﻿The genus Dryadaula Meyrick (Lepidoptera, Tineoidea, Dryadaulidae) in China, with descriptions of four new species and a world checklist

﻿Abstract Four new species of the genus Dryadaula Meyrick, 1893 from China are described: Dryadaulaauriformissp. nov., D.flavostriatasp. nov., D.hirtiglobosasp. nov. and D.securiformissp. nov. Photographs of adults and genitalia of the new species are provided. DNA barcodes of D.auriformissp. nov., D.hirtiglobosasp. nov. and D.securiformissp. 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.


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, 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

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.

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 M 3 or CuA 1 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.
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 lajev, 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 dorsoapically. 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.

Distribution. China (Hainan).
Etymology. The specific name is derived from the Latin auriformis, meaning earshaped, 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) (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.
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.

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.  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-hiltlike 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) (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 yellowishwhite 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.
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.
Distribution. China (Hainan). Etymology. The specific name is derived from the Latin word securiformis, referring to the battle-axe-shaped left valva.

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.