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Research Article
The genus Dryadaula Meyrick (Lepidoptera, Tineoidea, Dryadaulidae) in China, with descriptions of four new species and a world checklist
expand article infoLin-Lin Yang‡, Hou-Hun Li§
‡ Institute of Plant Protection, Henan Academy of Agricultural Sciences, Zhengzhou, China
§ Nankai University, Tianjin, China

Corresponding author: Hou-Hun Li ( lihouhun@nankai.edu.cn )

Academic editor: Andrew Mitchell
© 2021 Lin-Lin Yang, Hou-Hun Li.
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: Yang L-L, Li H-H (2021) The genus Dryadaula Meyrick (Lepidoptera, Tineoidea, Dryadaulidae) in China, with descriptions of four new species and a world checklist. ZooKeys 1074: 61-81. https://doi.org/10.3897/zookeys.1074.73067
ZooBank: urn:lsid:zoobank.org:pub:A921393D-58B0-48E1-8F10-1A9A163D8B91
Open Access

Abstract

Four new species of the genus Dryadaula Meyrick, 1893 from China are described: Dryadaula auriformis sp. nov., D. flavostriata sp. nov., D. hirtiglobosa sp. nov. and D. securiformis sp. nov. Photographs of adults and genitalia of the new species are provided. DNA barcodes of D. auriformis sp. nov., D. hirtiglobosa sp. nov. and D. securiformis sp. nov. are given. A key to the species in China and a detailed checklist for the genus with all 49 known species in the world are presented.

Keywords

Asymmetrical genitalia, COI, morphology, taxonomy

Introduction

The family Dryadaulidae was proposed by Regier et al. (2015) on the basis of a molecular phylogenetic study of the Tineoidea. It currently includes two genera: Dryadaula Meyrick, 1893 (45 species with worldwide distribution) and Brachydoxa Meyrick, 1917 (two species distributed in the Oriental Region). The New Zealand genera Eschatotypa Meyrick, 1880 (three species), Eugennaea Meyrick, 1915 (one species) and Sagephora Meyrick, 1888 (six species) were also considered to belong to this group (Robinson and Nielsen 1993).

The genus Dryadaula was established by Meyrick (1893) with D. glycinopa Meyrick, 1893 as the type species. It comprises 45 species, distributed in all zoogeographical regions. Most of these species have been placed in subfamilies of Tineidae when originally described. Dryadaula was introduced as the senior name of Thermocrates Meyrick, 1936 by Robinson (1988) and of Archimeessia Zagulajev, 1970, Chorocosma Meyrick, 1893, Cyane Chambers, 1873, Choropleca Durrant, 1914, Diachalastis Meyrick, 1920, Ditrigonophora Walsingham, 1897, Opsodoca Meyrick, 1919 and Strophalinga Gozmány & Vári, 1973 by Robinson and Nielsen (1993). The main reason for this situation is that Dryadaula is difficult to diagnose externally. However, significant features can be seen when members of Dryadaula are dissected: segment VIII and genitalia are strongly modified and asymmetrical in the male, the oviscapt is greatly reduced and the anterior apophyses are rudimentary or absent in the female.

Before this study, only one species D. epischista (Meyrick, 1936) in the genus Dryadaula was reported from Hong Kong, China (Robinson 1988). We herein describe four new species in China, with illustrations of adults and genitalia and provide a key to the known Chinese species. A world checklist of the genus is also provided to facilitate the further study.

Materials and methods

The holotypes of D. flavostriata sp. nov. and D. hirtiglobosa sp. nov. were collected using sweep nets in the daytime; other type specimens were collected under 250-W high-pressure mercury lamps on white sheets at night. The type specimens are deposited in the Insect Collection, College of Life Sciences, Nankai University, Tianjin, China (NKU).

Genitalia dissection and mounting methods follow Li (2002), while head and wing dissections were carried out following the methods described by Lee and Brown (2006). Photographs of the adults were taken with a Leica M205A stereomicroscope, and those of genitalia were taken with a Leica DM750 microscope plus Leica Application Suite 4.6 software. All photographs were refined with Photoshop CS5 software. Protocols for total DNA extraction and mitochondrial COI gene amplification followed that described in our previous study (Yang and Li 2021). Degrees of intra- and interspecific variation of DNA barcode fragments were calculated under the Kimura 2-parameter model using MEGA X. Terminology used in the description of the adult, vein venation and male genitalia follows Robinson and Nielsen (1993) and that of female genitalia follows Regier et al. (2015).

Abbreviations used in the text are as follows

ANIC Australian National Insect Collection, CSIRO Division of Entomology, Canberra, Australia;

BPBM Bernice Pauahi Bishop Museum, Honolulu, Hawaii, United States of America;

coll. Baldizzone collection of Giorgio Baldizzone, Asti Italy;

coll. Heindel collection of Richard Heindel, Günzburg, Germany;

coll. Scholz collection of Axel Scholz, Illerberg, Germany;

coll. Sutter collection of Reinhard Sutter, Bitterfeld, Germany;

LMNH Latvian Museum of Natural History, Riga, Latvia;

MCZ The Museum of Comparative Zoology, Harvard University, United States of America

NHM The Natural History Museum, London, United Kingdom;

NKU Insect Collection, College of Life Sciences, Nankai University, Tianjin, China;

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany [former: IPE: Institut für Pflanzenschutzforschung, Eberswalde, Germany; and DEI: Deutsches Entomologisches Institut, Deutsche Akademie der Landwirtschaftswissenschaften zu Berlin, Eberswalde, Germany];

SEL/HNU Systematic Entomology Laboratory, Hannam University, South Korea;

TL Type locality;

TM Transvaal Museum, Pretoria, The Republic of South Africa;

ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia;

ZMHB Museum für Naturkunde der Humboldt-Universität (Wolfram Mey), Berlin, Germany;

ZMUC Zoological Museum, University of Copenhagen, Denmark.

