The Symmocinae and Holcopogoninae in the Canary Islands and Madeira, with descriptions of 13 new species (Lepidoptera: Autostichidae)

We recognize 20 species of Symmocinae, concluding that all belong in the genus Chersogenes Walsingham, 1908 with the synonyms Epanastasis Walsingham, 1908, syn. rev., Ambloma Walsingham, 1908, syn. nov. and Thanatovena Gozmány, 1957. Thirteen species are described as new: Chersogenes variabilis Falck & Karsholt, sp. nov. (Spain: Gran Canaria), Chersogenes pseudocanariensis Falck & Karsholt, sp. nov. (Spain: Gran Canaria), Chersogenes subextricata Falck & Karsholt, sp. nov. (Spain: Tenerife), Chersogenes gomerae Falck & Karsholt, sp. nov. (Spain: La Gomera), Chersogenes nigra Falck & Karsholt, sp. nov. (Spain: Gran Canaria), Chersogenes hermiguae Falck & Karsholt, sp. nov. (Spain: La Gomera), Chersogenes mercedella Falck & Karsholt, sp. nov. (Spain: Tenerife), Chersogenes duabusalis Falck & Karsholt, sp. nov. (Spain: Fuerteventura and Lanzarote), Chersogenes aguiari Falck & Karsholt, sp. nov. (Portugal: Selvagens Islands), Chersogenes coxi Falck & Karsholt, sp. nov. (Spain: Fuerteventura), Chersogenes lanzarotae Falck & Karsholt, sp. nov. (Spain: Lanzarote), Chersogenes fuerteventurae Falck & Karsholt, sp. nov. (Spain: Fuerteventura), Chersogenes indistincta Falck & Karsholt, sp. nov. (Spain: Fuerteventura and Lanzarote). Chersogenes extricata (Gozmány, 1964), sp. rev., comb. nov. is removed from synonymy of Chersogenes ( Epanastasis ) canariensis (Rebel, 1906) and reinstated as a distinct species. Chersogenes excellens (Gozmány, 1977), syn. nov., comb. nov. is synonymized with Chersogenes klimeschi (Gozmány, 1975), comb. nov. Symmoca canariensis Rebel, 1906 and Epanastasis eupracta Gozmány, 1988 are newly combined as Chersogenes canariensis (Rebel, 1906), comb. nov. and Chersogenes eupracta (Gozmány, 1988), comb. nov. The following North African species are formally transferred from Epanastasis : Chersogenes arenbergerorum (Gozmány, 1988), comb. nov., Chersogenes enigmatica (Gozmány, 1964), comb. n., Chersogenes eremicola (Gozmány, 1988), comb. nov., Chersogenes erroris (Gozmány, 1962), comb. nov., Chersogenes friedeli (Gozmány, 1988), comb. nov., Chersogenes tunesica (Gozmány, 1988), comb. nov., and Chersogenes vetustella (Zerny, 1935), comb. nov. Two species of Holcopogoninae, Turatia iranica Gozmány, 2000 and Hesperesta hartigi (Turati, 1934) are recorded as new to the Canary Islands. Two of the new species, C. duabusalis , sp. nov. and C. aguiari , sp. nov. have brachypterous males. Photographs of the adults of all species are shown. Photographs of the genitalia of the new species are provided. All of the new species are barcoded. Analyses of DNA barcodes show that the identifications and distinctiveness of each species as well-supported and genetically isolated.


Introduction
The Autostichidae, as currently understood, is a medium-sized family of gelechioid moths with about 800 described species. It includes six or seven subfamilies (Heikkilä et al. 2014, p. 585;Wang & Li, 2020, pp. 323-324), three of which (Oegoconiinae, Symmocinae and Holcopogoninae) occur in the West Palaearctic region, including the Canary Islands and Madeira. In a previous paper (Falck et al. 2021) we dealt with the Oecogoniinae, and here we treat the Symmocinae and Holcopogoninae of these islands.
The Symmocinae are distributed mainly in dryer areas of the western Palaearctic. In his revision Gozmány (2008) recognized 230 species, and relatively few additional species are known from the Afrotropical and Oriental regions. Holcopogoninae are a smaller subfamily with less than 40 species distributed in Eurasia and Africa.
Although the Autostichidae, and especially the two first mentioned subfamilies, are diverse in the Canary Islands they have received surprisingly little attention in the lepidopterological literature. Rebel (1896Rebel ( , 1906 described two species of Symmocinae, and Walsingham (1908) two further species, and then very little happened until Gozmány (1964Gozmány ( , 1975Gozmány ( , 1988) described four additional symmocine species. Klimesch (1985) reviewed the then known species, and they were again revised by Gozmány (2008) in the "Microlepidoptera Palaearctica" series.
In the present paper we revise the taxonomy at both genus and species level of the Symmocinae of the Canary Islands and Madeira, reducing the number of genera from three to one, and raising the number of species from seven (Vives Moreno, 2014, pp. 109-110) to 20, by describing 13 new species.
The Holcopogoninae of the Canary Islands have received even less attention. Only one species was previously recorded from the islands. We add here the records of two additional species.
Female: Wingspan 14-16.5 mm. Differs from male by the slightly shorter antenna and the broader forewing.
Variation: C. variabilis exhibits considerable variation. In both sexes the ground colour varies from almost pure off-white to dark grey and the wing pattern may be absent.
DNA barcodes ( Figure 87): We obtained full length DNA barcode (658 bp) from eighteen specimens and DNA barcode fragments of 581 bp, 605 bp and 612 bp from three specimens. The barcodes fall within Barcode Index Numbers (BIN) BOLD: ADT9918 (nineteen specimens) and ADT9915 (two specimens). The maximum intraspecific p-distance is high 2.36%. The minimum pdistance to the nearest neighbour C. canariensis is 5.35%.
