The genus Apatema Walsingham, 1900 in the Canary Islands and Madeira, with description of 13 new species

The genus Apatema Walsingham, 1900 in the Canary Islands and Madeira is revised based on external morphology, genitalia and DNA barcodes. We recognize 18 species, 13 of which are described as new: Apatema confluellum Falck & Karsholt, sp. n. (Spain: Tenerife), Apatema minimum Falck & Karsholt, sp. n. (Spain: Lanzarote and Fuerteventura), Apatema lapalmae Falck & Karsholt, sp. n. (Spain: La Palma), Apatema sallyae Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema lanzarotae Falck & Karsholt, sp. n. (Spain: Lanzarote), Apatema mixtum Falck & Karsholt, sp. n. (Spain: Tenerife), Apatema pseudolucidum Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema griseum Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema grancanariae Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema brunneum Falck & Karsholt, sp. n. (Spain: Tenerife), Apatema transversum Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema stadeli Falck & Karsholt, sp. n. (Spain: Tenerife) and Apatema skulei Falck & Karsholt, sp. n. (Spain: Gran Canaria). The monotypic Dysallomima Gozmány, 2008, syn. n. is treated as a new synonym of Apatema and its type species Lampros coarctella Rebel, 1896 is transferred as Apatema coarctella (Rebel, 1896) comb. n. Apatema helleri (Rebel, 1910), comb. n. is transferred from the Blastobasidae. A. mediopallidum Walsingham, 1900 is removed from the list of Lepidoptera of the Canary Island. Morphology of all species is described and figured in detail. All species occurs in the Canary Islands and only one, A. fasciata (Stainton, 1859) occurs both in these islands and in the Madeiran archipelago. Records of A. mediopallidum Walsingham, 1900 from the Canary Islands are based on misidentifications. Analyses of DNA barcodes support the identification distinctiveness of the species as each species appear monophyletic, well-supported and genetically isolated in the analyses. The only exception is A. coarctella , which could not be rendered monophyletic with respect to a single A. mediopallidum specimen, when sequences from non-Canarian Apatema were included in the analysis. We argue that this result is an artefact.


Introduction
Oegoconiinae are a small group of Gelechioidea, most of which are distributed in the Western Palearctic region.They were included by GOZMÁNY (2008) in his revision of the Palearctic Symmocidae.According to recent molecular studies, e. g.HEIKKILÄ et al. (2013) Oegoconiinae should be regarded as a subfamily of Autostichidae.WALSINGHAM (1900: 220) erected the genus Apatema for a new species, A. mediopallidum, which he described from France: Corse.In a treatment of Microlepidoptera of the Canary Islands he subsequently transferred Gelechia fasciata Stainton, 1859 and Lampros coarctella Rebel, 1896 (as a synonym of fasciata) to Apatema and described a new species, A. lucidum (Walsinghm, 1908: 945).During most of the 20 th century most specimens from outside of the Macaronesia Islands were referred to A. mediopallidum.Apatema species are rather small Gelechioidea having blackish forewings with whitish or yellowish markings, their genitalia are rather simple and similar between the species, and it has only slowly become evident that the genus is more diverse that hitherto suspected.
The Lepidoptera of the Canary Islands are considered well-known.Similar to other oceanic islands the fauna of the Canary Islands is less diverse than that of the adjacent continental countries (Morocco, Portugal), but includes a number of endemic species.Recent field work by the first author (e. g.FALCK et al., 2019;FALCK & KARSHOLT, 2019) has revealed a number of hitherto unrecorded or even undescribed species of Lepidoptera occurring in these islands.In the present paper we revise the genus Apatema in the Canary Island and the Madeiran Archipelago.
Many Canarian Apatema species are quite easily identified based on adult habitus (e. g. A. sallyae Falck & Karsholt, sp. n. and A. lucidum Walsingham), while others are difficult to identify without genitalia dissections (e. g. A. mixtum Falck & Karsholt, sp. n. and A. fasciata (Stainton)).When dissecting the male genitalia it is very important to separate the cornuti groups (Fig. 28) to ensure correct identification.In the female genitalia (Fig. 48) some species (e. g. A. stadeli Falck & Karsholt, sp. n.) have a slightly twisted ductus bursae.This gives the impression of a longitudinal fold of the anterior part of ductus bursae in the genital-slide, despite being an artefact, which we will refer to as "a longitudinal fold".
