (S. aureolalis to S. rectacutus) while intraspecic
genetic distances in Spinosuncus range from 0% (S. contractalis) to 2.7% (S. aureolalis).
e BI and ML analyses of the concatenated dataset inferred congruent topologies
with only subtle dierences in posterior probability and bootstrap values probability
(Figure 1). e monophyly of Spinosuncus is robustly supported (PP = 1.00, BS = 98).
Within Spinosuncus, three well-supported clades are identied. e clade S. aureolalis
+ S. quadracutus, clade S. curvisetaceus + S. praepandalis and clade S. brevacutus + (S.
rectacutus + S. contractalis) are each recovered with robust supports (PP = 1.00, BS =
100). Clade Aglaops youboialis + (Pseudopagyda homoculorum + P. acutangulata) is in a
sister position to clade Spinosuncus with robust support as well (PP = 1.00, BS = 77).
Distances between Spinosuncus and Pseudopagyda range from 10.2% to 13.3%, and
between Spinosuncus and Aglaops from 9.0% to 12.9%.
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. e 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
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
120
Table 2. Pairwise distances of the COI barcode region based on Kimura-2-parameter model (intraspecic distances are highlighted in bold).
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
1LEP0132 Spinosuncus
aureolalis
2LEP0146 Spinosuncus
aureolalis 0.027
3LEP0001 Spinosuncus
quadracutus 0.044 0.037
4LEP0002 Spinosuncus
quadracutus 0.046 0.035 0.002
5LEP0129 Spinosuncus
curvisetaceus 0.111 0.092 0.109 0.107
6LEP0006 Spinosuncus
praepandalis 0.115 0.100 0.107 0.105 0.041
7LEP0131 Spinosuncus
praepandalis 0.117 0.107 0.105 0.107 0.050 0.024
8LEP0009 Spinosuncus
brevacutus 0.129 0.104 0.118 0.116 0.083 0.101 0.109
9LEP0010 Spinosuncus
brevacutus 0.131 0.106 0.120 0.118 0.085 0.103 0.111 0.002
10 LEP0156 Spinosuncus
brevacutus 0.125 0.104 0.114 0.116 0.083 0.101 0.105 0.003 0.005
11 LEP0134 Spinosuncus
rectacutus 0.138 0.112 0.120 0.118 0.094 0.100 0.109 0.049 0.047 0.049
12 LEP0155 Spinosuncus
rectacutus 0.138 0.112 0.120 0.118 0.094 0.100 0.109 0.047 0.046 0.047 0.005
13 LEP0133 Spinosuncus
contractalis 0.131 0.106 0.118 0.116 0.089 0.095 0.099 0.049 0.048 0.049 0.027 0.025
14 LEP0135 Spinosuncus
contractalis 0.127 0.106 0.114 0.112 0.089 0.095 0.095 0.049 0.048 0.049 0.027 0.025 0.003
15 LEP0153 Spinosuncus
contractalis 0.127 0.106 0.114 0.112 0.089 0.095 0.095 0.049 0.048 0.049 0.027 0.025 0.003 0.000
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
16 LEP0068 Aglaops
youboialis 0.129 0.112 0.118 0.116 0.090 0.100 0.104 0.094 0.096 0.094 0.098 0.098 0.100 0.100 0.100
17 LEP0116 Pseudopagyda
homoculorum 0.131 0.124 0.133 0.130 0.102 0.114 0.114 0.124 0.126 0.124 0.118 0.118 0.120 0.116 0.116 0.085
18 LEP0011 Pseudopagyda
acutangulata 0.129 0.122 0.126 0.126 0.108 0.122 0.120 0.124 0.126 0.122 0.130 0.130 0.122 0.118 0.118 0.092 0.058
19 LEP0126 Pseudopagyda
acutangulata 0.129 0.122 0.126 0.126 0.108 0.122 0.120 0.124 0.126 0.122 0.130 0.130 0.122 0.118 0.118 0.092 0.058 0.000
20 LEP0080 liptoceras
sinense 0.149 0.138 0.135 0.133 0.129 0.138 0.134 0.131 0.133 0.131 0.137 0.135 0.131 0.129 0.129 0.096 0.118 0.123 0.123
21 LEP0087 Placosaris
rubellalis 0.133 0.124 0.135 0.137 0.100 0.110 0.119 0.114 0.116 0.110 0.116 0.120 0.116 0.116 0.116 0.087 0.112 0.112 0.112 0.127
22 LEP0157 Ostrinia
furnacalis 0.142 0.124 0.120 0.118 0.126 0.130 0.137 0.116 0.118 0.116 0.133 0.133 0.130 0.130 0.130 0.108 0.114 0.132 0.132 0.120 0.112
23 LEP0158 Paratalanta
ussurialis 0.143 0.126 0.143 0.141 0.122 0.133 0.139 0.122 0.124 0.122 0.124 0.128 0.124 0.124 0.124 0.102 0.120 0.112 0.112 0.135 0.106 0.102
24 LEP0074 Pseudebulea
fentoni 0.148 0.135 0.148 0.146 0.159 0.161 0.159 0.161 0.158 0.161 0.170 0.170 0.161 0.158 0.158 0.158 0.133 0.143 0.143 0.161 0.145 0.145 0.157
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
122
lines usually arched to CuA2 then obviously angled or concave near the tornus. Di-
agnostic characters in the male genitalia are the short and stout, strongly sclerotized
uncus distally with two spines or teeth, the lamellate, distally inated sella set with
n-shaped setae forming editum, the dorsally inated sacculus with the dorsal mar-
gin sclerotized and usually spinulose, the distally broad and usually spinulose phal-
lus, and the spine-like cornuti appear funnel-shaped in the distal end of the vesica.
e female genitalia are characterized by the strongly sclerotized lamella postvagi-
nalis 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 (usu-
ally less than 24 mm). However, some Spinosuncus species have a similar body size to
Pseudopagyda, but they can still be dierentiated by the wavy or dentate lines on the
wings dorsally, especially the sinuate (rather than oblique, or slightly curved as in Pseu-
dopagyda) anterior part of the postmedial line near the costa. In the male genitalia, the
sclerotized uncus, the n-shaped setae (editum) of the sella, and the inated sacculus
distinguish Spinosuncus from Pseudopagyda. In the female genitalia, the long and slen-
der ductus bursae is distinct from the extremely short ductus bursae of Pseudopagyda.
