Research Article |
Corresponding author: Dandan Zhang ( zhdd61@163.com ) Academic editor: Colin Plant
© 2021 Lanbin Xiang, Kai Chen, Dandan Zhang.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Xiang L, Chen K, Zhang D (2021) Revision and phylogeny of the genus Loxoneptera Hampson, 1896 (Lepidoptera, Crambidae, Pyraustinae), based on morphology and molecular data. ZooKeys 1036: 75-98. https://doi.org/10.3897/zookeys.1036.63814
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The genus Loxoneptera Hampson, 1896 is revised based on external appearance and genitalia. It is comprised of eleven species, of which three are described as new species from China: L. crassiuncata Chen & Zhang, sp. nov., L. triangularis Chen & Zhang, sp. nov., and L. rectacerosa Chen & Zhang, sp. nov.; six species are proposed as new combinations: L. carnealis (Swinhoe, 1895), comb. nov., L. medialis (Caradja, 1925), comb. nov., L. pentasaris (Meyrick, 1932), comb. nov., L. bipunctalis (Hampson, 1912), comb. nov., L. brevipalpis (Snellen, 1890), comb. nov., and L. dichroma (Moore, 1888), comb. nov. A new replacement name, L. hampsoni Chen & Zhang, nom. nov., is proposed for L. carnealis Hampson, 1896, the type species of the genus, because it is a secondary homonym of L. carnealis (Swinhoe, 1895), comb. nov. External characters and genitalia morphology of all species are figured. Nucleotide sequences of COI, 16S rRNA, 28S rRNA, and EF-1α were used for the molecular analysis and phylogeny of Loxoneptera species.
Calamochrous, China, molecular phylogeny, new combinations, new species
The genus Loxoneptera was established as a monotypic genus by
Within the additional Chinese specimens collected, three undescribed species of Loxoneptera were recognised. Moreover, a few species of Calamochrous and Anania Hübner, 1823 were found to be congeneric with species of Loxoneptera. The aim of this study is to diagnose Loxoneptera based on external and genital characters, to clarify the species included in the genus, and to provide a preliminary phylogenetic hypothesis based on selected genetic markers.
The material studied, including the types of the newly described species, are all deposited at the Museum of Biology, Sun Yat-sen University, China (SYSBM) except those stored in the following institutions: Insect Collection of the College of Life Sciences, Nankai University, China (
Ten species in four genera were included in the molecular phylogenetic analyses (Table
Genus | Species | Voucher | Locality | GenBank accession number | References | |||
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COI | 16S | EF-1α | 28S | |||||
Eumorphobotys | eumorphalis | SYSULEP0046 | Fujian | MG739574 | MG739586 | MG739598 | MG739609 |
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SYSULEP0047 | Fujian | MG739575 | MG739587 | MG739599 | MG739610 |
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concavuncus | SYSULEP0042 | Yunnan | MG739571 | MG739583 | MG739595 | MG739606 |
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SYSULEP0175 | Guangxi | MG739581 | MG739593 | MG739604 | MG739616 |
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horakae | SYSULEP0043 | Sichuan | MG739572 | MG739584 | MG739596 | MG739607 |
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SYSULEP0172 | Sichuan | MG739580 | MG739592 | N/A | MG739615 |
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Loxoneptera | hampsoni | SYSULEP0166 | Hainan | MG739579 | MG739591 | MG739603 | MG739614 |
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SYSULEP0174 | Hainan | MW736545 | MW736550 | MW736555 | MW728364 | Present study | ||
albicostalis | SYSULEP0162 | Yunnan | MG739578 | MG739590 | MG739602 | MG739613 |
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medialis | SYSULEP0096 | Hainan | MG739576 | MG739588 | MG739600 | MG739611 |
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SYSULEP0171 | Guangdong | MW736546 | MW736551 | MW736556 | MW728365 | Present study | ||
SYSULEP0173 | Guangdong | MW736547 | MW736552 | N/A | N/A | Present study | ||
rectacerosa | SYSULEP0170 | Yunnan | MW736548 | MW736553 | N/A | N/A | Present study | |
carnealis | SYSULEP0044 | Guizhou | MG739573 | MG739585 | MG739597 | MG739608 |
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SYSULEP0186 | Yunnan | MW736549 | MW736554 | MW736557 | MW728366 | Present study | ||
Sclerocona | acutella | SYSULEP0152 | Macau | MG739577 | MG739589 | MG739601 | MG739612 |
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Euclasta | stoetzneri | SYSULEP0334 | Shannxi | MT738696 | MT734412 | MT724335 | MT734404 |
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The sequences were aligned using Clustal W (
The concatenated dataset of four genes consisted of 2511 nucleotide positions (658 for COI, 463 for 16S rRNA, 619 for 28S rRNA, and 771 for EF-1α). Both BI and ML analyses of the concatenated dataset inferred congruent topologies with only subtle differences in posterior probability and bootstrap values probability (Fig.