Results

DNA Barcoding

The holotype of Dryadaula auriformis sp. nov., a paratype of D. hirtiglobosa sp. nov. and two paratypes of D. securiformis sp. nov. were successfully sequenced and yielded a barcode of 604 bp. Complementary public sequences of D. heindeli Gaedike & Scholz (BOLD:AAL1778, n = 4), D. terpsichorella (Busck) (BOLD:AAF9987, n = 139) and D. visaliella (Chambers) (BOLD:ACA7671, n = 57; BOLD:AAV6731, n = 17; BOLD:AAV6730, n = 10) from BOLD systems were used to calculate the genetic distance barcode divergence. Sequence divergences are presented in Table 1. The sampled specimens of D. visaliella within three Barcode Identification Numbers (BINS) might represent different species, as members of them show higher divergences from each other and were not well distinguished by barcodes.

Table 1.
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Percentage of divergence in the cytochrome c oxidase subunit I (COI) gene sequences of the Dryadaula species.

1 2 3 4 5 6 7 8
1 D. auriformis sp. nov. – – – – – – – –
2 D. hirtiglobosa sp. nov. 18.55 – – – – – – –
3 D. securiformis sp. nov. 19.18 6.06 0 – – – – –
4 D. heindeli 17.74–17.95 13.61–13.81 14.31–14.81 0–0.17 – – – –
5 D. terpsichorella 18.64–19.83 10.41–12.05 9.97–11.92 9.97–10.73 0–0.35 – – –
6 D. visaliella (ACA7671) 15.20–21.36 9.33–10.79 10.00–12.79 11.04–13.85 9.33–12.49 0–1.98 – –
7 D. visaliella (AAV6731) 18.74–22.70 10.33–11.26 10.62–11.64 15.29–18.58 11.12–14.49 3.34–13.27 0–0.51 –
8 D. visaliella (AAV6730) 20.57–21.22 9.20–9.62 10.22–10.99 13.18–14.43 10.19–11.89 6.93–9.65 11.51–13.67 0–0.34

Genetic distances (%) were corrected with the Kimura two-parameter (K2P) substitution model using MEGA X; extreme values of intraspecific and interspecific distances are given (the numbers in bold are the intraspecific distances).

Taxonomic accounts

Dryadaula Meyrick, 1893

Dryadaula Meyrick, 1893: 559. Type species: Dryadaula glycinopa Meyrick, 1893, by monotypy. TL: Australia (New South Wales).

Cyane Chambers, 1873: 112. Synonymised by Robinson and Nielsen 1993: 55. Type species: Cyane visaliella Chambers, 1873, by monotypy. TL: United States (Kentucky).

Chorocosma Meyrick, 1893: 560. Synonymised by Robinson and Nielsen 1993: 55. Type species: Chorocosma melanorma Meyrick, 1893, by monotypy. TL: Australia (Sydney).

Ditrigonophora Walsingham, 1897: 117. Synonymised by Robinson and Nielsen 1993: 55. Type species: Ditrigonophora marmoreipennis Walsingham, 1897, by original designation. TL: Grenada (Balthazar).

Choropleca Durrant, 1914: 366. Objective replacement name for Cyane Chambers, 1873. Synonymised by Robinson and Nielsen 1993: 55.

Opsodoca Meyrick, 1919: 270. Synonymised by Robinson and Nielsen 1993: 55. Type species: Opsodoca metrodoxa Meyrick, 1919, by original designation. TL: Guyana.

Diachalastis Meyrick, 1920: 363. Synonymised as Choropleca Durrant by Clarke, 1971: 221. Synonymised by Robinson and Nielsen 1993: 55. Type species: Diachalastis tetraglossa Meyrick, 1920, by monotypy. TL: Fiji.

Thermocrates Meyrick, 1936: 620. Synonymised by Robinson, 1988: 74. Type species: Thermocrates epischista Meyrick, 1936, by monotypy. TL: Japan (Kyushu).

Archimeessia Zagulajev, 1970: 658. Synonymised by Robinson and Nielsen 1993: 55. Type species: Archimeessia zinica Zagulajev, 1970, by original designation. TL: Azerbaijan.

Strophalinga Gozmány & Vári, 1973: 9. Synonymised by Robinson and Nielsen 1993: 55. Type species: Tinea glycinocoma Merick, 1932, by original designation. TL: Ethiopia.

Diagnosis

Dryadaula is a small-sized moth, with wingspans of no more than 20 mm. It can be recognised by the following characters: head (Figs 1a–4a, 5) with erect piliform scales, transfrontal suture inverted V-shaped; antennae 0.7× length of forewing, scape without pecten, flagellomeres with single annulus or two annuli of contrasting-coloured scales; labial palpus spatulate, bearing lateral bristles; forewing often brightly coloured, with venation (Fig. 6) complete, CuP weak; hind-wing with M3 or CuA1 absent; female with single frenulum bristle; segment VIII reduced and highly modified, usually asymmetrical in male; male genitalia (Figs 7–10) strongly asymmetrical, incorporating part of sternum VII and sternum VIII; aedeagus fused with right valva; gnathos absent; uncus lobes usually fused; female (Fig. 11) oviscapt reduced, posterior apophyses short, anterior apophyses rudimentary or absent, sternum VIII hardly developed.

Distribution

Worldwide; the distribution of each species is given in Table 2.

Table 2.
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World checklist of the genus Dryadaula Meyrick, 1893.