Figure 1. Phylogenetic hypothesis inferred from Bayesian inference. Numbers on branches indicate
Bayesian posterior probabilities (values ≥ 0.8 are labelled) and ML bootstrap values probabilities (values
≥ 50% are labelled), respectively.
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 123
Description. Head. Frons oblique, yellowish brown, with white lateral bands. Ver-
tex with moderately raised scales projecting between antennae. Labial palpus obliquely
upturned, exceeding frons by 2/3 length of head or slightly less, third palpomere por-
rect, 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
agellomeres in male. orax. With appressed scales, yellow. Legs unmodied. 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; reti-
naculum 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 fe-
male. Wing venation (Figure 2) in forewing with cell about half length of wing; R1 free,
from 4/5 of anterior margin of cell, R2 free but adjacent to stem of R3+R4 in about basal
half, R3 and R4 stalked to about 2/3, R4 to just before apex, R5 parallel to stalked R3+R4
at base then diverging; M1 moderately close to R5 at base, M2 widely separate from M1,
closing vein concavely curved; M2, M3 and CuA1 from posterior angle of cell, M3 closer
to M2 at base than to CuA1, then diverging; CuA2 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+R1 and Rs anastomosing for 1/3 beyond end of discal cell, Rs and M1 short-stalked,
closing vein concave, angled medially; M3 closer to M2 at base than to CuA1, parallel
with M2 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
bid 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, la-
mellate, distally inated, set with modied setae (editum), varying from n-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 at, densely setose. Sinus vaginalis well developed, membra-
nous, 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
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
124
Figure 2. Wing venation of Spinosuncus praepandalis.
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 drop-
shaped 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 oc-
cur 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 ailand.
Etymology. e generic name is a compound word that refers to the uncus distally
with spines (“spinosus” in Latin). e resultant name is masculine in gender.
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 125
Spinosuncus contractalis (Warren, 1896), comb. n.
Figs 3, 10, 19, 28
Paliga contractalis Warren, 1896, 18(6): 123.
Microstega contractalis (Warren) Bänziger, 1995: 270.
Material examined. Type material. Lectotype, 1♂; Khasis, Warren Type, Pyralidae
Brit. Slide No. 8677 (NHMUK), designated by Bänziger (1995).
Other material examined. CHINA, Hainan: 2♂, 1♀, Bawangling, Changji-
ang, 19.12N, 109.08E, alt. 161 m, 22.VII.2014, leg. Cong Peixin, Hu Sha and Liu
Linjie, genitalia slide no. ZDD12049 (♂) (NKU); 1♀, Bawangling, 11.VI.2010,
leg. Kang Li, genitalia slide no. SYSU0185; 1♂, 1♀, Jianfengling, 5.VI.2010, leg.
Kang Li, genitalia slide no. SYSU0174 (♂); 1♀, Jianfengling, 18.75N, 108.85E, alt.
969 m, 12.IX.2013, leg. Xie Weicai, genitalia slide no. SYSU0067; 1♂, Bangxi Re-
serve, 19.37N, 109.10E, alt. 97 m, 2.IX.2013, leg. Chen Xiaohua, genitalia slide no.
SYSU0017; 1♂, Nankai Town, Baisha, 19.05N, 109.40E, alt. 294 m, 19.V.2013,
leg. Li Jinwei, genitalia slide no. SYSU0065; 1♂, Sanya Village, Fanjia, 19.25N,
109.65E, alt. 302 m, 27.X.2013, leg. Chen Kai and Chen Xiaohua, genitalia slide no.
SYSU0040; 1♀, Mt. Diaoluoshan, alt. 500 m, 24.V.2014, leg. Xu Dan and Xu Lijun,
genitalia slide no. SYSU0914; 1♀, Wuzhishan Natural Reserve, 18.88N, 109.65E,
alt. 742 m, 21.V.2015, leg. Cong Xinpei, Guan Wei and Hu Sha (NKU); Yunnan:
3♂, Bawan, Baoshan, alt. 1040 m, 9.VIII.2007, leg. Zhang Dandan, genitalia slide
no. SYSU0019; 5♂, 1♀, Baihualing, Baoshan, alt. 1520 m, 11,13.VII.2007, leg.
Zhang Dandan, genitalia slides no. CXH12155 (♂), SYSU0039 (♂), SYSU0047
(♂), SYSU0073 (♀); 2♂, Baihualing, Mt. Gaoligongshan, Baoshan City, 25.30N,
98.80E, alt. 1473 m, 29.VII.2013, leg. Liu Shurong, Teng Kaijian and Wang Yuqi
(NKU); 1♂, Baihualing, Mt. Gaoligongshan, Baoshan City, 25.30N, 98.80E, alt.
1473 m, 7.VIII.2014, leg. Liu Shurong, Rong Hua and Teng Kaijian (NKU); 1♂,
Dahaoping, alt. 2020 m, 6.VIII.2007, leg. Zhang Dandan; 1♂, Jingpozhai, Na-
bang, Yingjiang, 24.71N, 97.39E, alt. 231 m, 3.VIII.2013, leg. Liu Shurong, Teng
Kaijian and Wang Yuqi (NKU); 2♂, 1♀, 55 km site, Xishuangbanna Natural Re-
serve, 23.V.2015, leg. Zhang Zhenguo, genitalia slide no. ZDD12053 (♂, molecular
voucher no. SYSU-LEP0153) (NKU); 1♂, Yexiang Valley, Xishuangbanna, 22.17N,
100.87E, alt. 762 m, 18.VII.2014, leg. Guan Wei, Liu Shurong and Wang Xiuchun
(NKU); 2♂, 1♀, Yexiang Valley, Xishuangbanna, 22.17N, 100.87E, alt. 762 m, 10–
12.VII.2015, leg. Bai Xia and Teng Kaijian, genitalia slide no. ZDD12048 (♂, mo-
lecular voucher no. SYSU-LEP0135) (NKU); 1♂, Guanping, Mengyang, alt. 1200
m, 20.VIII.2005, leg. Ren Yingdang, genitalia slide no. CYP12056 (NKU); 1♂,
Nanla River, Bubang, Mengla, 21.59N, 101.58E, alt. 652 m, 15.VII.2013, leg. Liu
Shurong, Teng Kaijian and Wang Yuqi (NKU); 1♀, Yaoqu Town, Xishuangbanna,
alt. 780 m, 26.V.2015, leg. Tao Manfei, genitalia slide no. SYSU0913, molecular
voucher no. SYSU-LEP0133; Tibet: 1♀, Medog, alt. 1103 m, 8.VII.2013, leg. Li
Jinwei, genitalia slide no. SYSU0915.