Diagnosis: C. variabilis resembles other members of the genus, especially C. extricata. It is distinguished by the robust appearance and the rounded apex of the forewing. In the male genitalia the short straight sacculus and two cornuti groups are characteristic. In the female genitalia the rounded posterior margin of the lamella antevaginalis, the short colliculum and the narrow ductus bursae are characteristic.
Biology: Early stages unknown. Most specimens were collected at light, but some were disturbed from varied vegetation during the daytime, from the beginning of August until the end of September, at altitudes from 80 to 1400 m.
Distribution: Widely distributed on the island of Gran Canaria, Spain, except coastal areas. Etymology: The species is named after the variable adults. Remarks: Interestingly it seems that the different populations are highly local. Specimens near the city of Moya all belong to the grey form with a distinct wing pattern, while further south near the city of Fontanales the specimens are with a white or light grey ground colour, and with indistinct or totally without a wing pattern, at the southernmost location, Pie de la Cuesta the specimens are dark grey with an indistinct wing pattern. The molecular analyses show high intraspecific divergence, but without any correlation between the different populations of C. variabilis.
Description: Male. Wingspan 10.5-13 mm. Labial palp upturned, segment 2 creamy white dorsally and medially, ventrally with a dark brown scale tuft, extending beyond the base of segment 3, segment 3 half the length of segment 2, creamy white, laterally mottled with black scales. Antenna black with indistinct grey rings. Head and neck yellowish brown; thorax and tegula yellowish brown, mottled with brown scales. Forewing yellowish brown, with a distinct broad dark brown costal line reaching apex; dorsal 1/3 light brown mottled with a few black scales; at 1/3 one or two diffuse brownish black spots, at 2/3 two distinct black discal spots; termen with distinct black spots; fringe grey. Hindwing grey with grey fringe. Abdomen yellowish brown. Female. Wingspan 12 mm. Differs from the male by the uniformly yellowish-brown colour of the head, neck, thorax, tegula and forewing.
Variation: The spots may be more or less distinct, sometimes forming two indistinct oblique fasciae.
Female genitalia ( Figure 77): Papilla analis long, distally pointed, posterior apophysis slender, twice as long as papilla analis; anterior apophysis slightly longer than half the length of posterior apophysis; tergum VIII sub-rectangular, sternum VIII with median fissure widening anteriorly; lamella antevaginalis sub-rectangular, broad, posterior margin laterally slightly concave with a broad medial Vshaped invagination; antrum cup shaped; colliculum quite narrow and sclerotized; ductus bursae membranous, about twice as broad as colliculum and slightly narrowing anteriorly; corpus bursae membranous oval, signum with about 15 small spines.
DNA barcodes ( Figure 87): We obtained full length DNA barcodes (658 bp) from three specimens. The barcodes fall within Barcode Index Number (BIN) BOLD: ADF2619. The maximum intraspecific p-distance is 0.31%. The barcodes of C. sophroniellus, C. pseudocanariensis and C. canariensis cluster together in the neighbour-joining tree, and the minimum p-distance between them is 3.04% (C. sophroniellus vs. C. pseudocanariensis), 4.13% (C. sophroniellus vs. C. canariensis) and 4.89% (C. pseudocanariensis vs. C. canariensis). The nearest neighbour to them is C. variabilis with a 5.35% divergence.
Diagnosis: The yellowish-brown colour, the distinct dark brown costal line (in male) and the long segment 2 of the labial palp separates C. sophroniellus from similar species such as C. canariensis and C. pseudocanariensis. In the male genitalia the sickle-shaped apex of the sacculus and only one cornuti group with very long cornuti are characteristic. It differs from C. canariensis by the sickle-shaped apex of the sacculus, and by having only one group of very long cornuti. It differs from C. pseudocanariensis in the slightly broader valva and in having fewer cornuti. In the female genitalia the shape of the posterior margin lamella antevaginalis is characteristic. It differs from from C. canariensis in the concave postero-lateral margin of ostium and the less pronounced invagination. It closely resembles C. pseudocanariensis and there is no reliable difference in the female genitalia between the two species. Adults of C. sophroniellus are very characteristic and are easily separated from other members of the genus.
Biology: Early stages unknown. Most of the examined specimens were caught during evening sunshine, and a few were collected at light during late March and April at altitudes ranging from about 500-1000 m.
Distribution: Known only from the holotype from the island of Tenerife and a few specimens from the surroundings of Teror and one specimen from Valleseco in the northern part of Gran Canaria.
Remarks: Epanastasis sophroniellus was described by Rebel (1894) on the basis of a single male collected by John Henry Leech in April 1885 on the island of Tenerife, deposited in the Walsingham collection (NHMUK). In a later publication Rebel (1896, p. 128, pl. III) provides a more detailed description based on a series of specimens from nearby Teror at the northern part of Gran Canaria collected the 10th of May 1895 by the Danish lepidopterologist Wilhelm von Hedemann.
The spelling "sophroniella" is an unnecessary gender-agreement variant of the original sophroniellus in the literature.
DNA barcodes ( Figure 87): We obtained full length DNA barcodes (658 bp) from six specimens. The barcodes fall within Barcode Index Number (BIN) BOLD: ADF2781. The maximum intraspecific p-distance is 1.38%. The results are presented in the DNA barcodes section of C. sophroniellus.
Diagnosis: C. pseudocanariensis is similar to C. canariensis, q.v. Biology: Early stages unknown. Most of the specimens were attracted to light, but most of the males were found nectaring on flowers of Todaroa montana Webb ex Christ on a rainy and humid day. The adults of C. pseudocanariensis were collected from January to late June at altitudes from 450 m to 1400 m.
Distribution: Known only from mountain regions in the southern half of the island of Gran Canaria, Spain.
Etymology: The species name is derived from combining the Greek word øåõäï (pseudo = false) and canariensis, referring to its similarity to C. canariensis.