The results of the current phylogenetic analyses support that the undescribed species (here named as L. rectacerosa sp. nov.) should be placed in Loxoneptera, and that L. carnealis (Swinhoe, 1895) comb. nov. and L. medialis (Caradja, 1925) comb. nov. should be transferred from Calamochrous Lederer, 1863 to Loxoneptera. Within the genus, L. medialis + L. rectacerosa form a sister group with robust support (PP = 1.00, BS = 99), while L. carnealis is the sister group to L. medialis + L. rectacerosa (PP = 1.00, BS = 100). Loxoneptera albicostalis is associated with the clade L. carnealis + (L. medialis + L. rectacerosa), although with relatively low support (PP = 0.78, BS = 52). Loxoneptera hampsoni is the first-diverging species with strong support in the BI analysis (PP = 0.93), but with relatively low support in the ML analysis (BS = 65).
Pairwise distances of the barcoding region (COI) are given in Table
Pairwise distance of the COI barcoding region based on Kimura-2-parameter model.
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | ||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | LEP0046 Eumorphobotys eumorphalis | ||||||||||||||||
2 | LEP0047 Eumorphobotys eumorphalis | 0.000 | |||||||||||||||
3 | LEP0042 Eumorphobotys concavuncus | 0.072 | 0.072 | ||||||||||||||
4 | LEP0175 Eumorphobotys concavuncus | 0.070 | 0.070 | 0.006 | |||||||||||||
5 | LEP0043 Eumorphobotys horakae | 0.078 | 0.078 | 0.068 | 0.066 | ||||||||||||
6 | LEP0172 Eumorphobotys horakae | 0.074 | 0.074 | 0.068 | 0.066 | 0.003 | |||||||||||
7 | LEP0166 Loxoneptera hampsoni | 0.096 | 0.096 | 0.112 | 0.106 | 0.119 | 0.119 | ||||||||||
8 | LEP0174 Loxoneptera hampsoni | 0.096 | 0.096 | 0.112 | 0.106 | 0.119 | 0.119 | 0.000 | |||||||||
9 | LEP0162 Loxoneptera albicostalis | 0.087 | 0.087 | 0.091 | 0.091 | 0.107 | 0.103 | 0.087 | 0.087 | ||||||||
10 | LEP0096 Loxoneptera medialis | 0.087 | 0.087 | 0.094 | 0.092 | 0.099 | 0.098 | 0.090 | 0.090 | 0.077 | |||||||
11 | LEP0171 Loxoneptera medialis | 0.087 | 0.087 | 0.092 | 0.090 | 0.099 | 0.097 | 0.092 | 0.092 | 0.077 | 0.002 | ||||||
12 | LEP0173 Loxoneptera medialis | 0.087 | 0.087 | 0.092 | 0.090 | 0.099 | 0.097 | 0.092 | 0.092 | 0.077 | 0.002 | 0.000 | |||||
13 | LEP0170 Loxoneptera rectacerosa | 0.077 | 0.077 | 0.089 | 0.090 | 0.101 | 0.099 | 0.099 | 0.099 | 0.080 | 0.044 | 0.042 | 0.042 | ||||
14 | LEP0044 Loxoneptera carnealis | 0.101 | 0.101 | 0.103 | 0.107 | 0.118 | 0.118 | 0.112 | 0.112 | 0.080 | 0.067 | 0.065 | 0.065 | 0.082 | |||
15 | LEP0186 Loxoneptera carnealis | 0.101 | 0.101 | 0.103 | 0.105 | 0.117 | 0.117 | 0.108 | 0.108 | 0.077 | 0.064 | 0.062 | 0.062 | 0.079 | 0.003 | ||
16 | LEP0152 Sclerocona acutella | 0.099 | 0.099 | 0.113 | 0.113 | 0.119 | 0.119 | 0.108 | 0.108 | 0.108 | 0.106 | 0.104 | 0.104 | 0.102 | 0.124 | 0.124 | |
17 | LEP0334 Euclasta stoetzneri | 0.106 | 0.106 | 0.118 | 0.118 | 0.135 | 0.135 | 0.111 | 0.111 | 0.118 | 0.106 | 0.104 | 0.104 | 0.101 | 0.115 | 0.115 | 0.118 |
Loxoneptera Hampson, 1896: 405. Type species: Loxoneptera carnealis Hampson, 1896, by original designation.