Species Distribution Depository of type
1 acrodisca (Meyrick, 1917): 79. (Choropleca) Guyana NHM (LT)
TL: Guyana (Mallali).
2 amentata (Meyrick, 1919): 271. (Opsodoca) Guyana NHM (HT)
TL: Guyana (Bartica). Figs: Clarke (1970: pl. 34, fig. 2 adult and male genitalia).
3 advena (Zimmerman, 1978): 326. (Choropleca) United States BMH (HT)
TL: United States (Hawaii). Figs: Zimmerman (1978: fig. 156-A male genitalia; fig. 481-A adult).
4 anthracorma Meyrick, 1915: 369. Australia NHM (LT and PLT)
TL: Australia (Victoria). Figs: Robinson and Nielsen (1993: fig. 63 adult; fig. 70 male genitalia; figs 71, 72 female genitalia); Robinson (2009: fig. 2 adult).
5 auriformis sp. nov. China NKU (HT and PT)
TL: China (Hainan). Figs 1, 7.
6 boviceps (Walsingham, 1914): 366. (Choropleca) Mexico NHM (HT)
TL: Mexico (Guerrero). Figs: Robinson (2009: fig. 4 adult).
7 brontoctypa (Meyrick, 1880): 259. (Ereunetis) Australia NHM (LT and PLT)
TL: Australia (Sydney).
8 castanea Philpott, 1915: 201. New Zealand ?
TL: New Zealand (Bluff, Invercargill).
9 catorthota (Meyrick, 1917): 80. (Choropleca) Guyana NHM (LT and PLT)
TL: Guyana (Mallali).
10 caucasica (Zagulajev, 1970): 662. (Archimeessia) Azerbaijan, Poland, Russia, Sweden ZIN (HT)
TL: Azerbaijan (Artschevan). Figs: Zagulajev (1970: fig. 6 female genitalia); Zagulajev (1979: fig. 64 adult; fig. 65 female genitalia); Sachkov (1995: fig. 7 male genitalia); Jaworski et al. (2012: fig. 1 adult); Gaedike (2015: pl. 1, fig. 1 adult; drawings, male genitalia 1; drawings, female genitalia 1).
11 discatella (Walker, 1864): 1021. (Gelechia) Brazil NHM (HT)
TL: Brazil.
12 epischista (Meyrick, 1936): 621. (Thermocrates) China (Hong Kong), Japan NHM (HT)
TL: Japan (Kyushu). Figs: Robinson (1988: fig. 1 adult; fig. 2 abdominal pelt; figs 3, 4 male genitalia); Sakai (2013: fig. 3–12–13 adult).
13 epixantha (Turner, 1923): 184. (Erechthias) Australia ANIC
TL: Australia (Queensland).
14 flavostriata sp. nov. China NKU (HT)
TL: China (Guangxi). Figs 2, 8.
15 germana (Walsingham, 1914): 367. (Choropleca) Mexico NHM (HT)
TL: Mexico (Guerrero).
16 glycinocoma (Meyrick, 1932): 120. (Tinea) Ethiopia NHM (LT and PLT)
TL: Ethiopia. Figs: Gozmány and Vári (1973: fig. 6 male genitalia).
17 glycinopa Meyrick, 1893: 559. Australia NHM (LT)
TL: Australia (New South Wales). Figs: Robinson and Nielsen (1993: fig. 64 adult; fig. 136 wing venation).
18 heindeli Gaedike & Scholz, 1998: 106. Belgium, France, Germany, Italy, Netherlands, Norway, Spain, Switzerland SDEI (HT and PT); coll. Scholz (PT); coll. Heindel (PT); coll. Sutter (PT)
TL: Germany (Bayem). Figs: Gaedike and Scholz (1998: fig. 1 adult; figs 3–6 male genitalia; fig. 7 female genitalia; figs 10–15 larva, chaetotaxy and pupa); Gaedike (2015: pl. 1, fig. 5 adult; drawings, male genitalia 5; drawings, female genitalia 5).
19 hellenica (Gaedike, 1988): 331. (Archimeessia) Greece ZMUC (HT and PT); SDEI (PT).
TL: Greece (Peloponnese). Figs: Gaedike (1988: figs 22–26 male genitalia); Gaedike (2015: pl. 1, fig. 7 adult; drawings, male genitalia 7; drawings, female genitalia 7).
20 hirtiglobosa sp. nov. China NKU (HT and PT)
TL: China (Guangxi). Figs 3, 9.
21 irinae (Savenkov, 1989): 94. (Archimeessia) Austria, Bulgaria, Latvia, Poland, Slovakia, LMNH
TL: Latvia. Figs: Savenkov (1989: figs. 1–3); Pastorális et al. (2011: figs 1, 2 adults; fig. 3 male genitalia); Jaworski et al. (2014: fig. 5 adult; fig. 6 larva; fig. 7 larval shelter; fig. 8 pupal case); Gaedike (2015: pl. 1, fig. 3 adult; drawings, male genitalia 3; drawings, female genitalia 3).
22 isodisca (Meyrick, 1917): 80. (Choropleca) Guyana NHM (LT)
TL: Guyana (Bartica, Mallali).
23 koreana Roh & Byun, 2020: 222 South Korea SEL/HNU (HT and PT)
TL: South Korea (Jeollanam-do). Figs: Roh et al. (2020: figs. 1, 2 adult; fig. 3: wing venation; fig. 4 male genitalia; fig. 5 female genitalia).
24 marmoreipennis (Walsingham, 1897): 118. (Ditrigonophora) Grenada NHM (LT)
TL: Grenada (Balthazar).
25 melanorma (Meyrick, 1893): 560. (Chorocosma) Australia NHM (HT)