Female genitalia (Figures 79): Papilla analis long, distally pointed, posterior apophysis slender, twice as long as papilla analis; anterior apophysis slightly longer than half the length of posterior apophysis; tergum VIII sub-rectangular, sternum VIII with median fissure widening anteriorly; lamella antevaginalis sub-rectangular, broad, posterior margin laterally straight with a broad medial V-shaped invagination; antrum cup shaped; colliculum quite narrow and sclerotised; ductus bursae membranous, about 3 times broader than colliculum and narrowing anteriorly; corpus bursae membranous oval, signum with about 15 small spines.
DNA barcodes ( Figure 87): We obtained full length DNA barcode (658 bp) from eleven specimens and DNA barcode fragments of 519 bp, 605 bp, 629 bp, 630 bp, 634 bp, 647 bp and 648 bp from seven specimens. The barcodes fall within Barcode Index Numbers (BIN) BOLD: ADT8333 (thirteen specimens from northern Gran Canaria, El Hierro and Tenerife), ADI4064 (four specimens from southern Gran Canaria) and AEN3712 (one specimen from central Gran Canaria), the minimum p-distance between the BINs is 1.92% (ADI4064 vs. AEN3712), 2.08% (ADI4064 vs. ADT8333) and 2.22% (ADT8333 vs. AEN3712). The maximum intraspecific p-distance is very high 3.10%. The results are presented in the DNA barcodes section of C. sophroniellus.
Diagnosis: C. canariensis resembles C. sophroniellus (q.v.) and especially C. pseudocanariensis, and it is not possible to distinguish specimens from the southern part of Gran Canaria without dissection of the genitalia. In the male genitalia the short hook-shaped apex of the sacculus and two cornuti groups are characteristic. In the female genitalia the laterally straight posterior margin of lamella antevaginalis and the bulbous ductus bursae are characteristic.
Biology: Early stages unknown. Most specimens were collected at light, but some were disturbed from varied vegetation during daytime, from March to September, at altitudes from sea level to 1800 m.
Remarks: In his paper "Fünfter Beitrag zur Lepidopterenfauna der Kanaren" Rebel (1906) described Symmoca canariensis from one male specimen collected at Güímar, Tenerife by W. White. It was examined by Walsingham (1908, pp. 949-950) and compared with the type specimen of C. sophroniellus. He concluded that adults of these two highly variable species only were separable by the difference in labial palps. In later literature e. g. Klimesch (1985, p. 136) and Gozmány (2008, p. 190) C. canariensis was misinterpreted as C. sophroniellus.
It appears that the males of C. canariensis occur in two geographical forms in Gran Canaria, in the northern part they are similar to specimens from El Hierro and Tenerife, in the southern part they are similar to males of C. pseudocanariensis. The molecular analyses show high intraspecific divergence in COI, and there seems to be some correlation between the different populations from Gran Canaria in the DNA and in the adult appearance. Although the divergence between two of the populations is above the 2% threshold suggested as a putative guideline for species delimitation by Hebert et al. (2003), we hesitate to describe them as new species, because we were unable to find any constant difference in the genitalia. Description: Male. Wingspan 14-15.5 mm. Labial palp upturned, segment 2 white mottled with black and brown scales, especially ventrally and laterally, ventrally a small brownish scale tuft, segment 3 almost as long as segment 2, whitish mottled with dark grey scales. Antenna almost as long as forewing, black. Head and neck grey mottled with white; thorax and tegula greyish brown with a few white scales. Forewing greyish white, mottled with ochreous, brown and black scales, fringe grey. Hindwing grey with grey fringe. Abdomen greyish.

Chersogenes subextricata
Female: Unknown. Variation: One of the specimens has the forewing grey with a broad dark grey costal line reaching the apex; median 1/3 light grey mottled with ochreous scales; dorsal 1/3 grey, darker brownish grey towards termen; one black discal spot. C. subextricata is probably a variable species.
Diagnosis: In the male genitalia the sickle-shaped sacculus, the short appendix and only one group of cornuti are characteristic. For separation from similar species see under C. extricata.
Biology: Early stages unknown. The specimens were attracted to light from the beginning of November until the beginning of December, at altitudes between 100 m and 200 m.
Distribution: Only known from the island of Tenerife, Spain. Etymology: The species is named C. subextricata because of its similarity to C. extricata.
Description: Male. Wingspan 14-16.5 mm. Labial palp upturned, segment 2 white, laterally and ventrally mottled with dark brown and black scales, segment 3 almost as long as segment 2, whitish mottled with brown and black. Antenna almost as long as forewing, black with indistinct grey rings. Head and neck greyish white mottled with brown; thorax and tegula ochreous mottled with brown especially towards neck. Forewing grey mottled with brown and black; from the base to near apex a whitish mottled with ochreous median streak, at 1/3 and before 2/3 bordered by two black dots; along termen and apex 6-8 black dots; fringe grey. Hindwing grey with grey fringe. Abdomen greyish brown. THE SYMMOCINAE AND HOLCOPOGONINAE IN THE CANARY ISLANDS AND MADEIRA, WITH DESCRIPTIONS OF 13 NEW SPECIES Female: Wingspan 13 mm. Differs from the male in being brachypterous, the forewing being shorter and broader, the hindwing slenderer with a pointed apex. Forewing dark brownish.
Variation: C. extricata exhibits considerable variation. The ground colour varies from whitish grey or dark grey to yellowish brown or ochreous, the median streak is often indistinct or missing, the distal black spots are sometimes confluent, or the black spots may be totally absent.
DNA barcodes ( Figure 87): We obtained full length DNA barcodes (658 bp) from four specimens and DNA barcode fragments of 633 bp from one specimen. The barcodes fall within Barcode Index Number (BIN) BOLD: ADF2779. The maximum intraspecific p-distance is 1.61%. The results are presented in the DNA barcodes section of C. subextricata.