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
126
Diagnosis. Within the genus, S. contractalis resembles S. rectacutus and S. bre-
vacutus 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 (approxi-
mately as long as the length of the corpus bursae). However, it can be dierentiated
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 genita-
lia, it is characterized by the curved sclerotized notches anterolaterally on the sinus
vaginalis. e dierences between S. contractalis and S. brevacutus are given in the
diagnosis of the latter species.
Redescription. Head. As for the genus. orax. Yellow. Legs as described for the
genus. Wingspan 18–22 mm. Wings yellow, lines fulvous. Forewing broadly trian-
gular 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 CuA2, then to 2/3 of posterior
margin; subterminal line from distal end of R2, arched to about 4/5 of CuA2, 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 M1 to distal third
of CuA2, bent inwardly to basal third of CuA2, then straight to near end of 2A;
subterminal line from distal third of RS, arched, tapering to CuA2, then concave to
distal end of 1A; fringe as in forewing. Abdomen. Yellow dorsally, apical margin of
segments tinged with white. Male genitalia (Figure10). 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; cos-
tal 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 inated, set with
several narrow, n-shaped setae forming editum, each seta with apex evenly divided
into several laments. 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 (Figure10C). Female genitalia (Figure 19). Posterior apophy-
sis with small expansion at basal third; anterior apophysis with small expansion
beyond basal half. Sinus vaginalis with two curved, sclerotized notches anterolat-
erally; 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 approxi-
matively drop-shaped; accessory bursa arising from posterior 1/3 of corpus bursae,
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 127
Figures 3–9. Adults of Spinosuncus spp. 3 S. contractalis, male (Dahaoping, Yunnan) 4 S. rectacutus,
holotype, male (Weng’ang Town, Guizhou) 5 S. brevacutus, holotype, male (Weng’ang Town, Guizhou)
6 S. praepandalis, male (Weng’ang Town, Guizhou) 7 S. curvisetaceus, paratype, male (Tongmu Village,
Fujian) 8 S. aureolalis, male (Bubang, Yunnan) 9 S. quadracutus, paratype, male (Mt. Limu, Hainan).
Scale bars: 5.0 mm.
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, ailand.
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
128
Spinosuncus rectacutus sp. n.
http://zoobank.org/B77B43A4-3F97-4157-ACD1-BD646B953C4D
Figs 4, 11, 20, 28
Material examined. Type material. Holotype ♂ (Fig. 4); CHINA, Guizhou: Weng’ang
Town, Maolan Reserve, Libo, 25.25N, 107.90E, alt. 814 m, 25.VII.2015, leg. Chen
Kai, genitalia slide no. SYSU0060. Paratypes: Hubei: 1♂, Pingbaying, Xianfeng, alt.
1280 m, 21.VII.1999, leg. Li Houhun et al., genitalia slide no. ZDD12055 (NKU);
3♂, Mahe Town, Xianfeng, alt. 400 m, 24–26.VII.1999, leg. Li Houhun et al., geni-
talia slide no. ZDD12056 (NKU); 1♂, Maobaqu, Lichuan, alt. 700 m, 29.VII.1999,
leg. Li Houhun et al., genitalia slide no. ZDD12057 (NKU); Guangxi: 1♂, Nanchao,
Yachang Forest Farm, Leye, alt. 1160 m, 26.VII.2004, leg. Xu Jiasheng (NKU); 1♂,
Huaping, Yachang Forest Farm, Leye, alt. 910 m, 28.VII.2004, leg. Xu Jiasheng, geni-
talia slide no. CYP12058 (NKU); 15♂, 2♀, Nonggang, Longzhou, 22.47N, 106.96E,
alt. 271 m, 19.IV.2012, leg. Li Jinwei, genitalia slides no. SYSU0036 (♀), 0188
(♀), 0009, 0014, 0024, 0041, 0042, 0043, 0044, 0054, 0055; 8♂, 2♀, Nonggang,
Longzhou, alt. 188 m, 25,27,28,31.VII.2011, leg. He Guiqing, genitalia slides no.
SYSU0189 (♂), 0194 (♀), 0979 (♀); 1♂, Nonggang Reserve, 21.VIII.2011, leg. Yang
Lijun, genitalia slide no. SYSU0053; 1♂, Nonggang Reserve, 21.VIII.2011, leg. Cheng
Muchun, genitalia slide no. CXH12165; 1♂, Nonggang, Longzhou, 20.VIII.2011,
leg. Cheng Muchun; 1♂, Nonggang, Longzhou, alt. 280 m, 29.VII.2012, leg. Yang
Xiaofei (NKU); 1♂, Sanlian, Longzhou, alt. 180 m, 1.VIII.2011, leg. He Guiqing;
1♂, Tongling Valley, 23.02N, 106.65E, alt. 535 m, 22.VII.2013, leg. Chen Xiao-
hua, genitalia slide no. SYSU0259; 1♂, Longrui Reserve, 18.VIII.2011, leg. Li Jin-
wei, genitalia slide no. SYSU0021; 1♂, Longrui Reserve, 19.VIII.2011, leg. Zhang
Dandan; 2♂, Bangliang, Jingxi, 1,5.VIII.2010, leg. Huang Jianhua, genitalia slide no.