TL: Australia (Sydney).
26 mesosticha (Turner, 1923): 184. (Erechthias) Australia ANIC
TL: Australia (Queensland).
27 metrodoxa (Meyrick, 1919): 271. (Opsodoca) Guyana NHM (LT)
TL: Guyana (Bartica, Mallali). Figs: Clarke (1970: pl. 34, fig. 1 adult, wing venation and male genitalia).
28 minuta Gaedike, 2007: 160. Greece, Turkey ZMHB (HT), ZMUC (PT)
TL: Turkey (Mugla). Figs: Gaedike (2007: fig. 1 adult; figs 12, 13 male genitalia; fig. 14 female genitalia); Gaedike (2015: pl. 1, fig. 4 adult; drawings, male genitalia 4; drawings, female genitalia 4).
29 multifurcata Gaedike, 2000: 358. Russia ZIN (HT and PT)
TL: Russia (Primorskij kraj). Figs: Gaedike (2000: figs 5–7 male genitalia; fig. 8 female genitalia).
30 murenula (Meyrick, 1924): 65. (Choropleca) Peru NHM (LT and PLT)
TL: Peru (Jurimaguas, Iquitos).
31 myrrhina Meyrick, 1905: 243. New Zealand NHM (HT)
TL: New Zealand. Robinson (2009: fig. 3 adult).
32 napaea Meyrick, 1905: 244. Australia NHM (HT)
TL: Australia (Tasmania).
33 nedae (Gaedike, 1983): 125. (Infurcitinea) Croatia, Cyprus, Greece, Turkey coll. Baldizzone (HT and PT); SDEI (PT)
TL: Greece. Figs: Baldizzone (1983: figs 3–7 male genitalia; fig. 8 female genitalia); Gaedike (2015: pl. 1, fig. 8 adult; drawings, male genitalia 8; drawings, female genitalia 8).
34 pactolia Meyrick, 1901: 577. Denmark, France, Germany, Great Britain, New Zealand, Netherlands, Portugal, Switzerland. NHM (LT)
TL: New Zealand. Figs: Gaedike and Scholz (1998: fig. 2 adult; figs 8, 9 male genitalia); Gaedike (2015: pl. 1, fig. 6 adult; drawings, male genitalia 6; drawings, female genitalia 6).
35 panscia Meyrick, 1917: 81. (Choropleca) Ecuador NHM (LT)
TL: Ecuador (Huigra).
36 placens Meyrick, 1920: 363. (Hectacma) Australia NHM (HT)
TL: Australia (Queensland).
37 poecilta Walsingham, 1914: 366. (Choropleca) Mexico NHM (HT)
TL: Mexico (Guerrero).
38 rhombifera Meyrick, 1917: 82. (Choropleca) Guyana NHM (HT)
TL: Guyana (Mallali).
39 securiformis sp. nov. China NKU (HT and PT)
TL: China (Hainan). Figs 4, 5, 6, 10 and 11.
40 selenophanes (Meyrick, 1880): 259. (Ereunetis) Australia NHM (LT)
TL: Australia (Queensland).
41 sublimis (Meyrick, 1917): 81. (Choropleca) Colombia NHM (LT)
TL: Colombia (La Crumbre).
42 terpsichorella (Busck, 1910): 134. (Cyane) Fiji, Hawaii, Rapa. NHM (HT)
TL: Hawaii (Honolulu). Figs: Clarke (1971: pl. 28, figs. a, b adult; fig. 172 male genitalia, coremata and 8th segment; fig. 173 wing venation and female genitalia); Zimmerman (1978: fig. 157 head and wing venation; fig. 159 adult, male and female genitalia; fig. 160 abdomen and male genitalia); Robinson (2009: fig. 5 adult).
43 trapezoides (Meyrick, 1935): 579. (Tinea) Japan NHM (LT)
TL: Japan (Tokyo). Figs: Sakai (2013: figs 3–12–14 adult; fig. Tin12 female genitalia).
44 tripudians (Meyrick, 1924: 65). (Choropleca) Peru NHM (LT)
TL: Peru (Jurimaguas).
45 ussurica Gaedike, 2000: 358. Russia ZIN (HT and PT)
TL: Russia (Primorskij kraj). Figs: Gaedike (2000: figs. 1–3 male genitalia; fig. 4 female genitalia).
46 visaliella (Chambers, 1873): 113. (Cyane) Canada, United States MCZ (ST)
TL: United States (Kentucky). Figs: Zimmerman (1978: fig. 158 wing venation); Regier et al. (2015: fig. 6F adult; fig. 10 head, wing venation, male and female genitalia).
47 zinica (Zagulajev, 1970: 661). (Archimeesia) Azerbaijan, Russia ZIN (HT and PT); NHM (PT)
TL: Azerbaijan. Figs: Zagulajev (1970: fig. 1 head; fig. 2 wing venation; fig. 3 legs; fig. 4 male genitalia; fig. 5 female genitalia); Gaedike (2015: pl. 1, fig. 2 adult; drawings, male genitalia 2; drawings, female genitalia 2).
48 zygodes Meyrick, 1918: 44. (Tinea) South Africa TM (HT)
TL: South Africa (Natal). Figs: Janse (1968: pl. 69, fig. 3 adult, fig. 4 male genitalia; pl. 109: fig. 7 wing venation; pl. 111, fig. 16 labial palpi and maxillary palpi; pl. 113, fig. 9 male genitalia); Gozmány and Vári (1973: fig. 58 male genitalia)
49 zygoterma Meyrick, 1917: 82. (Choropleca) Colombia, Ecuador NHM (LT)
TL: Colombia (La Crumbre).