Diagnosis: The three species C. extricata, C. subextricata and C. gomerae are closely related, and it is not possible to separate them without dissection of the genitalia or barcoding. The group is distinguished from the similar looking species C. canariensis and C. pseudocanariensis by the lack of a scale tuft ventrally on segment 2 of the labial palp. In the male genitalia of C.extricata the sickleshaped sacculus, the short appendix and the two group of cornuti are characteristic. It is distinguished from C. subextricata in having two groups of cornuti, and from C. gomerae in the more curved apex of the sacculus and a larger number of cornuti in both groups of cornuti.
Biology: Early stages unknown. The specimens were attracted to light during October and November, at altitudes from 430 m to 1400 m. The only known female was found running on the ground near the light.
Distribution: Known only from the mountain area in the central part of the island of Gran Canaria, Spain.
Description: Male. Wingspan 14-17 mm. Labial palp upturned, segment 2 white, laterally and ventrally mottled with dark brown and black scales, segment 3 almost as long as segment 2, whitish mottled with brown and black. Antenna almost as long as forewing, black with indistinct grey rings. Head and neck greyish white mottled with brown; thorax and tegula ochreous mottled with brown especially towards neck. Forewing grey mottled with brown and black; from the base to near apex a whitish mottled with ochreous median streak, at 1/3 and before 2/3 bordered by two black dots; along termen and apex 6-8 black dots; fringe grey. Hindwing grey with grey fringe. Abdomen greyish brown.
Variation: C. gomerae exhibits considerable variation. The ground colour varies from whitish grey or dark grey to yellowish brown or ochreous, the median streak is often indistinct or missing, the distal black spots are sometimes confluent, or the black spots may be totally absent.
DNA barcodes ( Figure 87): We obtained full length DNA barcodes (658 bp) from three specimens. The barcodes fall within Barcode Index Number (BIN) BOLD: AEP5785. The maximum intraspecific p-distance is 0.16%. The results are presented in the DNA barcodes section of C. subextricata.
Diagnosis: In the male genitalia the sickle-shaped sacculus, the short appendix and two groups of cornuti are characteristic. For separation from similar species see under C. extricata.
Biology: Early stages unknown. All specimens of the type series were attracted to light from late October until the beginning of November at altitudes from 570 m to 820 m.
Distribution: Only known from a few scattered localities in the mountain region of the island of La Gomera, Spain.

Chersogenes nigra
Description: Male. Wingspan 8.5 mm. Labial palp upturned, segment 2 blackish, posteriorly white and mottled white ventrally, segment 3 almost as long as segment 2, blackish. Antenna as long as the length of forewing, black with indistinct grey rings. Head and neck dark brown mottled light grey; thorax and tegula blackish brown. Forewing dark brown mottled with black scales; at 1/5 and 1/2 two broad, diffuse, lighter brown fasciae; fringe grey. Hindwing dark brown with grey fringe. Abdomen dark greyish brown.
Female: Wingspan 8.5 mm. Differs from male by the broader forewing and the more clearly marked fasciae.
Diagnosis: C. nigra does not resemble other members of the genus, but the female is superficially similar to some species of Scythris Hübner, 1825, especially in the S. petrella species-group (Bengtsson, 1997, p. 138). The small size and dark colour are characteristic. In the male genitalia the short straight sacculus, the upturned apex of the valva and the lack of cornuti are characteristic. In the female genitalia the short papilla analis, the narrow segment VIII, the narrow concave lamella antevaginalis and the lack of signum are characteristic.
Biology: Early stages unknown. Both specimens were attracted to light. Distribution: Only known from the type locality, Pie de la Cuesta, Gran Canaria, Spain. Etymology: The species is named after its dark colour.
Female: Wingspan 12-15 mm. Differs from the male by a more prominent wing pattern. Variation: C. eupracta is a highly variable species. The colour of the forewing varies from yellowish to brownish and grey, and the mottling may be absent or very prominent. The wing pattern is sometimes almost absent, giving a uniform appearance of the wing, and in other specimens the wing pattern is very pronounced and forming two distinct, dark brown fasciae, especially in the females.
Diagnosis: The yellowish-brown colour of the forewing and the very contrasting dark grey hindwing separates C. eupracta from most species in the genus. It closely resembles C. mercedella and can be distinguished by the two ochreous spots before and beyond the middle. In the male genitalia the slender and bent apex of the sacculus, and the long, narrow transtilla projection are characteristic. It differs from C. mercedella by the evenly bent apex of the sacculus and the longer transtilla projection. In the female genitalia the twisted membraneous part of ductus bursae and the large ductus seminalis are characteristic.
Biology: Early stages unknown. The specimens were collected from the beginning of June until the end of October mainly at light, but C. eupracta was also observed flying actively in the afternoon sunshine in great numbers, both in a mixed forest and in open mountain areas at altitudes from 110 m to 1400 m.
Distribution: Widely distributed on the island of Gran Canaria and from one locality in the western part of the island of Tenerife, Spain. Description: Male. Wingspan 9 mm. Labial palp upturned, segment 2 grey mottled with white scales, segment 3 grey, half the length of segment 2. Antenna black with indistinct grey rings. Vertex whitish; head with laterally, grey and white scale tufts; neck grey mottled with white scales, thorax and tegula greyish mottled with brown scales. Forewing grey, apically mottled with black scales; basally with blackish spot at costa; at 1/3 an indistinct, oblique, black fascia, in the middle part mottled with ochreous scales; at 2/3 an indistinct, oblique black fascia; in the discal area an ochre-coloured spot. fringe grey. Hindwing grey with grey fringe. Abdomen greyish brown.