In external appearance, the species of Loxoneptera are similar to species of Eumorphobotys Munroe & Mutuura, 1969 in the long and porrect labial palpus, the usually concolorous wings with no obvious pattern and the straight termen of forewing, but can be best distinguished by the triangular uncus, the rod-shaped dorsal projection of transtilla bearing long and thick hair at the apex, and the hook-shaped ventral sella in the male genitalia. In the female genitalia, the ductus bursae of Loxoneptera is shorter and stouter than that of Eumorphobotys. These two genera are also different in the shape of the signum, if present a nearly rhomboid signum with connected carina, or reduced into a keel-like carina in Loxoneptera, and a narrowly rhomboid signum with carina interrupted in Eumorphobotys. Eighth sternite in males of Loxoneptera is slightly sclerotised, with two slender and sclerotised anterolateral processes.
Head. Frons oblique, slightly protruding. Vertex with moderately raised scales projecting between antennae. Labial palpus ~ 2–2.5 × eye diameter; second segment obliquely upward, third segment long and porrect. Maxillary palpus small. Thorax. Legs unmodified usually, outer spur 1/3 to 1/2 the length of inner spur, sometimes outer spur minute. Wings. Forewing elongated triangular, termen obliquely straight to slightly curved; discal cell ~ 1/2 length of wing, R1 from ~ 3/4 of anterior margin of cell, R3 and R4 stalked to more than half of R4, R5 free from anterior angle of cell, parallel to stalked R3+R4 at base, then diverging, discocellular veins concavely curved, M1 close to R5 at base, free from discocellular veins and close to anterior angle of cell, M2, M3 and CuA1 from posterior angle of cell, CuA2 from 4/5 of the posterior margin of cell, 1A faintly sinuate to tornus; 2A forming complete loop and distally recurved before joining 1A; usually only with orbicular and reniform stigmata, sometimes no pattern. Hindwing fan-shaped, termen rounded; discal cell less than half length of wing, Sc+R1 and Rs anastomosed to half of Rs, discocellulars concave, M2, M3 and CuA1 from posterior angle of discal cell, CuA2 from 4/5 of the posterior margin of cell; without obviously spot. Abdomen. Eighth sternite in male with two slender and sclerotised anterolateral processes, pointed or slightly stout (Fig.
Male genitalia. Uncus triangular, glabrous or with few hair-like setae. Tegumen trapezoid. Saccus nearly triangular. Transtilla with developed ventral process, extending a rod-shaped projection dorsad, usually long, curved, and slender, and terminal part with many long hairs. Valva tongue-shaped; dorsal sella membranous, ventral sella usually with a hook-shaped, strongly sclerotised process, dorso-distal sella presented as a sclerite and usually extended as a long, hook-shaped, sclerotised process; editum absent or not obvious; sacculus broad. Juxta with basal part rivet-shaped, remainder usually with two long and slender bifid arms. Phallus tubular, vesica with spine-shaped cornuti and sometimes deciduous cornuti.
Female genitalia. Ovipositor lobes flat, densely setose. Anterior apophyses longer than posterior apophyses. Antrum sclerotised, cup-shaped or bowl-shaped; colliculum well developed and sclerotised; ductus seminalis entering near anterior end of colliculum; ductus bursae short and stout, almost as long as length of corpus bursae; corpus bursae oval, appendix bursae oval or absent, signum nearly rhomboid, with a carina not interrupted in middle, sometimes signum reduced into a carina, sometimes absent.
China, India, Indonesia, Malaysia.