Biology

The larvae of some species are detritivores or feed on lichens and fungi. The biology of Dryadaula was reviewed or summarised by Robinson and Nielsen (1993), Gaedike (2015) and Regier et al. (2015).

Key to Chinese Dryadaula species, based on the male

1 Forewing dull ochreous brown patterned with yellow-brown and white streaks (Robinson 1988: fig. 1) D. epischista
– Forewing white patterned with black spots or patches 2
2 Subscaphium absent 3
– Subscaphium present 4
3 Uncus lobes without process; vinculum without additional lobe; left valva clavate; right valva with dorsal lobe globular apically, ventral lobe with three prominences (Fig. 7) D. auriformis sp. nov.
– Uncus lobes with a rectangular process; vinculum with a lobe; left valva sub-oval; right valva with dorsal lobe vaulted, having a finger-like process apically, ventral lobe digitate, without prominence (Fig. 8) D. flavostriata sp. nov.
4 Modification attached to vinculum is receptacle-shaped, with a sharp horn and a drumstick-like process; left valva irregular in shape; right valva with dorsal lobe having a subquadrate and a digitate process, ventral lobe crescent; juxta pocket-like (Fig. 9) D. hirtiglobosa sp. nov.
– Modification attached to vinculum comprising of a Y-shaped sclerotisation and a receptacle-shaped sclerite; left valva battle axe-shaped; right valva with dorsal and ventral lobes slender, S-shaped; juxta elliptical (Fig. 10) D. securiformis sp. nov.

Dryadaula auriformis sp. nov.

http://zoobank.org/816AD20C-6E06-4A79-885F-9B8A00F93290
Figures 1, 7

Type material

Holotype : China: • ♂; Hainan Province, Mt. Jianfeng (18°44'N, 108°52'E); alt. 787 m; 1.vi.2015; leg. Peixin Cong; genitalia slide No. DNAYLL18124. Paratype: China: • 1 ♂; Hainan Province, Mt. Jianfeng; alt. 745 m; leg. Xia Bai; genitalia slide No. XMR18217.

Differential diagnosis

The new species is externally similar to D. zinica (Zagulajev, 1970), but can be separated from it by the male genitalia structures. In D. auriformis sp. nov., the left valva is narrowed and clavate, the right valva is bilobate, the bullet-like sternum VIII is smooth in the male genitalia, whereas in D. zinica, the left valva is broad, the right valva is not divided and the sternum VIII bears long and thin bristles on outer margin.

Description

Adult (Fig. 1): Wingspan 8.5 mm in holotype, 9.0 mm in paratype. Vertex and frons smoky grey, tinged with black scales anterior of antenna. Antenna with scape white, except for a black spot at dorsal base; flagellum with alternate yellowish-white and cinereous annulations, cinereous on dorsal surface of basal 2–4 flagellomeres, with three narrow cinereous bands towards apex. Labial palpus spatulate; yellowish-white, first palpomere and basal 3/4 of second palpomere black on outer surface, third palpomere black at base on inner surface, with three black dots on outer surface. Thorax and tegula blackish-brown in anterior 1/2, white in posterior 1/2. Forewing ground colour white, irrorate with blackish-brown scales, edged with bright ochreous yellow scales along of termen and markings; patterned with black markings: costa with a wedge-shaped spot at base, a rectangular spot at 2/5, an obscure dot at middle, an oval patch from 3/5 to 4/5; cell with an obscure irregular spot at distal 1/4, tending to coalesce with oval costal patch; fold with irregular stripes at base, basal 1/3 and 2/3, obliquely inward towards dorsum; an interrupted terminal line around apex then along termen to tornus; cilia white in basal 1/2, grey in distal 1/2, with individual scales dark-tipped. Hind-wing and cilia grey. Legs greyish-white, tibia black on outer surface, tarsus black on dorsal surface, except for end of each tarsomere.

Male genitalia (Fig. 7). Uncus lobes small, ear-shaped, bearing dense setae dorso-apically. Subscaphium not developed. Tegumen somewhat broad. Vinculum narrowed, deeply arched at middle, without additional lobe. Saccus not developed. A complicated, irregular, sclerotised modification attached to vinculum anteriorly, possibly part of segments VII and VIII; its left part rectangular, with a stout digitation, its right part stem-like. Sternum VIII articulated with vinculum at left, articulated with left valva dorso-basally; somewhat bullet-like, narrowly rounded and folded apically, triangularly folded at 1/3 on ventral margin. Valvae strongly asymmetrical. Left valva clavate, bent outwards; its basal part skirt-like, arched anteriorly, distal part a globular, setose lobe, a small, digital, setose lobe at distal 1/3. Right valva bilobate: dorsal lobe with a thumb-like process articulated with juxta at base, middle part curved like a gooseneck, distal part globular, setose; ventral lobe with three prominences, one stout, finger-like, one slightly twisted, horn-shaped and one hammer-shaped. Juxta irregular in shape. Aedeagus a curved horn with a stout base; cornutus absent.

Figures 1–4. 

Adults of Dryadaula species 1 Dryadaula auriformis sp. nov., male holotype 1a lateral view of head 2 D. flavostriata sp. nov., male holotype 2a lateral view of head 3 D. hirtiglobosa sp. nov., male holotype 3a lateral view of head 4 D. securiformis sp. nov., male holotype 4a lateral view of head. Scale bars: 1.0 mm.

Female. Unknown.

Distribution

China (Hainan).

Etymology

The specific name is derived from the Latin auriformis, meaning ear-shaped, referring to the ear-shaped uncus lobes.

DNA barcode

One DNA barcode from the holotype was generated and deposited in GenBank and BOLD systems: MZ711361/ DRYAD001-21. Dryadaula auriformis sp. nov. is clearly distinguishable by its DNA barcode from its congeners, the minimum divergence (Table 1) to the nearest species, D. heindeli Gaedike & Scholz, is 17.74–17.95%.