DNA barcodes (Figure 87): We obtained full length DNA barcodes (658 bp) from two specimens. The barcodes fall within Barcode Index Number (BIN) BOLD: AEM6648. The intraspecific p-distance is 0.00%. The results are presented in the DNA barcodes section of C. eupracta.
Diagnosis: The small size and grey colour with two dark oblique fasciae of the forewing separates C. hermiguae from other species in the genus. In the male genitalia the long appendix, the long narrow projection of the transtilla and the 90º degree angle of the sacculus are very characteristic.
Biology: Early stages unknown. The specimens were attracted to light. Distribution: Known only from the type locality in the northern part of the island of La Gomera, Spain.
Etymology: The species is named after its type locality, Hermigua. Description: Male. Wingspan 7-10.5 mm. Labial palp upturned, segment 2 dark brown, ventrally yellowish, segment 3 longer than the half the length of segment 2, yellowish, mottled with dark brown scales. Antenna black with indistinct grey rings. Head, neck, tegula and thorax yellowish brown. Forewing yellowish brown, mottled with light brown and dark brown scales; mottled with black scales at dorsum in distal half, in discal area and apically; fringe grey. Hindwing dark grey with grey fringe. Abdomen yellowish brown.

Chersogenes mercedella
Female: Unknown. Variation: The mottling with black scales of the forewing is sometimes almost absent, otherwise forming two indistinct, oblique, black fasciae.
DNA barcodes (Figure 87): We obtained a full-length DNA barcode (658 bp) from one specimen and DNA barcode fragments of 635 and 627pb from two specimens. The barcodes fall within Barcode Index Number (BIN) BOLD: AEC2599. The maximum intraspecific p-distance is 0.96%. The results are presented in the DNA barcodes section of C. eupracta.
Diagnosis: The yellowish-brown colour of the forewing and the very contrasting dark grey hindwing separates C. mercedella from most of the species in the genus. It closely resembles C. eupracta, q. v.
Biology: Early stages unknown. The specimens were attracted to light. Distribution: Known only from the type locality in the eastern part of the island of Tenerife, Spain. Etymology: The species is named after its type locality, Las Mercedes. Female: Wingspan 7-8 mm. Differs from the male by the shorter antenna, the broader forewing and the distinctly concave apical half of costa. Variation: Ground colour varies from beige, grey and to dark brown, often present is an indistinct, median, beige streak from base to about 2/3.
DNA barcodes ( Figure 87): We obtained full length DNA barcodes (658 bp) from three specimens. The barcodes fall within Barcode Index Number (BIN) BOLD: ADV1498. The maximum intraspecific p-distance is 0.64%. The results are presented in the DNA barcodes section of C. klimeschi.
Diagnosis: The rudimentary hindwing separates C. duabusalis from most species in the genus. It resembles C. aguiari and is distinguished by the broader wings and larger size. In the male genitalia the small transtilla projection, the very short appendix and the bent phallus with few robust cornuti are characteristic. They differ from C. aguiari by the longer appendix, the shorter gnathos, the posteriorly broader phallus and by having two cornuti groups. In the female genitalia the crown-shaped lamella antevaginalis and the irregular signum plate are characteristic.
Biology: Early stages unknown. Most of the specimens were netted in open grass areas during daytime in full sunshine, and a few were attracted to light, all from February, March and November at altitudes ranging from sea level to 400 m.
Distribution: Known only from the islands of Fuerteventura and Lanzarote, Spain. Etymology: The species is named after the Latin words duabus alis (= two wings) referring to the adult appearance, as appearing to have only two wings. Paratypes: 2 11, same data as holotype, genitalia slide 3801APF (AFA, ZMUC). Description: Male. Wingspan 5 mm. Labial palp light grey-brown, slightly upcurved, segment 2 broadest in apical part; segment 3 almost three times shorter than segment 2, with pointed apex. Antenna slightly longer than forewing, ringed dark brown and whitish. Head and thorax [partly denuded] whitish grey-brown. Forewing very slender, pointed, whitish grey with some grey-brown tipped scales. Hindwing rudimentary. Abdomen rather stout, yellowish white.
DNA barcodes: Barcoding failed twice. Diagnosis: C. aguiari resembles C. duabusalis (q. v.). In the male genitalia the short appendix and the bent, rather slender phallus with one cornuti group are characteristic.
Biology: Early stages unknown. The three type specimens were sifted from soil near Schizogyne sericea (L.f.) DC. (Asteraceae) growing in shade.
Distribution: Only known from the type locality Selvagem Grande in the Selvagens Islands, Portugal.
Etymology: The species is dedicated to António M. Franquinho Aguiar, who placed the known specimens of this new species at our disposal and contributed so much to the knowledge of the entomofauna of the Madeira Islands.
Remarks: The Selvagens Islands, situated between the Canary Islands and Madeira, have a very low diversity of lepidopteran fauna. Aguiar & Karsholt (2006, p. 13) listed only 24 lepidopteran species from these islands. None of them are endemic to the islands but are either widespread or also occur on the Canary Islands. The geological history of the Selvagens Islands is most closely related with that of the Canary Islands (Geldmacher et al., 2001). C. aguiari is thus the first endemic Lepidoptera species for the Selvagens Islands, which has 59 endemic arthropod species (UNESCO, 2017).
The islands are of volcanic origin and were never connected to the continent. They are very old, about 29 million years, and very eroded and have probably not been submerged during the last 4-5 million years (Geldmacher et al. 2001), which is crucial for the terrestrial fauna. The largest of the islands, and type locality of C. aguiari, Selvagem Grande, has an area of only 245 ha and raises to a rather flat plateau with altitudes up to 163 m. It is dominated by low vegetation and strong winds, giving only limited shelter for winged insects like Lepidoptera.
Variation: It appears that there is some general geographic variation in the ground colour. It varies from whitish (Gran Canaria), greyish with beige (La Gomera and Tenerife) and to dark grey (El Hierro), the spots are sometimes very diffuse and often totally absent.