ZDD12047 (NKU); Guizhou: 2♂, Xian’nv’dong, Dashahe, Daozhen, alt. 600 m,
28–29.V.2004, leg. Hao Shulian, genitalia slide no. CYP12057 (NKU); 1♂, Baishao
Village, Qinggangtang, Suiyang, alt. 800 m, 11.VIII.2010, leg. Du Xicui, genitalia
slide no. CYP12063 (NKU); 1♂, Baishao, Kuankuoshui, alt. 800 m, 10.VIII.2010,
leg. Du Xicui, genitalia slide no. SYSU0187; 1♂, Mt. Leigongshan, 26.35N, 108.15E,
alt. 1198 m, leg. Chen Xiaohua, genitalia slide no. SYSU0057, molecular voucher
no. SYSU-LEP0155; 1♂, Dongdai, Shuizu Town, Limingguan, Libo, alt. 720 m,
19.VII.2015, leg. Li Jia’en and Yang Meiqing, genitalia slide no. ZDD12050, mo-
lecular voucher no. SYSU-LEP0134 (NKU); 1♀, Weng’ang Town, Maolan Reserve,
Libo, 25.25N, 107.90E, alt. 814 m, 25.VII.2015, leg. Chen Kai, genitalia slide no.
SYSU0072; Chongqing: 2♂, 1♀, Xiaonanhai, Qianjiang, alt. 370 m, 21.VII.2012,
Xu Lijun and Zhang Jun, genitalia slides no. SYSU0186 (♂), 0193 (♀).
Diagnosis. Spinosuncus rectacutus resembles S. contractalis and S. brevacutus, for
which details are provided in the diagnosis of S. contractalis. It can be best distin-
guished 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. e distal spines of the uncus are straight and longer than those
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 129
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. orax. Yellow. Legs as described for the ge-
nus. Wingspan 18–22.5 mm. Wing pattern as in S. contractalis. Abdomen. Yellow dor-
sally, 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; tran-
stilla 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 inat-
ed, set with several narrow, n-shaped setae forming editum, each seta with apex evenly
divided into several laments. 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. e specic name is derived from the Latin recti- for straight and acu-
tus, pointed, referring to the straight, pointed spines of the uncus.
Distribution.(Figure 28). China (Hubei, Guangxi, Guizhou, Chongqing).
Spinosuncus brevacutus sp. n.
http://zoobank.org/D2E5D324-0CEB-46A5-B31E-4B520E704B72
Figs 5, 12, 21, 28
Material examined. Type material. Holotype ♂ (Fig. 5); CHINA, Guizhou:
Weng’ang Town, Maolan Reserve, Libo, 25.25N, 107.90E, alt. 814 m, 25.VII.2015,
leg. Chen Kai, genitalia slide no. SYSU0056, molecular voucher no. SYSU-LEP0009.
Paratypes: Jiangxi: 1♂, Main Peak, Mt. Jinggangshan, 28.IV.2011, leg. Liu Ping
and Mei Yan, genitalia slide no. CXH12192; 1♂, Main Peak, Mt. Jinggangshan,
30.VI.2011, leg. Yang Lijun, genitalia slide no. CXH12161; 1♂, Main Peak, Mt.
Jinggangshan, 1.IX.2011, leg. Cheng Muchun, genitalia slide no. SYSU0064; 1♂,
1♀ (Abdomen lost), Reservoir of Mt. Jinggangshan, 19.IX.2010, leg. Tong Bo,
Zhang Dandan and Zhao Shuang; 1♂, 1♀, Mt. Guanggushan, Wuzhifeng Town,
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
130
Shangyou, 25.92N, 114.05E, alt. 846 m, 22.VI.2015, leg. Chen Kai, genita-
lia slides no. SYSU0015 (♂), 0062 (♀, molecular voucher no. SYSU-LEP0156);
1♂, Mt. Jiulianshan, Longnan, 24.54N, 114.46E, alt. 625 m, 28.IV.2012, leg. Li
Jinwei, genitalia slide no. SYSU0049; Hunan: 1♀, Visitors’center, Taoyuandong,
26.47N, 114.04E, alt. 870 m, 20.V.2014, leg. Chen Xiaohua, genitalia slide no.
SYSU0063; Guizhou: 1♂, Maolan Reserve, 1.IX.2011, leg. Li Jinwei, genitalia slide
no. CXH12162; 4♂, Maolan Reserve, 25.13N, 107.87E, alt. 797 m, 12.VII.2013,
leg. Chen Xiaohua, genitalia slides no. SYSU0020, 0023, 0074, 0910; 1♂, 2♀,
Weng’ang Town, Maolan Reserve, Libo, 25.25N, 107.90E, alt. 814 m, 25.VII.2015,
leg. Chen Kai, genitalia slides no. SYSU0046 (♂), 0071 (♀, molecular voucher no.
SYSU-LEP0010), 0978 (♀).
Diagnosis. Spinosuncus brevacutus is similar to S. contractalis and S. rectacutus.
Dierences with S. rectacutus are given in the diagnosis of S. rectacutus. It can be dis-
tinguished from S. contractalis by the minute and weakly outwardly curved spines of
the apical uncus, the concave margin between those spines, the moderately setose tran-
stilla 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.
Description. Head. As for the genus. orax. Yellow. Legs as described for the
genus. Wingspan 19–24 mm. Wing pattern as in S. contractalis. Abdomen. Yellow dor-
sally, 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 me-
dium width, length approximately 2.5× its maximal width; transtilla with dorsal mar-
gin 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 inated and upcurved, set
with several narrow, n-shaped setae forming editum, each seta with apex evenly di-
vided into several laments. Juxta heart-shaped, distal half divided. Phallus as in S. con-
tractalis. 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 some-
what 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. e specic 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).
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 131
Figures 10–12. Male genitalia of Spinosuncus spp. 10 S. contractalis, Hainan (genitalia slide no.
SYSU0017) 11 S. rectacutus, Guangxi (genitalia slide no. SYSU0044) 12 S. brevacutus, Guizhou (genitalia
slide no. SYSU0910). A: Whole genitalia. B: Base of valva dorsally. C: Apex of phallus. Scale bars: 0.5 mm.
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
132
Spinosuncus praepandalis (Snellen, 1890), comb. n.
Figs 6, 13, 22, 28
Botys praepandalis Snellen, 1890: 573–574.
Material examined. Type material. Lectotype, 1♀; Sikkim, O. Miller., [18]89, collec-
tion of H. J. Elwes, Pyralidae Brit. Slide no. 9711 (NHMUK).