Dryadaula flavostriata sp. nov.

http://zoobank.org/59375061-706B-4229-A4C2-B919D4DA48AB
Figures 2, 8

Type material

Holotype : China: • ♂; Guangxi Province, Nanning City, Mt. Daming (23°24'N, 108°30'E); alt. 1250 m; 23.v.2011; leg. Linlin Yang & Yinghui Mou; genitalia slide No. YLL11112.

Differential diagnosis

Dryadaula flavostriata sp. nov. is similar to D. caucasica (Zagulajev, 1970), but differs from it by the forewing peppered with more ochreous yellow scales that form stripes between fold and dorsum; the male genitalia with short uncus that is equipped with a rectangular process at the left and the sub-oval left valva with processes of different shapes, not bearing thorns or long bristles. In D. caucasica, the forewing has dark grey-brown longitudinal stripes between fold and dorsum; in the male genitalia, the elongate uncus has no process and the fluted left valva has dense, long bristles along outer ventral margin and a globular sclerotisation which is densely thorned.

Description

Adult (Fig. 2): Wingspan 11.5 mm in holotype. Vertex and frons yellowish-white. Antenna with scape brightly white, margined with black anteriorly; flagellum with dorsal surface alternating black and white in basal 1/3, black in medial 1/3, alternating black and white every two annuli in distal 1/3, ventral surface white in basal 2/3, alternating black and white every two annuli in distal 1/3. Labial palpus spatulate; first palpomere black, second palpomere white on inner surface, black on outer surface; third palpomere white tinged ochreous yellow, with a black dot at middle of outer surface. Thorax and tegula blackish-brown in anterior 1/2, brightly white tinged with blackish-brown in posterior 1/2. Forewing brightly white, with scattered grey and black scales; patterned with black markings that are bordered with ochreous yellow and ochreous yellow markings that are tinged with black: costa with a wedge-shaped spot at base, a semicircular spot at 1/3, a semicircular patch at 3/4 and an arc line at apex, diffused greyish-black smudges amongst spots; cell with a narrowed, oval spot at distal 1/4, suffused with ochreous yellow anterolaterally, tending to coalesce with semicircular costal patch; fold with irregular ochreous yellow stripes at base, basal 1/3 and 2/3, tinged with black, obliquely inwards towards dorsum; four or five black dots from apex to tornus along termen, forming a broken terminal line; cilia white in basal 1/2, grey in distal 1/2. Hind-wing and cilia dark grey. Foreleg dark grey, tarsus yellowish-white on inner surface; mid-leg femur greyish-black on outer surface, pale yellow on inner surface, tibia and tarsomeres yellowish-white at end; hind-leg yellowish-white on inner and ventral surface, tibia grey on outer and dorsal surface, tarsus greyish-black on outer and dorsal surface, yellowish-white at end of each tarsomere.

Figures 5, 6. 

Dryadaula securiformis sp. nov., male paratype, slide No. DNAYLL18121 5 head 6 Wing venation. Scale bars: 0.2 mm (head); 1.0 mm (wing venation).

Figures 7, 8. 

Male genitalia of Dryadaula species 7 D. auriformis sp. nov., holotype, slide No. DNAYLL18124 8 D. flavostriata sp. nov., holotype, slide No. YLL11112. Scale bars: 0.25 mm. (U = uncus lobes; T = tegumen; V = vinculum; S = saccus; S VIII = sternum VIII; Vr = right valva; Vl = left valva; J = juxta; A = aedeagus).

Male genitalia (Fig. 8). Uncus lobes fused into a shield, bearing long hairs dorsally, slightly concave at middle on posterior margin, asymmetrical, left lobe slightly longer than right, with a smooth rectangular process. Subscaphium not developed. Tegumen simple, forming a narrow ring with vinculum. Vinculum broad, posterior margin strongly sclerotised, with a setose lobe at right. Saccus subtriangular, asymmetrical. A complicated, irregular, sclerotised modification attached to vinculum anteriorly, possibly part of segments VII and VIII; its left part somewhat triangular, right part chestnut-shaped. Sternum VIII articulated with vinculum at left, articulated with left valva dorso-basally, strongly modified into a large pocket, broadly rounded apically, with a large hammer-like process at middle, with a tuft of non-deciduous hairs. Valvae strongly asymmetrical. Left valva larger than right one, sub-oval; costal margin with a broad, vertical bridge at middle, a digitate process at distal 1/3, a larger subrectangular process at 1/6, a slender, curved process at end that is about 2/3 length of valva, with a setose fan-shaped lobe at middle; ventral margin with an oblique, digitate process at middle and a horned process at end. Right valva divided into two parts: dorsal lobe large, vaulted, with a finger-like process apically and a slender, decurved, setose lobe at middle; ventral lobe digitate, hooked apically. Juxta inflated and rounded, with a U-shaped process on left, an arced band on right. Aedeagus as long as saccus, simple, expanded in basal 1/4, sinuate in S-shape in distal 3/4; cornutus absent.

Female. Unknown.

Distribution

China (Guangxi).

Etymology

The specific name is derived from the Latin prefix flav-, meaning yellowish and the Latin word striatus, stripe, referring to the forewing with inwardly oblique ochreous yellow stripes between fold and dorsum.

Dryadaula hirtiglobosa sp. nov.

http://zoobank.org/3BC5ADCC-97DA-4685-B9C4-39D6C013B943
Figures 3, 9

Type material

Holotype : China: • ♂; Guangxi Province, Nanning City, Mt. Daming (23°24'N, 108°30'E); alt. 1250 m; 23.v.2011; leg. Linlin Yang & Yinghui Mou; genitalia slide No. YLL13026. Paratypes: China: • 1 ♂; Zhejiang Province, Jingning She Autonomous County, Wangdongyang Wetland Reserve (27°24'N, 119°23'E); alt. 1174 m; 16.viii.2018; leg. Shuai Yu et al.; genitalia slide No. DNAYLL18170 • 1 ♂; Zhejiang Province, Jiangshan City, Mt. Xianxia, Shuangxikou Town, Laofoyan Village (28°22'N, 118°40'E); alt. 400 m; 26.v.2017; leg. Shuonan Qian and Jiaen Li; genitalia slide No. DNAYLL18169.