DNA barcodes (Figure 87): We obtained full length DNA barcodes (658 bp) from twelve specimens and DNA barcode fragments of 591 bp and 623 bp from two specimens. The barcodes fall within Barcode Index Numbers (BIN) BOLD: AEH2995 (one specimen from Gran Canaria), AEH2996 (one specimen from Gran Canaria), AEH2997 (three specimens from Gran Canaria), ADF2780 (one specimen from Gran Canaria), ADU2892 (two specimens from Tenerife), ADY9824 (one specimen from Tenerife and three specimens from La Gomera) and AEW3196 ( two specimens from El Hierro). The maximum intraspecific p-distance is very high 5.20%. The barcodes of C. klimeschi, C. duabusalis, C. coxi and C. lanzarotae cluster together in the neighbour-joining tree and the minimum pdistances between them are 5.29% (C. klimeschi vs. C. duabusalis), 5.81% (C. klimeschi vs. C. coxi), 5.35% (C. klimeschi vs. C. lanzarotae), 6.17% (C. duabusalis vs. C. lanzarotae), 5.81% (C. duabusalis vs. C. coxi) and 3.93% (C. coxi vs. C. lanzarotae). The nearest neighbour to them is C. variabilis, with a 6.57% divergence. Diagnosis: C. klimeschi resembles other members of the genus, especially pale specimens of C. coxi. It is distinguished by the smaller scale tuft ventrally on segment 2 of the labial palp. In the male genitalia the long gnathos, the short transtilla projection and three cornuti groups are characteristic.
Biology: Early stages unknown. The specimens were netted from low vegetation during the daytime in full sunshine or attracted to light from almost all months of the year at altitudes ranging from sea level to 895 m.
Distribution: Only known from the Canary Islands: Gran Canaria, La Gomera, El Hierro and Tenerife, Spain.
Remarks: Despite the high intraspecific divergence in COI and variation in adult appearance we were not able to find any morphological differences in the genitalia between the separate populations. We consider C. excellens as a synonym based on similar morphology both in the adult appearance and the genitalia. Gozmány (2008, p. 413) figures the male genitalia without the transtilla projection, whereas Klimesch (1985, p. 146) illustrated them (from the same genitalia slide) with the transtilla projection on the right side. If the transtilla projections are small they are easily hidden by the costal margin of the valva. During dissection of the genitalia it is important to place both the valva and the phallus in the correct position to recognise the diagnostic details such as the small transtilla projection, juxta and the number of cornuti groups. The best way to recognize the number of cornuti groups is to leave the phallus in situ.
Description: Male. Brachypterous. Wingspan 8-9 mm. Labial palp upturned, segment 2 white with small dark grey scale tuft ventrally, segment 3 slightly shorter than segment 2, whitish dark grey especially ventrally. Antenna as long as forewing, dark grey. Head greyish. Neck whitish. Thorax grey mottled with white. Tegula dark grey basally, greyish white distally. Forewing very slender and pointed; ground colour grey, heavily mottled with white and with scattered black scales; at 1/3 a diffuse ochreous spot; fringe grey. Hindwing very slender, short and pointed, grey with grey fringe. Abdomen greyish.
DNA barcodes: Due to the lack of fresh material we were not able to barcode any specimens. Diagnosis: C. brachyptera resembles C. coxi and especially C. klimeschi in the colour and the wing pattern. It is distinguished by being brachypterous. In the male genitalia the long gnathos, the short transtilla projection, the relatively long and slender phallus with one cornuti group are characteristic. It differs from C. klimeschi in the shorter gnathos, the longer and slenderer phallus and in having only one cornuti group with longer spines.
Distribution: Known only from the southern coastal areas of the island of Tenerife, Spain. Description: Male. Wingspan 9.5-11 mm. Labial palp upturned, segment 2 white, ventrally and laterally dark grey and with dark grey scale tuft ventrally, segment 3 shorter than segment 2, whitish mottled with dark grey ventrally and laterally. Antenna almost as long as forewing, grey with indistinct whitish rings. Head and neck whitish. Thorax and tegula whitish with a few scattered black scales. Forewing slender and pointed; ground colour whitish, grey at costa and dorsum, in the middle part at 1/3 and 2/3 and in the apical area mottled with relatively few black scales; at 1/3 and 2/3 diffuse ochreous spots; fringe grey. Hindwing grey with grey fringe. Abdomen light grey.

Chersogenes coxi
Female: Wingspan 10 mm. Differs from the male by being brachypterous, the forewing being shorter and broader, the distinctly concave apical half of costa and the hindwing slenderer with pointed apex. Forewing ochreous; at 1/3 a broad, diffuse brownish fascia, apical half mottled with a few light to dark brown scales.
Variation: C. coxi exhibits considerable variation. The ground colour varies from white or grey to yellowish brown or ochreous. The scattered black scales may be almost absent.
DNA barcodes (Figure 87): We obtained full length DNA barcodes (658 bp) from six specimens and DNA barcode fragments of 632 bp and 619 bp from two specimens. The barcodes fall within Barcode Index Numbers (BIN) BOLD: ADU1582 and ADT9919. The maximum intraspecific pdistance is very high, 3.86%. The results are presented in the DNA barcodes section of C. klimeschi.
Diagnosis: C. coxi resembles other members of the genus, especially C. klimeschi q.v. and C. lanzarotae. It is distinguished from the latter by the usual greater number of black scales. In the male genitalia the short transtilla projection, the short and broad phallus with the drop-like apex and one cornuti group are characteristic.
Biology: Early stages unknown. Most of the specimens were attracted to light and a few were disturbed from varied vegetation during the daytime during the months of October, November and February at altitudes ranging from sea level to 400 m.