Other material examined. CHINA, Hubei: 1♂, 1♀, Shayuan, Hefeng, alt. 1260
m, 15,17.VII.1999, leg. Li Houhun, genitalia slides no. ZDD02388 (♂), 02389
(♀) (NKU); Hunan: 1♀, Jiangping, Mt. Hupingshan, Shimen County, alt. 480 m,
6.V.2002, leg. Yu Haili (NKU); Sichuan: 1♂, Wannian Temple, Mt. E’meishan,
29.59N, 103.38E, alt. 830 m, 14.VII.2014, leg. Guan Wei, Liu Shurong and Wang
Xiuchun (NKU); Chongqing: 1♂, Dawopu, Mt. Simianshan, 28.58N, 106.35E,
alt. 1059 m, 12.VII.2016, leg. Chen Kai; 1♂, Tiantangba, Mt. Simianshan, 28.64N,
106.35E, alt. 921 m, 13.VII.2016, leg. Chen Kai; 1♂, Mt. Jinfoshan, alt. 1700 m,
13.VII.2010, leg. Du Xicui and Shi Shengwen, genitalia slide no. SYSU0191; 1♀,
Wuli Town, Qianjiang, alt. 870 m, 23.VII.2012, leg. Xu Lijun and Zhang Jun, geni-
talia slide no. SYSU0196; Guizhou: 1♂, Heiwan, Jiangkou, alt. 600 m, 28.VII.2001,
leg. Li Houhun and Wang Xinpu, genitalia slide no. ZDD02061 (NKU); 2♂, Huguo
Temple, Mt. Fanjingshan, alt. 1390 m, 28.V.2002, leg. Wang Xinpu, genitalia slide no.
CYP12041 (NKU); 3♂, Weng’ang Town, Maolan Reserve, Libo, 25.25N, 107.90E,
alt. 814 m, 25.VII.2015, leg. Chen Kai, genitalia slide no. SYSU0038, molecular
voucher no. SYSU-LEP0006; 1♂, Weng’ang Town, Libo, alt. 1345 m, 18.VII.2015,
leg. Wan Jiping; Yunnan: 4♂, 2♀, Mt. Jizushan, Binchuan, 25.93N, 100.38E, alt.
1831 m, 29.VI.2012, leg. Li Jinwei, genitalia slides no. CXH12156 (♂), SYSU0045
(♂), 0076 (♀), 0980 (♀); 5♂, 1♀, Mt. Gaoligongshan, Baoshan, 24.82N, 98.78E, alt.
1700 m, 22.V.2016, leg. Duan Yongjiang, genitalia slides no. SYSU0190 (♂), 0195
(♀, molecular voucher no. SYSU-LEP0131); 2♀, Dahaoping, Tengchong, alt. 2020
m, 6.VIII.2007, leg. Zhang Dandan, genitalia slides no. SYSU0078, 0183; 1♂, Pi-
anma Village, Lushui, Nujiang, alt. 1889 m, 16.VIII.2015, leg. Wei Xueli; 1♀, Malipo
County, alt. 1098 m, 4.VI.2015, leg. Tao Manfei, genitalia slide no. SYSU0911; Ti-
bet: 1♂, 1♀, Hanmi, Medog, alt. 2380 m, 9.VIII.2003, leg. Wang Xinpu and Xue
Huaijun, genitalia slide no. CYP12062 (♂) (NKU); 1♂, Shangzayü, Nyingchi, alt.
1936 m, 16.VIII.2015, leg. Xu Dan. INDIA: 1♂, India, Sikkim, Elwes, collection
of H. J. Elwes, Pyralidae Brit. Slide no. 8674 (NHMUK); 1 ♂, Sikkim, O. Miller.,
[18]89, collection of H. J. Elwes (NHMUK).
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 dif-
ferentiated 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 bid
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-
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 133
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. curvi-
setaceus) and the long and slender ductus bursae, which is more than twice as long as
the diameter of the corpus bursae, diers from that of the species described above (the
ductus bursae is almost as long as the length of the corpus bursae). e dierences be-
tween S. praepandalis and S. curvisetaceus are given in the diagnosis of the latter species.
Redescription. Head. As for the genus. orax. Yellow. Legs as described for the ge-
nus. 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 CuA2.
Abdomen. Yellowish to brown, apical margin of segments tinged with white. Male geni-
talia (Figure 13). Uncus tapering towards apex; distal 3/4 bid, 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 o), 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 inated, set with
a few broad, n-shaped setae forming editum, each seta with apex evenly divided into
several laments. Juxta pentagonal, weakly bid 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 dis-
tinct 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 trans-
verse 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, globu-
lar; 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.
Distribution. (Figure 28). China (Hubei, Hunan, Sichuan, Chongqing, Guizhou,
Yunnan, Tibet), India.
Spinosuncus curvisetaceus sp. n.
http://zoobank.org/5E9A3861-D420-43A5-9B85-343125D46FCB
Figs 7, 14, 23, 28
Material examined. Holotype ♂; CHINA, Jiangxi: Mt. Sanqingshan, Jinsha Coun-
ty, Shangrao, alt. 380–390 m, 20.IV.2007, leg. Bai Haiyan and Du Xicui, genitalia
slide no. ZDD12058 (NKU). Paratypes: Fujian: 1♂, Tongmu Village, Mt. Wuyis-
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
134
han, 3.V.2014, leg. Yang Xiaofei, genitalia slide no. ZDD12051 (NKU); Jiangxi: 1♂,
Shiguling Power Plant, Mt. Sanqingshan, Jinsha County, Shangrao, alt. 410–420 m,
15.IV.2007, leg. Bai Haiyan and Du Xicui, genitalia slide no. CYP12066 (NKU); 6♂,
Mt. Sanqingshan, Jinsha County, Shangrao, alt. 380–390 m, 19, 20.IV.2007, leg. Bai
Haiyan and Du Xicui, genitalia slides no. CYP12060, 12074, ZDD12026 (NKU);
1♂, Shixi Town, Fengxin, 28.44N, 114.54E, alt. 506 m, 22.IX.2012, leg. Yang Li-
jun, genitalia slide no. CXH12167; 1♀, Nanfengmian Reserve, Qianmo Village, Sui-
chuan, 26.28N, 114.06E, alt. 816 m, 19.VI.2015, leg. Chen Kai, genitalia slide no.