Figures 9, 10. 

Male genitalia of Dryadaula species 9 D. hirtiglobosa sp. nov., holotype, slide No. YLL13026 10 D. securiformis sp. nov., paratype, slide No. DNAYLL18173. Scale bars: 0.25 mm. (U = uncus lobes; T = tegumen; V = vinculum; Su = Subscaphium ; S VIII = sternum VIII; Vr = right valva; Vl = left valva; J = juxta; A = aedeagus).

Differential diagnosis

The new species is externally close to D. irinae (Savenkov, 1989), but differs from it by the shape of valva in the male genitalia: in D. hirtiglobosa sp. nov., the left valva has a blade-shaped setose lobe apically and a sword hilt-like process subapically on ventral margin and the right valva is bilobate; in D. irinae, the left valva is divided into two parts, without blade-shaped setose lobe or sword hilt-like process, the right valva is not bilobate.

Description

Adult (Fig. 3): Wingspan 11.0 mm in holotype, 9.5–10.5 mm in paratypes. Vertex cream white, frons pale greyish-brown. Antenna with scape brightly white, margined with black anteriorly; flagellum with ventral surface bearing white scales on alternate annuli, dorsal surface alternating black and white in basal 2/3, white with three black bands in distal 1/3. Labial palpus spatulate; smoky white on inner surface, black on outer surface, except for yellowish-white at end of third palpomere. Thorax and tegula blackish-brown in anterior 1/2, white tinged with blackish-brown in posterior 1/2. Forewing white, with scattered black and brown scales, patterned with black markings that are edged with ochreous yellow: costa with a wedge-shaped stripe in basal 1/6, a semicircular spot at 1/3, an ambiguous dot at middle, a semicircular patch at 3/4, an obscure, irregular greyish fuscous dot near apex and a dot at apex; cell with a narrowed, oval spot at distal 1/3, more or less coalesced with semicircular costal patch; fold with irregular stripes at base, basal 1/3 and 2/3, powdered with ochreous yellow scales, obliquely inwards towards dorsum; terminal line black, not continuous, around apex, then along termen to tornus; cilia greyish-white, with some black dots. Hind-wing and cilia dark grey. Foreleg greyish-black, tarsomeres white at end; mid-leg femur greyish-black on outer surface, pale yellow on inner surface, tarsomeres yellowish-white at end; hind leg with tibia yellowish-white on inner and ventral surface, grey on outer and dorsal surface, tarsomeres yellowish-white on inner and ventral surface, greyish-black on outer and dorsal surface, brightly white at end.

Male genitalia (Fig. 9). Uncus lobes fused into a shield, bearing long hairs, slightly concave at middle on posterior margin, asymmetrical. Subscaphium ribbon-like. Tegumen asymmetrical, left part broader than right part, forming a narrow ring with vinculum. Vinculum arched, narrowed, equipped with a globular lobe bearing long hairs at middle. Saccus not developed. A receptacle-shaped, sclerotised modification attached to vinculum anteriorly, possibly part of segments VII and VIII, with a sharp horn and a drumstick-like process at right on posterior margin. Sternum VIII articulated with vinculum at left, articulated with left valva dorso-basally; strongly modified, folded, concave at middle on costal margin, convex in hillock shape on ventral margin; broad in basal 3/4, with a digitate basal process, narrowed and thumb-like in distal 1/4, bearing long scales apically. Valvae strongly asymmetrical. Left valva larger than right one, irregular in shape, with a blade-shaped setose lobe apically; costal margin with a triangular protuberance at base and a 1/2-round protuberance near apex; ventral margin with a sword-hilt-like process subapically; inner surface with a finger-like lobe at base, a small, subquadrate, smooth process at middle near costal margin and a hillock-shaped smooth process near apex; apodeme distinct. Right valva divided into two parts: dorsal lobe strongly sclerotised, expanded and convex dorso-apically, with a large subquadrate and a small digitate process, with a vaulted, setose lobe ventro-apically; ventral lobe crescent, with a slender, hooked process distally, a membranous, hillock-shaped process that bears dense setae at middle of costal margin, a horned process at base of inner surface. Juxta broad, pocket-like, with a curved thorn-like process at end of right side. Aedeagus short, as long as modification of vinculum, slightly expanded and membranous basally; sclerotised near lateral sides, with a small process on right; tapered to pointed apex; cornutus absent.

Female. Unknown.

Distribution

China (Guangxi, Zhejiang).

Etymology

The specific name is derived from the Latin prefix hirt-, from hirtus meaning hairy and the Latin word globosus, globular, referring to the vinculum equipped with a globular lobe bearing long hairs.

DNA barcode

One DNA barcode from a paratype was generated and deposited in GenBank and BOLD systems: MZ711362/ DRYAD002-21. The minimum divergence (Table 1) to its nearest species, D. securiformis sp. nov., is 6.06%.