1 | Forewing reddish brown; hindwing black-brown in male, with a triangular patch presented near the posterior angle of cell | 2 |
– | Forewing colour paler, not reddish brown; hindwing pale yellow, triangular patch absent | 3 |
2 | Costal band of forewing white, fringe white and with basal 1/4 black-brown (Fig. |
L. albicostalis |
– | Costal band of forewing brown, fringe pale yellow and with basal half black-brown (Fig. |
L. crassiuncata |
3 | Forewing with pale yellow stripes between veins, posterior margin with a small triangular indentation and a group of black-brown scales in male (Fig. |
L. hampsoni |
– | Forewing without pale yellow stripe between veins, posterior margin arc-shaped, without indentation and a group of black-brown scales in male; juxta normal | 4 |
4 | Distal part of phallus with a long and pointed spine, longer than the length of phallus (Fig. |
L. carnealis |
– | Distal part of phallus without spine, or spine shorter than the length of phallus | 5 |
5 | Distal part of juxta with a strongly sclerotised and narrowly triangular process (Fig. |
L. triangularis |
– | Distal part of juxta without process | 6 |
6 | Distal end of phallus densely decorated with short spines (Fig. |
L. pentasaris |
– | Distal end of phallus not decorated with short spines | 7 |
7 | Dorsal margin of valva forming a break angle subapically (Fig. |
L. rectacerosa |
– | Dorsal margin of valva without break angle | 8 |
8 | Dorsal margin of valva convex, dorso-distal sella extended outwards and not beyond the end of valva (Fig. |
L. medialis |
– | Dorsal margin of valva somewhat concave, dorso-distal sella extended ventrad and beyond the ventral margin of valva | 9 |
9 | Forewing with a stripe along posterior margin of cell (Fig. |
L. dichroma |
– | Forewing absent stripe on posterior margin of cell; wall of phallus not sclerotised | 10 |
10 | Distal part of phallus with a heavily sclerotised, spiny and thumb-shaped cornutus (Fig. |
L. brevipalpis |
– | Distal part of phallus with a weakly sclerotised, slice-shaped cornutus (Fig. |
L. bipunctalis |
Loxoneptera carnealis
Hampson, 1896: 406, fig. 219 (a junior secondary homonym of Notaspis carnealis Swinhoe, 1895). TL: India (Sikkim). TD:
Type material. Type ♂, Sikkim, O. Müller [Coll.], Pyralidae Brit. Mus. Slide No. 9752 (
Adults of Loxoneptera spp. 2 L. hampsoni nom. nov., male (Tibet) 3 L. hampsoni nom. nov., female (Hainan) 4 L. albicostalis, male (Yunnan) 5 L. crassiuncata sp. nov., paratype, male (Yunnan) 6 L. carnealis, male (Yunnan) 7 L. triangularis sp. nov., holotype, male (Yunnan) 8 L. rectacerosa sp. nov., holotype, male (Yunnan) 9 L. medialis, male (Guangdong). Scale bars: 5.0 mm.
Other material examined. China. Hainan: 1♂, Mt. Limushan, 5.V.2011, leg. Zhang Dandan & Yang Lijun; 1♂1♀, Mt. Limushan, 6.V.2011, leg. Zhang Dandan & Yang Lifeng, genitalia slide no. SYSU0117 (♂), no. SYSU0130 (♀); 1♂1♀, Mt. Limushan, 19.17°N, 109.73°E, alt. 662 m, 20.V.2013, leg. Li Jinwei, genitalia slide no. SYSU0929 (♂), no. SYSU0991 (♀), molecular voucher no. LEP0166 (♂), no. LEP0174 (♀). Yunnan: 2♂, Mengla, Xishuangbanna, 4, 6.X.2004, leg. R. L. Kitching, genitalia slide no. FCEL0003 (FCEL). Tibet: 1♂, 80K, Medog County, 29.66°N, 95.49°E, alt. 2059 m, 8.VIII.2017, leg. Qi Mujie & Yang Xiaofei (
Loxoneptera hampsoni is easily distinguished from other Loxoneptera species as follows: forewing with distinct, black-brown and point-like orbicular and reniform stigmata, bearing pale yellow stripes between veins, and veins with ochre-brown scales forming streaks; dorsal sella with a long and slender rod-shaped extension in the male genitalia.
Head. Frons brown, with white lateral bands. Vertex brown, mixed with some white erected scales. Labial palpus dark brown, with white scales on ventral side. Maxillary palpus brown. Antennae brown. Thorax. Dorsal side, patagia and tegula yellowish brown, ventral side grey white. Foreleg yellowish brown, dorsal tarsus grey white; ventral femur and tibia of midleg and hindleg grey white, others pale yellow. Wings. Wingspan 29.0–36.0 mm. Forewing termen straight, a small triangular indentation presented on 1/3 of posterior margin in male, and with a group of black-brown scales; yellowish brown, mixed with ochre-brown scales, pale yellow stripes presented between veins, and veins covered with ochre-brown scales forming streaks; orbicular stigma appearing as a black point, reniform stigma black, small and round; fringe white, basal 1/5 black-brown. Hindwing in male black-brown on terminal area, remaining areas pale yellow, a triangular patch present near posterior angle of cell, slightly concave and densely covered with pale brown scales; in female pale yellow, mixed with ochre-brown scales on termen; fringe brown in male, pale yellow in female. Abdomen. Dorsal side of abdomen black-brown, ventral side grey white; 5th abdominal segment with a group of pale yellow scales on each side in male; sternite VIII in male slightly sclerotised with two pointed anterolateral processes.
Male genitalia
(Fig.