SYSU0061, molecular voucher no. SYSU-LEP0129; Guangxi: 1♂, Jiuniutang, Mt.
Mao’ershan, alt. 550 m, 20.IV.2002, leg. Hao Shulian and Xue Huaijun, genitalia slide
no. ZDD02245 (NKU); 1♂, Huawang Villa, Jinxiu, alt. 550 m, 13.IV.2002, leg. Hao
Shulian and Xue Huaijun, genitalia slide no. ZDD02241 (NKU).
Diagnosis. Spinosuncus curvisetaceus resembles S. praepandalis in wing pattern. e
wingspan of S. curvisetaceus is usually smaller than that of S. praepandalis, S. aureolalis
and S. quadracutus, but larger than in S. contractalis, S. rectacutus and S. brevacutus. e
ground colour of the wings is paler than that of S. praepandalis. In the male genitalia,
it can be dierentiated from S. praepandalis by the straight mediobasal margin of the
distal teeth of the uncus (curved in S. praepandalis), the curved setae on the transtilla
dorsally (straight in S. praepandalis) and the expanded, rectangular distal half of sac-
culus, with sclerotized and densely spinulose dorsal margin. In the female genitalia, the
anterior apophysis is thicker than that of S. praepandalis. It can be distinguished from
other Spinosuncus species (except S. praepandalis) by the distally strongly bid uncus,
forming two sclerotized, large excurved teeth bearing hair-like setae basally, two thick
needle-shaped setae on the transtilla dorsally and the absence of sclerotized, streak-like
or hook-like notches anterolaterally on the sinus vaginalis.
Description. Head. As for the genus. orax. Yellowish brown. Legs as described
for the genus. Wingspan 24–26 mm. Wing pattern as in S. praepandalis, ground col-
our 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 bid, 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 max-
imal 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 spinu-
lose, distally twisted; sella long and slender, rod-like, upcurved, distally set with few
broad, n-shaped setae, each seta with apex evenly divided into several laments.
Juxta shield-shaped, distal half divided medially. Phallus as in S. praepandalis. Fe-
male 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
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 135
Figures 13–14. Male genitalia of Spinosuncus spp. 13 S. praepandalis, Guizhou (genitalia slide no.
SYSU0038) 14 S. curvisetaceus, Fujian (genitalia slide no. ZDD12051). A: Whole genitalia. B: Base of
valva dorsally. C: Apex of phallus. Scale bars: 0.5 mm.
times as long as length of corpus bursae; colliculum narrower at anterior end. Cor-
pus bursae small, ovoid; accessory bursa arising from posterior 1/3 of corpus bursae;
rhombic signum as in S. praepandalis; second signum absent.
Etymology. e specic 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).
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
136
Spinosuncus aureolalis (Lederer, 1863), comb. n.
Figs 8, 15–16, 24–25, 28
Botys aureolalis Lederer, 1863: 473.
Pyralis ochrealis Moore, 1877: 614.
Microstega aureolalis (Lederer): Bänziger, 1995: 270.
Material examined. Type material. Lectotype of Pyralis ochrealis: 1♂; Sikkim,
Moore Coll. 94-106, Pyralidae Brit. Slide No. 8678 (NHMUK), designated by Bän-
ziger (1995).
Other material examined. CHINA, Guangxi: 2♂, Nonggang, Longzhou, alt.
188 m, 26.VII.2011, leg. He Guiqing, genitalia slide no. SYSU0909; Yunnan: 1♀,
Baihualing, Baoshan, alt. 1251 m, 13.VIII.2007, leg. Zhang Dandan, genitalia slide
no. SYSU0075; 2♂, Baihualing, Baoshan, alt. 1520 m, 11,13.VIII.2007, leg. Zhang
Dandan, genitalia slides no. SYSU0050, 0066; 1♂, 1♀, Mengla, alt. 800 m, 6,8.
VII.2012, leg. Kitching and Ashton, genitalia slide no. FCEL0002 (♀) (FCEL); 2♂,
1♀, Bubang, Xishuangbanna, 21.60N, 101.59E, alt. 656 m, 23.VII.2014, leg. Guan
Wei, Liu Shurong, Teng Kaijian and Wang xiuchun, genitalia slide no. ZDD12052
(♀, molecular voucher no. SYSU-LEP0146), ZDD12054 (♂) (NKU); 1♂, Nabang,
Yingjiang County, 24.75N, 97.56E, alt. 239 m, 27.V.2016, leg. Duan Yongjiang,
genitalia slide no. SYSU0958, molecular voucher no. SYSU-LEP0132; 1♂, Pianma
Village, Lushui, Nujiang, alt. 1889 m, 16.VIII.2015, leg. Wei Xueli, genitalia slide no.
SYSU0959; 1♂, Daxichang, Malipo County, alt. 1465 m, 7.VI.2015, leg. Tao Manfei,
genitalia slide no. SYSU0173.
Diagnosis. Spinosuncus aureolalis has a large wingspan (more than 26 mm). e
ground colour of the wings is the darkest within the genus. ough 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 n-
and 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 diag-
nostic. e appearance of S. aureolalis is most similar to that of S. quadracutus, both
having the same wing pattern. e dierences between these two species are given
in the diagnosis of S. quadracutus.
Redescription. Head. As for the genus. orax. 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-
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 137
Figures 15–16. Male genitalia of Spinosuncus aureolalis. 15 Yunnan (genitalia slide no. ZDD12054)
16Yunnan (genitalia slide no. SYSU0173). A: Whole genitalia. B: Base of valva dorsally. C: Projections
of sacculus. D: Apex of phallus. Scale bars: 0.5 mm.
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 me-
dioventrally. 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,
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
138
set with dense and at-lying spines (except distal half, Figs 15C, 16C) and two mod-
erately downcurved spines pointing towards juxta (sometimes the longer one absent,
Figure 16C); sella short and broad, distally inated, set with modied setae forming
editum, varying from n-shaped to thick needle-shaped, ventral margin upcurved,
thickened and sclerotized, distally spinose, ended in long, curved spine. Juxta shield-
shaped, 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 postvagi-
nalis 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.
Distribution. (Figure 28). China (Guangxi, Yunnan), India (Sikkim), ailand
(Chiang Mai).