Here we revise the Canarian Apatema species based on adult morphology and DNA barcodes (a 658 bp fragment of the mitochondrial COI gene).We provide detailed descriptions and diagnoses for all species and discuss their molecular diversity and potential phylogenetic relationships.

Material and methods
Most of the specimens were attracted to an 8 watt super actinic light.Some of the specimens especially from forest localities were obtained by netting in the afternoon sunshine and a few specimens were disturbed from the vegetation during daytime.Genitalia were dissected following Robinson (1976).Whole specimens were photographed with a Canon EOS 700D camera equipped with a Canon EF 100 mm objective.The genitalia slides were photographed using a Soptop CX40T Trinocular microscope in conjunction with a Toup Tek P10500A-E3 / E3ISPM05000KPA-E3 / 5.0MP USB3 camera.
The so-called DNA barcode fragment (a 658 bp fragment of the mitochondrial COI gene) was sequenced from all known Apatema species from the Canary Islands to detect genetically distinct clusters, and to obtain molecular data for new species.DNA samples (dried legs) were prepared according to the accepted standards and were processed at the Canadian Centre for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph).Genetic clusters are presented with their barcode index number (BIN; cf.RATMNASINGHAM & HERBERT, 2013).
To assess overall DNA diversity and relationships within the Canary Island Apatema we analysed the DNA barcodes under a Bayesian framework in MrBayes 3.2.(RONQUIST et al., 2011).The analysis was done utilising the model-jumping option (nst=mixed) allowing MrBayes to assess the best evolutionary model for the dataset.The rates were set to GAMMA, all other priors were left as default.The analysis was run for 10 million generations with sampling every 1000 generation, and the first 25% were used as burnin.The run files were subsequently assessed in Tracer 1.7.1 (part of the BEAST package, BOUCKAERT et al., 2019) to confirm that the two parallel analyses had converged and reached stationarity.The majority rule consensus tree was visualised in FigTree 1. 4.4. (RAMBAUT, 2014).We analysed the dataset in BEAST in conjunction with BEAUti (BOUCKAERT et al., 2019) under the GTR model for molecular evolution (the model found by MrBayes) with rates set to GAMMA and all other priors left as default.The analysis was run for 10 million generations with sampling every 5000 generations.The resultant trees were then imported into DensiTree (BOUCKAERT & HELED, 2014), with the first 25% used as burnin, to visualise potential conflict areas in the data.To test whether the Apatema species found on the Canary Island and Madeira potentially comprise a monophyletic group, we augmented our DNA barcode dataset with publicly available barcode sequences from four additional species of Apatema (16 sequences) and three species of the closely related genus Oegoconia (six sequences).The new dataset was analysed under a Bayesian framework in MrBayes 3.2.utilising the model-jumping option with rates set to GAMMA.The analysis was run for 20 million generations with sampling every 1000 generations, and the first 25% were used as burnin.The run files were subsequently assessed in Tracer 1.7.1, and the majority rule consensus tree was visualised in FigTree 1.4.4.In all Bayesian analyses a single specimen of each of the three species Epanastatis sophroniella (Rebel, 1894), E. canariensis (Rebel, 1906) and Ambloma klimeschi Gozmány, 1975 were used as outgroups.All specimens used in the molecular analyses are listed in Table 1.We explored the general genetic diversity in Apatema from the Canary Island by calculating uncorrected p distances in Mega X (KUMAR et al., 2018).We calculated the average uncorrected p distance within as well as between each putative species (as listed in Table 1 and the Taxonomic section); the values are listed in Table 2.We further divided A. coarctella into populations from Fuerteventura, Gran Canaria, Lanzarote, La Palma and Tenerife and separately calculated the average uncorrected p distance within and between the four populations; the values are listed in Table 3.Similarly, we divided A. fasciata into populations from Gran Canaria, Fuerteventura and Lanzarote and separately calculated the average uncorrected p distance within and between the three populations; the values are listed in Table 4.