14 The sternite VIII in male of L. albicostalis 15–19 Male genitalia of Loxoneptera spp. 15 L. hampsoni nom. nov., Hainan (genitalia slide no. SYSU0929) 16 L. albicostalis, Yunnan (genitalia slide no. ZDD12108) 17 L. crassiuncata sp. nov., Yunnan (genitalia slide no. FCEL0010) 18 L. carnealis, Guizhou (genitalia slide no. SYSU0165) 19 L. triangularis sp. nov., Yunnan (genitalia slide no. FCEL0004). Scale bars: 1.0 mm.
Male genitalia of Loxoneptera spp. 20 L. rectacerosa sp. nov., Yunnan (genitalia slide no. ZDD12059) 21 L. medialis, Guangdong (genitalia slide no. SYSU0987) 22 L. pentasaris, India (Pyralidae Brit. Mus. Slide No. 9747) 23 L. bipunctalis, India (Pyralidae Brit. Mus. Slide No. 9750) 24 L. brevipalpis, India (Pyralidae Brit. Mus. Slide No. 9748) 25 L. dichroma, India (Pyralidae Brit. Mus. Slide No. 9749). Scale bars: 1.0 mm.
Female genitalia
(Fig.
China (Hainan, Yunnan, Tibet), India.
The species is renamed after the last name of George Hampson, who proposed the genus Loxoneptera in 1896.
According to the characters of the male and female genitalia, Calamochrous carnealis (Swinhoe, 1895) is transferred to Loxoneptera in this paper, which creates a secondary homonym of Loxoneptera carnealis Hampson, 1896, the type species of Loxoneptera. The specific name of Loxoneptera carnealis Hampson, 1896 is not valid, therefore we give it a new replacement name, i.e., Loxoneptera hampsoni nom. nov.
Loxoneptera albicostalis Swinhoe, 1906: 415.
Type material. Type ♂, Padang, W. Sumatra, Pyralidae Brit. Mus. Slide No. 9753 (
Other material examined. China. Yunnan: 1♂, Jingpo Village, Nabang, Yingjiang Country, 24.71°N, 97.39°E, alt. 231 m, 2.VIII.2013, leg. Teng Kaijian et al., genitalia slide no. ZDD12108 (
In appearance, Loxoneptera albicostalis is extremely similar to L. crassiuncata and Eumorphobotys horakae Chen & Zhang, 2018 in the wing shape, the clean reddish brown forewing and the dark brown hindwing, but can be distinguished by the whiter costa of both wings, a group of dark brown scales on the posterior margin of forewing, and a group of scales on each side of the 5th abdominal segment in male. The underside of forewing in L. albicostalis is smoky brown, while that of E. horakae is pale yellow from anterior margin of cell to posterior margin. The male genitalia resemble that of L. crassiuncata but can be differentiated by the shorter and stouter uncus, the relatively longer and slender dorsal projection of transtilla, nearly triangular dorsal sella, the long and hook-shaped process of the ventral sella, as well as the absence of the spine-shaped cornutus in phallus.
Head. Frons brown. Vertex brown, mixed with some yellow erected scales. Labial palpus dark brown, with white scales on ventral side. Maxillary palpus brown. Antennae yellowish brown. Thorax. Dorsal side, patagia and tegula brown, ventral side grey white. Legs pale yellow to grey white; hindleg basal outer spur 1/5 of basal inner spur. Wings. Wingspan 32.0–36.0 mm. Forewing wide, reddish brown, without pattern; costal area white, mixed with pale brown scales at apex; termen straight; a small triangular indentation presented on the 1/3 of posterior margin in male, and with a group of black-brown scales; fringe white, with basal 1/4 black-brown. Hindwing black-brown, costa area pale yellow; a triangular patch presented near the posterior angle of cell, densely covered with pale brown scales; fringe black-brown. Underside smoky dark brown on forewing, and pale brown on posterior margin area. Abdomen. Dorsal side black-brown, ventral side grey white; abdominal segment V with a group of dark scales on each side in male; sternite VIII in male slightly sclerotised with two pointed anterolateral processes.
Male genitalia
(Fig.
Female genitalia. Unknown.
China (Yunnan), Indonesia (Sumatra), Malaysia.
The forewing colour of the type material of Loxoneptera albicostalis is pale yellow tinged with some reddish brown scales and differs from the specimen collected in China. No obvious difference could be found in the male genitalia between the type specimen and the Chinese specimen.
Type material. Holotype, ♂, China: Yunnan: Mengla, Xishuangbanna, 4.IX.2004, leg. R. L. Kitching, genitalia slide no. FCEL0010 (FCEL). Paratypes: China: Yunnan: 1♂, Mengla, Xishuangbanna, 28.IX.2004, leg. R. L. Kitching; 1♂, Mengla, Xishuangbanna, 29.IX.2004, leg. R. L. Kitching, genitalia slide no. FCEL0012 (FCEL).