Spinosuncus quadracutus sp. n.
http://zoobank.org/181BC4CF-DC36-4D53-9084-4AC91688D188
Figs 9, 17–18, 26–28
Material examined. Type material. Holotype ♂; CHINA, Hainan: Mt. Limushan,
19.16N, 109.73E, alt. 662 m, 20.V.2013, leg. Li Jinwei, genitalia slide no. SYSU0048,
molecular voucher no. SYSU-LEP0002. Paratypes: CHINA, Fujian: 1♂, Guadun,
Mt. Wuyishan, 27.74N, 117.64E, alt. 1220 m, 17.V.2012, leg. Li Jinwei, genitalia
slide no. SYSU0034; Hainan: 2♂, Mt. Limushan, 19.16N, 109.73E, alt. 662 m,
20.V.2013, leg. Li Jinwei, genitalia slide no. SYSU0032; 1♂, 1♀, Jianling Reserve,
18.87N, 110.27E, alt. 143 m, 8.IX.2013, leg. Chen Xiaohua, genitalia slides no.
SYSU0029 (♂), SYSU0035 (♀, molecular voucher no. SYSU-LEP0001); 1♀, Mt.
Diaoluoshan, 18.65N, 109.93E, alt. 98 m, 3.XI.2013, leg. Chen Kai and Chen Xiao-
hua, genitalia slide no. SYSU0912; 1♀, Nankai Town, Baisha, 19.05N, 109.24E, alt.
294 m, 19.V.2013, leg. Li Jinwei, genitalia slide no. SYSU0077.
Diagnosis. is species is indistinguishable from S. aureolalis in wing pattern. In
the male genitalia, it can be distinguished from S. aureolalis by the uncus with four
prominent pointed spines distally (the median two small and indistinct in S. aureo-
lalis), the blunt distal projection of sacculus (pointed in S. aureolalis) always set with
one long spine pointing towards juxta (often with two long spines in S. aureolalis,
Figure15C) and the more spinulose and with arched dorsal margin distal projection
(smooth, less arched in S. aureolalis, Figs 15C, 16C). In the female genitalia, it can
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 139
Figures 17–18. Male genitalia of Spinosuncus quadracutus. 17 Fujian (genitalia slide no. SYSU0034) 18
Hainan (genitalia slide no. SYSU0048). A: Whole genitalia. B: Base of valva dorsally. C: Projections of
sacculus. D: Apex of phallus. Scale bars: 0.5 mm.
be dierentiated from S. aureolalis by the more closely set dorsolateral extensions of
lamella postvaginalis and relatively larger and more closely set hook-like notches of the
sinus vaginalis anterolaterally (Figs 26B, 27B).
Description. Head. Frons brown, vertex with moderately raised scales project-
ing between antennae, labial palpus brown, white at base ventrally. Maxillary palpus
brown, with apex pale yellow. orax. Yellow. Legs as described for the genus. Wing-
span 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
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
140
Figures 19–21. Female genitalia of Spinosuncus spp. 19 S. contractalis, Hainan (genitalia slide no.
SYSU0185) 20 S. rectacutus, Guangxi (genitalia slide no. SYSU0979) 21 S. brevacutus, Guizhou (genitalia
slide no. SYSU0978). A–B: Ventral views. B: Posterad of colliculum. Scale bars: 1.0 mm.
from base to middle; laterally membranous and set with several setae ventrally; other-
wise 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 nar-
row, extended to near distal end of valva; sacculus with central caniniform projection
and distal cheliform projection, distal half set with dense setae ventrally, distal projec-
tion strongly sclerotized, set with dense and slightly raised spines and one moderately
downcurved spine pointing towards juxta; sella short and broad, distally inated, set
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 141
with modied setae forming editum, varying form n-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 bid. Phallus as in
S. aureolalis. Female genitalia (Figs 26, 27). Anterior apophysis sclerotized, slightly sin-
Figures 22–23. Female genitalia of Spinosuncus spp. 22 S. praepandalis, Yunnan (genitalia slide no.
SYSU0980) 23 S. curvisetaceus, Jiangxi (genitalia slide no. SYSU0061). A–B: Ventral views. B: Posterad
of colliculum. Scale bars: 1.0 mm.
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
142
Figures 24–25. Female genitalia of Spinosuncus aureolalis. 24 Yunnan (genitalia slide no. ZDD12052)
25Yunnan (genitalia slide no. FCEL0002) A–B: Ventral views. B: Posterad of colliculum. Scale bars: 1.0 mm.
uate in distal third; posterior apophysis oblong, slender, and strongly sclerotized. Sinus
vaginalis with two large, thick, hook-like notches anterolaterally; lamella postvaginalis
sclerotized, band-shaped, extended dorsolaterally to approximately 1/3 width of sinus
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 143
Figures 26–27. Female genitalia of Spinosuncus quadracutus. 26 Hainan (genitalia slide no. SYSU0912)
27Hainan (genitalia slide no. SYSU0035). A–B: Ventral views. B: Posterad of colliculum. Scale bars: 1.0 mm.
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
(Figure26A) or weak and wide separated (Figure 27A), other two angles densely bear-
ing spines, sometimes smooth medially (Figure 27A); second signum absent.
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
144
Etymology. e specic name is derived from the Latin quadri- (four) and acutus
(pointed), referring to the distal uncus with four pointed spines.
Distribution.(Figure 28). China (Fujian, Hainan)
Key to species of Spinosuncus
1 Wingspan large, usually more than 24 mm. Uncus with setae. Ductus bursae
at least twice as long as length or diameter of corpus bursae ........................ 2
– Wingspan small, usually less than 24 mm. Uncus without setae. Ductus bur-
sae approximately as long as length or diameter of corpus bursae ................ 5
2 Forewing with postmedial line thickened near costa. Uncus without teeth ven-
trally; transtilla with two to three thick needle-shaped setae; sella distally only
with n-shaped setae. Antrum membranous; sinus vaginalis without notch .... 3
– Forewing with postmedial line not thickened near costa. Uncus with two
caniniform teeth ventrally; transtilla with few normal setae; sella distally with
n-shaped and thick needle-shaped setae. Antrum granulated; sinus vaginalis
with two large, hook-like notches ...............................................................4
3 Wings with lines somewhat dentate; ground color dark yellow (Fig. 6). Transtilla
with two to three straight setae, the longer one about twice as long as the shorter
one (Fig. 13B); distal half of sacculus expanded, semicircular, with margin sparse-
ly set with teeth (Fig. 13A) ..............S. praepandalis (Snellen, 1890), comb. n.