Dryadaula securiformis sp. nov.

http://zoobank.org/14EFF095-5E6A-4C29-8513-243C3B127E8F
Figures 4, 5–6, 10, 11

Type material

Holotype : China: • ♂; Hainan Province, Mt. Jianfeng (18°44'N, 108°52'E); alt. 787 m; 5.iii.2016; leg. Qingyun Wang. Paratypes: China: • 8♂, 1♀; same data as holotype, except dated 4–8.iii.2016; genitalia slide Nos DNAYLL18121m, DNAYLL18122m, DNAYLL18172, DNAYLL18173, XMR18158, XMR18334, XMR18335 • 1♂; Hainan Province, Mt. Jianfeng; alt. 770 m; 29.v.2015; leg. Peixin Cong; genitalia slide No. DNAYLL18171 • 2♀; Hainan Province, Mt. Jianfeng, Fengminggu; alt. 954 m; 8.viii.2017; leg. Xia Bai; genitalia slide Nos DNAYLL18123, XMR18241.

Differential diagnosis

The new species resembles D. trapezoides (Meyrick, 1935), but the flagellum has three cinereous bands towards apex, the forewing has an obscure blackish stripe at middle of fold and the ostium bursae located at middle on anterior 1/3 of sternum VIII in the female genitalia. In D. trapezoides, the flagellum has two dark fuscous bands towards apex, the forewing has pale yellowish spots suffused with a few dark scales above fold at 1/3 and 3/5 and the ostium bursae opens at left of sternum VIII in the female genitalia.

Description

Adult (Fig. 4): Wingspan 8.0 mm in holotype, 8.0–9.0 mm in paratypes. Vertex snow white to greyish-white, frons and occiput pale to smoky white. Antenna with scape snow white, without pecten; flagellum with alternate white and cinereous annulations, with three narrow cinereous bands towards apex. Labial palpus spatulate; yellowish-white on inner surface, black on outer surface, except yellowish-white at end of third palpomere. Thorax and tegula greyish-white, dusted with black. Forewing ground colour white, irrorate with blackish-brown and greyish scales, edged with bright ochreous yellow scales along termen and markings; patterned with black markings: costa with a wedge-shaped spot at base, a semicircular spot at 2/5, a larger semicircular patch from 3/5 to 4/5, diffused greyish smudges amongst spots; cell with a ribbon-like spot at distal 1/4, tending to coalesce with semicircular costal patch; fold with an obscure stripe at middle, obliquely inwards towards dorsum, surrounding suffusion of ochreous yellow; an interrupted terminal line around apex then along termen to tornus; cilia grey, with individual scales dark-tipped. Hind-wing and cilia grey. Legs yellowish-white, tibia black on outer surface, tarsus black on dorsal surface, except for end of each tarsomere.

Figure 11. 

Female genitalia of Dryadaula securiformis sp. nov., paratype, slide No. DNAYLL18123. Scale bars: 0.25 mm.

Male genitalia (Fig. 10). Uncus lobes fused into a terminally concave shield, bearing stout hair on caudal margin. Subscaphium an elongate band. Tegumen somewhat broad, with a hummocky process on left and caudal margins, respectively. Vinculum narrowed, sinuate, with a setose lobe at middle. Saccus not developed. A complicated, irregular, sclerotised modification attached to vinculum anteriorly, possibly part of segments VII and VIII; comprising of a Y-shaped sclerotisation and a receptacle-shaped sclerite that is enlarged and auricular at left. Sternum VIII articulated with vinculum at left, fused with left valva dorso-basally; oblong, folded, bottle-shaped, rounded and bearing long hairs apically, equipped with a spindle-shaped setose ridge beyond middle. Valvae strongly asymmetrical. Left valva battle axe-shaped, with a small finger-like lobe and a setose globular lobe at base; apodeme distinct, two. Right valva small, complicated: basal part broad, C-shaped; with a ribbon-shaped sclerite articulated with juxta; distal part bilobate into one large and one small lobe, both lobes slender, S-shaped, with an oval, setose apex. Juxta elliptical. Aedeagus a curved horn, tapered from base to a hook-like end; cornutus absent.

Female genitalia (Fig. 11). Oviscapt reduced; anal papillae a pair of short, mastoid lobes. Posterior apophyses short, equalling the length of sternum VIII. Anterior apophyses slightly longer and stouter than posterior apophyses. Tergum VIII short, rectangular, intricately wrinkled antero-laterally, bearing short setae on posterior margin. Sternum VIII somewhat rounded, intricately wrinkled, bearing dense short setae on posterior margin. Ostium bursae located at middle on anterior 1/3 of sternum VIII. Antrum funnel-shaped. Ductus bursae slender, short. Corpus bursae irregularly elongate, without signum.

Distribution

China (Hainan).

Etymology

The specific name is derived from the Latin word securiformis, referring to the battle-axe-shaped left valva.

DNA barcode

DNA sequencing resulted in a barcode of 604 bp from two paratypes: MZ711363/ DRYAD003-21 and MZ711364/ DRYAD004-21. The minimum distance (Table 1) to the nearest neighbour, D. hirtiglobosa sp. nov., is 6.06%.

Checklist

Until this study, a total of 49 species have been described worldwide as identified in the checklist in Table 2. It is a taxonomic summary of the published works, contains type localities, depositories of types, distribution and available sources of figures for adults or genitalia that were given by previous researchers.

Acknowledgements

We acknowledge Dr. Davis Lees (NHM, UK) for helping to examine types deposited in the NHM and Dr. S. Yu. Sinev (ZIN, Russia) for assisting in checking related specimens deposited in ZIN when the corresponding author visited the Institute and to those who took part in the fieldwork. We’re grateful to Dr. Toshiya Hirowatari (Kyushu University, Japan), Dr. Reinhard Gaedike (Bonn, Germany), Dr. Bong-Kyu Byun (Hannam University, Korea) and Dr. Andrew Mitchell for their encouragement and valuable suggestions on the manuscript. This research was supported by the National Natural Science Foundation of China (No. 31702034) and Basic Scientific Research Project of Henan Academy of Agricultural Sciences (No. 2021ZC45).

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