Loxoneptera crassiuncata is similar to L. albicostalis in reddish brown forewing colour but male specimens can be distinguished by the unbroken posterior margin of forewing (without a small triangular indentation), without a group of black-brown scales, and abdominal segment V without a group of dark scales. In the male genitalia, it can be differentiated by the longer and slender uncus, the shorter and stouter dorsal projection of transtilla, the slender and rod-shaped dorsal sella, the relatively shorter and slightly curved process of the ventral sella, as well as the presence of a horn-shaped cornutus in phallus.
Head. Frons brown. Vertex brown. Labial palpus brown, with white scales on ventral side. Maxillary palpus brown, broadened distally with scales. Antennae dark brown. Thorax. Dorsal side, patagia and tegula brown, ventral side grey white. Legs yellowish white or pale yellow, dorsal of midlegs and hindlegs yellowish brown; hindleg with basal outer spur 1/4 of inner spur. Wings. Wingspan 29.0–31.0 mm. Forewing wide, termen nearly straight; reddish brown, brown at basal half of posterior portion, costal band brown, without pattern; fringe pale yellow, basal half and the posterior angle black-brown. Underside greyish brown. Hindwing black-brown, pale yellow on anterior margin; a triangular patch presented near the posterior angle of cell, the margin of triangular patch with pale yellow scales and the outer margin dentate; fringe black-brown. Underside greyish brown. Abdomen. Dorsal side of abdomen brown, ventral side pale yellow; sternite VIII in male slightly sclerotised with two pointed anterolateral processes.
Male genitalia
(Fig.
Female genitalia. Unknown.
China (Yunnan).
The specific name is derived from the Latin crassi- (thick) and uncatus (horn-shaped), referring to the shape of cornuti in the phallus.
Notaspis carnealis Swinhoe, 1895: 302.
Calamochrous carnealis (Swinhoe): Hampson, 1896: 420.
Type material. Type ♂, Khasi Hills., 95-224, Cherra Punji (
Other material examined. China. Guangdong: 6♂, Shimentai Reserve, Yingde, 27.V.2012, leg. Yang Lijun & Jia Qianju. Guizhou: 1♂, Maolan Reserve, 25.13°N, 107.87°E, alt. 797 m, 12.VII.2013, leg. Chen Xiaohua, genitalia slide no. SYSU0165, molecular voucher no. LEP0044; 1♀, Banzhai Village, Maolan Reserve, 25.23°N, 108.03°E, alt. 530 m, 11.VIII.2018, leg. Zheng Meiling et al. (
This species is similar to Loxoneptera triangularis in appearance, but can be distinguished by the following characters: forewing mixed with reddish brown scales, a distinct dark brown stripe appearing near posterior angle of cell; apex of hindwing with a dark brown patch; dorso-distal sella with a hook-shaped process; distal end of phallus with a spine-shaped process, longer than phallus length.
Head. Frons pale reddish brown, with white lateral bands. Vertex pale brown, mixed with some reddish brown erect scales. Labial palpus reddish brown, with white scales on ventral side. Maxillary palpus reddish brown, broadened distally with scales. Antennae yellowish brown. Thorax. Dorsal patagia and tegula ochre-brown, ventral side grey-white. Legs pale yellow to grey-white; hindleg basal outer spur 2/5 of basal inner spur. Wings. Wingspan 22.0–29.0 mm. Forewing yellowish brown, densely mixed with reddish brown scales; dark brown from costal margin to posterior margin of cell; costal margin white; orbicular stigma appearing as a black-brown point, reniform stigma black, appearing as a thick streak on discocellulars; a distinct dark brown stripe appearing near posterior angle of cell; fringe black-brown. Hindwing pale yellow, apex with a dark brown patch; fringe pale yellow. Underside of forewing pale yellow, black-brown from costal margin to posterior margin of cell. Abdomen. Dorsal side of abdomen black-brown, ventral side grey white; sternite VIII in male slightly sclerotised with two pointed anterolateral processes.
Male genitalia
(Fig.
Female genitalia
(Fig.
China (Guangdong, Guizhou, Yunnan), India.
Type material. Holotype, ♂, China: Yunnan: Mengla, Xishuangbanna, 7.X.2004, leg. R. L. Kitching, genitalia slide no. FCEL0004 (FCEL). Paratype: China: Yunnan: 1♂, Mengla, Xishuangbanna, 4.X.2004, leg. R. L. Kitching.