Distribution: Known only from the islands of Fuerteventura, Spain. Etymology: The species is dedicated to the late Dutch lepidopterist Anton Cox, who collected the first known specimens used for our study.
Remarks: Specimens from dune areas have a white or light grey ground colour while specimens from the rocky and mountain areas have a yellowish to ochreous ground colour. The molecular analyses show high intraspecific divergence and splitting into two well separated clusters, but without any clear correlation to the two populations of C. coxi, since the "dune population" is represented in both clusters. We did not find any differences in the genitalia between the populations. Description: Male. Wingspan 9-11 mm. Labial palp upturned, segment 2 white ventrally and laterally dark grey, ventrally with dark grey scale tuft, segment 3 shorter than segment 2, whitish mottled with dark grey ventrally and laterally. Antenna almost as long as forewing, grey with indistinct whitish rings. Head and neck brownish grey. Thorax brownish, ochreous towards abdomen. Tegula brownish, distally beige. Forewing slender and pointed; ground colour brownish grey, at the base and in middle half to near apex ochreous; a few scattered black scales in apical half; at 2/3 sometimes a small black dot near dorsum; fringe grey. Hindwing grey with grey fringe. Abdomen light brownish.

Chersogenes lanzarotae
Female: Unknown. Variation: The ground colour varies from grey to brown, the ochreous part is sometimes reduced to a narrow diffuse streak and the black scales are sometimes absent.
DNA barcodes ( Figure 87): We obtained full length DNA barcodes (658 bp) from three specimens and DNA barcode fragments of 652 bp from one specimen. The barcodes fall within Barcode Index Numbers (BIN) BOLD: AEC6345, AEC2598 and AEC5794. The maximum intraspecific p-distance is 2.24%. The results are presented in the DNA barcodes section of C. klimeschi.
Diagnosis: C. lanzarotae is characterized by the slender and pointed forewing, the scale tuft ventrally on segment 2 of the labial palps and the very few scattered black scales. It resembles C. coxi q. v. In the male genitalia the strongly curved and one small straight cornuti are very characteristic.
Biology: Early stages unknown. The specimens were all attracted to light in the months of October, November and February at altitudes ranging from sea level to 280 m.
Distribution: Known only from the island of Lanzarote, Spain. Etymology: The species is named after the island of Lanzarote. Remarks: C. lanzarotae is, like several other members of the genus, highly variable and with high intraspecific divergence in the DNA barcodes. Walsingham, 1908 (Figures 48, 73, 73a) Chersogenes victimella Walsingham, 1908. Proc. zool. Soc. Lond., 1907 Type locality: SPAIN, Tenerife, Santa Cruz.
Description: Male. Wingspan 12 mm. Labial palp straight and long, segment 2 with dark grey scale tuft ventrally, the whole segment covered by rough, long dark grey scales, white dorsally, segment 3 dark brown. Antenna almost as long as forewing, dark grey with indistinct whitish rings. Head, neck, thorax and tegula greyish brown mottled with white. Forewing slender and pointed, costa slightly concave in apical half; ground colour dark grey, from the base to about 2/3 a narrow yellowish white streak; at 1/3 a spot of black raised scales on each side of the streak, at 2/3 a smaller spot of black raised scales on each side of the streak; fringe grey. Hindwing dark brown with dark grey fringe. Abdomen brownish.
DNA barcodes ( Figure 87): We obtained full length DNA barcodes (658 bp) from two specimens. The barcodes fall within Barcode Index Number (BIN) BOLD: AEV1724. The intraspecific p-distance is 0.00%. The minimum p-distance to nearest neighbour (C. fuerteventurae) is 8.1%.
Diagnosis: C. victimella is very characteristic because of the long and roughly scaled labial palp and the black dots with raised scales. It resembles no other species in the genus. In the male genitalia the relatively stout transtilla projection, the sickle-shaped apex of the sacculus and the straight phallus with two cornuti groups are characteristic.
Biology: Early stages unknown. The examined specimens were disturbed from a stone-wall during the daytime.
Distribution: Only known from two locations in the south-eastern part of Tenerife, Spain. Description: Male. Wingspan 7-8 mm. Labial palp upturned, segment 2 light brown, ventrally with a small brown scale tuft, segment 3 dark brown and slightly shorter than segment 2. Antenna almost as long as forewing, brown. Head, neck, thorax and tegula greyish brown. Forewing dark beige mottled with dark brown especially along termen and apex, costa blackish; at 1/3 and 2/3 two small indistinct dark brown spots, costal spots located more basally; fringe grey. Hindwing grey with grey fringe. Abdomen dark beige.

Chersogenes fuerteventurae
Female: Wingspan 10 mm. The forewing more heavily mottled with brown. Variation: Only minor variation in the distinctiveness of the spots. Male genitalia (Figures 74, 74a): Uncus long, slender rectangular, apex spatulate; gnathos shorter than uncus, straight, apically hook-shaped, apex pointed; tegumen sub-triangular, anterior margin flatly U-shaped; valva simple, approximately 6 times longer than broad, upturned in distal third, apex rounded; sacculus about 2/3 length of valva, apically evenly bent upwards; appendix distinctly shorter than valva, slightly bent, apex pointed; juxta sub-rectangular, slightly narrower in the middle, in anterior half two small lateral lamella; saccus triangular; phallus slightly bent, broad; two cornuti THE SYMMOCINAE AND HOLCOPOGONINAE IN THE CANARY ISLANDS AND MADEIRA, WITH DESCRIPTIONS OF 13 NEW SPECIES groups almost confluent, anterior group with 14-16 spines of various length, posterior group with 3-4 short spines.
DNA barcodes (Figure 87): We obtained full length DNA barcodes (658 bp) from three specimens. The barcodes fall within Barcode Index Number (BIN) BOLD: ADT9917. The intraspecific p-distance is 0.00%. The minimum p-distance to nearest neighbour (C. victimella) is 8.01%.