– Wings with lines not dentate, background color pale yellow (Fig. 7). Transtil-
la with two curved setae, the longer one less than twice as long as the shorter
one (Fig. 14B); distal half of sacculus expanded, rectangular, with margin
densely set with teeth (Fig. 14A) ................................S. curvisetaceus sp. n.
4 Uncus distally with two thick teeth, with two very minute spines, often indis-
tinct (Figs 15A, 16A); sacculus with distal projection relatively pointed, often
bearing two long spines pointing towards juxta (Figs 15C, 16C) ..................
...................................................... S. aureolalis (Lederer, 1863), comb. n.
– Uncus distally with four slender spines, the lateral two about two times as
long as the median two (Figs 17A, 18A); sacculus with distal projection
blunt, bearing one long spine pointing towards juxta (Figs 17C, 18C)..........
...................................................................................S. quadracutus sp. n.
5 Uncus distally blunt, with two minute spines (Fig. 12A); transtilla dorsally
with a row of sparse setae (Fig. 12B) ..............................S. brevacutus sp. n.
– Uncus distally with two distinct spines laterally; transtilla dorsally densely
setose .......................................................................................................... 6
6 Distal spines of uncus excurved, lateral margins below spines strongly bulging;
distal half of sacculus with acinaciform process (Fig. 10A). Notches of sinus
vaginalis curved (Fig. 19B) ........... S. contractalis (Warren, 1896), comb. n.
– Distal spines of uncus straight, lateral margins below spines slightly bulging;
distal half of sacculus without acinaciform process (Fig. 11A). Notches of
sinus vaginalis straight (Fig. 20B) ....................................S. rectacutus sp. n.
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 145
Discussion
e results of the molecular analysis robustly support the monophyly of Spinosuncus. e
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 inated sella with n-shaped setae forming editum, the dorsally expanded sac-
culus 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. e relationship between Pseudopagyda +
Aglaops and Spinosuncus is well supported (PP = 1.00, BS = 77), but the clade Pseu-
dopagyda + Aglaops is only weakly supported (PP = 0.81, BS = 50). Morphologically,
Aglaops species are similar to Pseudopagyda in the campanulate uncus and the distally
inated sella (which is also similar in Spinosuncus), but other genitalia traits dier. e
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 inated 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. praepanda-
lis 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 furnaca-
lis (Guenée, 1854), Placosaris rubellalis (Caradja, 1925), liptoceras sinense (Caradja,
1925) and Aglaops youboialis (Munroe & Mutuura, 1968), all lack a forewing subtermi-
nal line. Placosaris rubellalis and liptoceras sinense have a rod-like sella similar to that
of Spinosuncus in the male genitalia, but the editum are dierent. 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 dierent from those of Spinosuncus species. At
present, it is impossible to conrm the generic position of Spinosuncus within the sub-
family 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. e monophyly of these three
species groups is well supported by the phylogenetic analysis (Figure 1). e aureolalis
group, comprising S. aureolalis and S. quadracutus, is well characterized by the later-
ally setose uncus, distally with two or four spines, ventrally with two large teeth; the
lamellate, distally inated sella with n- and needle-shaped setae forming editum; the
cheliform sacculus with a long spine pointing towards juxta; and the two thick, hook-
like notches anterolaterally on the sinus vaginalis. e contractalis group comprises S.
contractalis, S. rectacutus and S. brevacutus. is species group is well dened by several
characters: the glabrous, sclerotized uncus distally with two spines; a row of setae on the
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
146
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. e praepandalis group, comprising S. praepandalis and S. curvisetace-
us, can be recognized by the following characters: the bid 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
dierences from related species. e morphological dierences are given above in the
diagnoses of the new species. e 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 stud-
ies in Lepidoptera (Hebert et al. 2003, 2010, Yang et al. 2016). e low interspecic
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 intraspecic divergence was observed in S. aureolalis (2.7%). e 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 conspecic. Genital variation is observed
in two male specimens collected in Daxichang, Yunnan and Nonggang, Guangxi (genita-
lia slides no. SYSU0173, SYSU 0909, respectively), both places which are near the north
of Vietnam. e 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, ailand 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 supercially similar
to the three described species. Considering the lack of sucient generic revisions, es-
pecially 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 in-
terspecic and intraspecic dierences will certainly help to move ahead to natural
classications as opposed to articial arrangements. However, the understanding of the
Systematics of the new genus Spinosuncus Chen, Zhang & Li... 147
Figure 28. Distribution of Spinosuncus species in China. A Distance between Bubang and Nabang.
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 dierences which are dicult
to interpret as apo- or plesiomorphic. e use of genetic data will facilitate species
identication and help to understand the interspecic and intergeneric relationships.
It calls for more comprehensive investigations on Pyraustinae in the future in order to
understand this species-rich subfamily better.
Acknowledgements
Grateful thanks go to Mr Geo Martin (NHMUK) for helping to access type speci-
mens, to Dr Koen Maes (Agrobiosys International, Belgium) for giving helpful scientic
comments, to Dr Hans Bänziger (Chiang Mai University, ailand) for providing some
images of genitalia and specimens, and to Dr Xicui Du (Southwest University, China)
for providing some specimens. We are also grateful to Dr Wolfgang Speidel (Depart-
ment of Entomology, Zoologische Staatssammlung München, Germany), Dr Robert B.
Angus (NHMUK), Dr Richard Mally (University Museum of Bergen, Natural History
Collections, Norway) and Dr Bernard Landry (Muséum d’histoire naturelle, Geneva,
Switzerland) for their critical reviews and linguistic assistance on the manuscript.
is project was supported by the National Natural Science Foundation of China
(Grant No. 31672330), the Program of the Ministry of Science and Technology of
China. (2015FY210300) and the Basic Work Special Project of the National Ministry
of Science and Technologyof China(2013FY111500).
Kai Chen et al. / ZooKeys 799: 115–151 (2018)
148
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