Externally, Loxoneptera triangularis resembles L. carnealis in the wing shape, but can be distinguish by the smaller wings, and costal and posterior areas of hindwing dark brown. In the male genitalia, it can be differentiated by the process on the dorso-distal sella with a strongly sclerotised stick, distal part of juxta with a strongly sclerotised and narrowly triangular process, and distal phallus with a relatively short and hook-shaped spine.
Head. Frons pale yellow, with white lateral bands, basal white bands mixed with reddish brown scales. Vertex pale yellow. Labial palpus reddish brown, ventral side with white scales. Maxillary palpus reddish brown, broadened distally with scales. Antennae yellowish brown. Thorax. Dorsal side, patagia and tegula yellowish brown, mixed with reddish brown scales, ventral side grey white. Legs pale yellow. Wings. Wingspan 23.0–25.0 mm. Forewing pale yellow, termen dark brown, as well as from costal margin to posterior margin of cell, apex mixed with reddish brown scales; orbicular stigma weak, appearing as a dark brown point, reniform stigma black-brown and weak; fringe dark brown. Underside of forewing black from costal margin to posterior margin of cell. Hindwing pale yellow between CuA2 and M2, remainders dark brown, without pattern, fringe yellow brown. Abdomen. Dorsal side of abdomen pale brown, ventral side grey white; sternite VIII in male slightly sclerotised with two stout anterolateral processes.
Male genitalia
(Fig.
Female genitalia. Unknown.
China (Yunnan).
The specific name derived from the Latin triangularis, referring to the triangular process in the end of juxta.
Type material. Holotype, ♂, China: Yunnan: Yexianggu, Xishuangbanna, 22.17°N, 100.87°E, alt. 762 m, 18.VII.2014, leg. Teng Kaijian et al., genitalia slide no. ZDD12059, molecular voucher no. LEP0170 (
Loxoneptera rectacerosa resembles L. medialis in wing pattern, but the forewing of L. rectacerosa is brown from the costal margin to posterior margin of the cell, and white on costal margin, whereas it is pale yellow in L. medialis. In the male genitalia, dorsal margin of valva of L. rectacerosa makes a turn in the end, forming a distinct obtuse subapical angle; the process of the dorso-distal sella is smaller and shorter than that of L. medialis; distal end of phallus has a small and triangular sclerite, vesica is just with a group of spines.
Head. Frons pale yellow, with white lateral bands. Vertex pale yellow. Labial palpus brown, with white scales on ventral side. Maxillary palpus brown, broadened distally with scales. Antennae yellowish brown. Thorax. Dorsal side, patagia and tegula yellowish brown, ventral side grey white. Legs white to yellowish white. Wings. Wingspan 29.0 mm. Forewing brown, mixed with reddish brown scales, costal margin white, posterior area pale yellow; orbicular stigma weak, appearing as a dark brown point, reniform stigma absent; fringe black-brown. Hindwing pale yellow, without any spot, apex mixed with a few pale brown scales. Underside of forewing black on cell. Abdomen. Dorsal side of abdomen pale brown, ventral side grey white; sternite VIII in male slightly sclerotised with two stout anterolateral processes.
Male genitalia
(Fig.
Female genitalia. Unknown.
China (Yunnan).
The specific name derived from the Latin rect- (straight) and arcerosus (spine-shaped), referring to the shape of cornuti in phallus.
Calamochrous medialis Caradja, 1925: 363.
Type material. Holotype, ♂, Canton, Type, Car.[adja], Gen. Praep.[Prep.] EGM 3 (
Other material examined. China. Guangdong: 1♂, Dongmei Village, Potou District, Zhanjiang, 10.IV.2016, leg. Li Zhiqiang & Li Jun, genitalia slide no. SYSU0987, molecular voucher no. LEP0171; 1♀, Liuzhang Village, Beihe Country, Leizhou, 9.IV.2016, leg. Li Zhiqiang & Li Jun, genitalia slide no. SYSU0990, molecular voucher no. LEP0173. Hainan: 1♂, Jianling Reserve, 18.87°N, 110.27°E, alt. 143 m, 8.IX.2013, leg. Chen Xiaohua, genitalia slide no. SYSU0180, molecular voucher no. LEP0096.
The wing shape of Loxoneptera medialis is similar to L. rectacerosa but can be distinguished by the light yellow forewing and costal margin. In the male genitalia, it can be distinguished by longer spinous process on dorso-distal sella, distal end of phallus with a small and pointed spine, and vesica with two groups of short, spine-shaped cornuti.