Diagnosis: C. fuerteventurae is characterized by the size and the dark beige colouration and the distinct wing pattern. In the male genitalia the shape of the juxta and the broad phallus with two almost confluent cornuti groups are characteristic. In the female genitalia the shape of the antrum and colliculum and the lack of a signum are characteristic.
Biology: Early stages unknown. The males were netted or disturbed from varied vegetation during daytime in full sunshine, the females were found nectaring on flowers of Asteriscus sericeus DC. (Asteraceae) from late February and March.
Distribution: Known only from the island of Fuerteventura, Spain. Etymology: The species is named after the island of Fuerteventura where the type series was collected. Description: Male. Wingspan 12-16.5 mm. Labial palp upturned, segment 2 white, mottled with dark beige, ventrally a large dark beige scale tuft, segment 3 as long as segment 2, whitish mottled with beige and grey. Antenna as long as forewing, light brownish with indistinct grey rings. Head and neck beige mottled with dark grey. Thorax and tegula dark beige. Forewing beige mottled with dark grey and black, especially along costa and apex, cell more whitish; at 1/3 and 2/3 two small indistinct ochreous spots, bordered by scattered black scales. Hindwing grey with grey fringe. Abdomen beige.

Chersogenes indistincta
Female. Similar to male, but with shorter antenna. Variation: The ground colour varies from beige to grey heavily mottled with black, wing pattern more or less distinct or totally absent.
Diagnosis: C. indistincta resembles no other known species of the genus. It is characterized by the size, the beige colouration with indistinct wingpattern and the large scale-tuft on the labial palp. In the male genitalia the long saccus, the shape of juxta and the few cornuti in the anterior group are characteristic. In the female genitalia the narrow lamella antevaginalis and the heavily sclerotised signum with robust spines are characteristic.
Biology: Early stages unknown. The specimens were attracted to light in the months February, March, October and November at altitudes ranging from sea level to 610 m. One specimen was reared by chance from Lotus lancerottensis Webb & Berthel. (Fabaceae).
Remarks: The occurrence of T. iranica in the Canary Islands far away from its hitherto known distribution around the Persian Gulf is surprising. The identity of the disjunct populations was confirmed by their similar genitalia. However, further studies may reveal cryptic diversity, because specimens from Canary Islands and the Arab United Emirates differ by 3.85% in the DNA barcodes. The species probably also occurs through northern parts of the Sahara. The male and female genitalia are figured by Gozmány (2000, pp. 133, 151).
Distribution: Widely distributed in tropical and subtropical parts the world. However, records from some areas are based on misidentified specimens of similar looking species of Tineidae or Oecophoridae (Gozmány, 2000, p. 88).
Remarks: This genus and species were placed by Hodges (1998, p. 141) in a monotypic subfamily Oeciinae of his newly erected family Schistonoeidae. However, this action has not received general acceptance, and Gozmány (2000, p. 85) transferred it to the Holcopogonidae. The latter is currently considered as a subfamily, Holcopogoninae of the Autostichidae (Heikkilä et al. 2014, p. 585, L. Kaila in litt.).
Oecia oecophila was first recorded from Madeira by Carvalho (1995, p. 579) without locality and date. We have only seen the specimens listed above from that island.

Discussion
The molecular analyses support the taxonomic arrangement. All identified species are clearly genetically distinct from other species with uncorrected p-distance values between species ranging from 3.93% (between C. coxi and C. lanzarotae) to 12.08% (between C. coxi and C. fuerteventurae). The taxonomy of the Canarian symmocinae species is not entirely unproblematic, as several species exhibit high variability in the adult appearance and very high intraspecific values in COI, both between populations from separate islands of the Canary Islands (e. g. C. canariensis and C. klimeschi) but also between populations from the same island (e. g. C. klimeschi, C. variabilis and C. lanzarotae). A high intraspecific variation in COI between species from separate islands of the Canary Islands is commonly observed (Falck et al. 2021, p. 298;Falck et al. 2022, p. 108). It can be interpreted as a snapshot of the evolutionary process; however further studies may reveal cryptic species.
The Autostichidae occurring in the Canary Islands are largely restricted to two genera, Apatema Walsingham, 1900 with 18 species (see Falck et al. 2021) and Chersogenes with 19 species. In addition to these, only three species of Holcopogonidae are recorded from the islands. Moreover, we describe one species of Chersogenes from the Selvagens Islands, which belong to Portugal.
Based on our study of the morphology and DNA barcodes we conclude that all Symmocinae species found in the Canary Islands belong to a single genus, Chersogenes. Outside of the Canary and Selvagens Islands this genus is only known from North Africa, where seven species have been recorded under the genus synonym of Epanastasis. Our data do not allow us to conclude if Chersogenes species from North Africa or from the Canary Islands phylogenetically are most basal. It is, however, evident that the fauna of Chersogenes from the Canary and Selvagens Islands are much more diverse, compared with the species occurring in North Africa. They have probably been present for a very long time, having had time to evolve external differences in, e. g. labial palps, form and venation of the wings and brachyptery. This is also reflected in the DNA barcode, with large percentage distances between many of the species.
Hypotheses relating to brachyptery and its function in species of Lepidoptera are not uniform and each case should be looked at on its own merits (Sattler, 1991, p. 244). With the biology and larval host-plants being largely unknown it is not clear which factors benefit development of wing reduction in Chersogenes. Sattler, 1991, p. 248 comments that a number of Lepidoptera species with THE SYMMOCINAE AND HOLCOPOGONINAE IN THE CANARY ISLANDS AND MADEIRA, WITH DESCRIPTIONS OF 13 NEW SPECIES brachypterous females or with wing reduction in both sexes inhabit grassland, which constitutes a permanent, continuous habitat.