Head. Frons pale yellow, with white lateral bands. Vertex pale yellow. Labial palpus pale yellow, with white scales on ventral side. Maxillary palpus pale yellow, mixed with white scales, broadened distally with scales. Antennae yellowish brown. Thorax. Dorsal side, patagia and tegula yellowish brown, ventral side grey white. Legs yellowish white. Wings. Wingspan 25.0–30.0 mm. Forewing pale yellow, costal and terminal areas reddish brown; orbicular stigma weak, dark brown, reniform stigma weak, black-brown, appearing as a thick line on discocellulars; a weak, dark-brown stripe appearing between M2 and CuA1; fringe black-brown. Hindwing pale yellow, termen mixed with brown scales, without pattern. Underside of forewing pale yellow, without any spot. Abdomen. Dorsal side of abdomen black-brown, ventral side grey white; sternite VIII in male slightly sclerotised with bifurcate anterolateral processes.
Male genitalia
(Fig.
Female genitalia
(Fig.
China (Guangdong, Hainan).
Calamochrous pentasaris Meyrick, 1932: 317.
Type material. Holotype, ♂, [India] Datarpur, Hoshiarpur. Officer-in-charge, 21.12.1927, Pyralidae Brit. Mus. Slide No. 9747 (
Wingspan 28.0 mm. Loxoneptera pentasaris is best distinguished from other Loxoneptera species by greyish ochreous forewing with a white costal band, and without pattern. In the male genitalia, this species is similar to L. medialis in the shape of dorsal projection of transtilla, ventral sella and valva, but can be distinguished by the triangular dorsal sella, process of dorso-distal sella extending ventrad, distal margin of phallus densely decorated with short spines.
India.
Calamochrous bipunctalis Hampson, 1912: 1269.
Type material. Type ♂, S. India, Palani Hills [Palnis], Campbell 1907.365, Pyralidae Brit. Mus. Slide No. 9750 (
Wingspan 34.0 mm. In appearance, Loxoneptera bipunctalis is best distinguished from other Loxoneptera species by pale ochreous yellow forewing, two blackish orbicular stigmata, and interrupted postmedial line of forewing. In the male genitalia, this species is similar to L. brevipalpis and L. dichroma but can be distinguished by the longer process of dorso-distal sella and the weakly sclerotised, slice-shaped cornutus of phallus.
India.
Calamochrous brevipalpis Snellen, 1890: 599.
Type material. Holotype, ♂, Sikkim, O. Möller, Pyralidae Brit. Mus. Slide No. 9748 (
Wingspan 33.0 mm. This species is distinguished by dull luteous forewing suffused with ochreous scales and bearing indistinct orbicular and reniform stigmata, lustrous hindwing suffused with grey scales along the costa. In the male genitalia, this species is similar to L. dichroma in the shape of the dorsal projection of the transtilla, ventral sella and valva, as well as by the process of dorso-distal sella extended ventrad and beyond the ventral margin of valva. Loxoneptera brevipalpis can be distinguished by the thick and heavily sclerotised process of dorso-distal sella, and the heavily sclerotised, spiny, thumb-shaped cornutus.
India (Sikkim).
Ebulea dichroma Moore, 1888: 223.
Calamochrous dichroma (Moore): Snellen, 1890: 599.
Anania dichroma
(Moore):
Type material. Type ♂, Darjeeling [Darjiling], Pyralidae Brit. Mus. Slide No. 9749 (
Wingspan 34.0 mm. This species can be distinguished by having a brown stripe along posterior margin of the discal cell in forewing, and ventral-distal wall of phallus is weakly sclerotised and obliquely extended into a process.
India.
The monophyly of Loxoneptera is strongly supported by the results of the molecular analysis. The dorsal projection of the transtilla in the male genitalia is a putative synapomorphy for the genus. It is shared by eleven species of Loxoneptera and can be used to separate them from most other pyraustine genera. In addition, two provisional infrageneric groups of the species of Loxoneptera are recognised by proportional lengths of the dorsal projection of the transtilla with its distal hair. The tree topology (Fig.
After examining three specimens of Calamochrous chilonalis (the type species of Calamochrous) deposited in
Grateful thanks to Prof. Houhun Li (Nankai University, China) and Prof. Akihiro Nakamura (Forest Canopy Ecology Lab, Yunnan, China) for the loan of some specimens, to Dr. David Lees and Dr. Geoff Martin (both Natural History Museum, United Kingdom) and Dr. Mihai Stănescu (“Grigore Antipa” National Museum of Natural History, Romania) for helping to access specimens deposited at their institutions. We are also grateful to Dr. Robert B. Angus (Natural History Museum, London, United Kingdom) for critical reviews of the manuscript and for linguistic assistance. This project was supported by the National Natural Science Foundation of China (Grant No. 31672330), Program of the Ministry of Science and Technology of the People of Republic of China (2015FY210300).