Abbreviations used
GP Genitalia preparation JJ Collection of Jari Junnilainen, Vantaa, Finland PF Collection of Per Falck, Neksø, Denmark MNCN Collection of Antonio Vives, Museo Nacional de Ciencias Naturales, Madrid, Spain NHMUK The Natural History Museum, London, UK TL Type locality ZMUC Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark
The monotypic genus Dysallomima was erected by GOZMÁNY (2008: 62) for Lampros coarctella, which differs from other Apatema species by details of the genitalia in both sexes.In our analyses of the DNA barcodes coarctella is nested deeply within the genus Apatema, and we consider it as a specialized representative of this genus.
Biology: Almost nothing is know about the biology and the early stages are unknown.The first author reared one species from a piece of dead wood overgrown with fungi.
Distribution: Western Palaearctic region.Our work demonstrates that the genus Apatema is much more diverse in the Canary Islands that hitherto believed.Field work by the first author has resulted in freshly collected material of both sexes from most species.This has enabled us to identify 18 separate species, which differ in both morphology and DNA.These are described in detail below.Description: Adult.Wingspan 7-11 mm.Labial palp slender, upturned, segment 2 pale-grey, laterally dark-grey at base and towards apex, medially near apex with a dark-grey spot, segment 3 black, dorsally pale-grey.Antenna black, with indistinct rings in females.Head and neck yellowish brown, thorax grey mottled with dark brown especially anteriorly; tegula pale brown.Forewing pale greyish brown mottled with brown, especially along costa and towards apex, base black; at 1/3 and 1/2 with two larges, diffuse black spots; apical spot yellowish grey, indistinct; fringe grey.Hindwing palegrey, with pale-grey fringe.
DNA barcodes (Figs 64-66, Tables 1-2): Three specimens were barcoded with some internal variation in COI (uncorrected p = 0.0061).The specimens are placed as sister to A. minimum.A. confluellum is clearly differentiated from all other species with uncorrected p distance to other species ranging from 0.0935 and 0.0986 (A.pseudolucidum and A. minimum) to 0.1207 (A.fasciata).Barcode Index Number: ADS6252.
Diagnosis: The greyish brown colour and the two large spots in the forewing separate A. confluellum from all other known Apatema species.In the male genitalia the nearly straight valva, the relatively long sacculus and the few cornuti in cornuti group I and II are characteristic.In the female genitalia the parallel-sided anterior part of ductus bursae and the small longitudinal fold are characteristic.
Biology: Early stages unknown.The specimens were attracted to light during March, May-June and November-December at altitudes ranging from sea level to 500 m.
Distribution: Only known from the island of Tenerife, Spain.Etymology: The name (a noun in opposition) is derived from combining the Latin word confluus (= confluent) with the diminutive suffixellum.Description: Adult.Wingspan 7-7.5 mm.Labial palp slender, upturned, segment 2 white, basally black, segment 3 white with few blackish scales ventrally.Antenna black.Head, neck and thorax creamy white; tegula creamy white, basally black.Forewing ground colour black, centrally a regular, slightly oblique creamy white fascia almost reaching dorsum; costal spot creamy white, merging with tornal spot, forming a distinct outer fascia; fringe grey.Hindwing grey with grey fringe.
DNA barcodes (Figs 64-66, Tables 1-2): Five specimens were barcoded with very high internal variation in COI (uncorrected p = 0.0317).The specimens are sister to A. confluellum.A minimum is clearly differentiated from all other species with uncorrected p distance to other species ranging from 0.0986 (A.confluellum) to 0.1352 (A.grancanariae).In fact, A. minimum appears to be the genetically most isolated species displaying more than 10% COI divergence (uncorrected p = 0.1) to all species except A. confluellum.Barcode Index Numbers: ADS4524, AEC2840.
Diagnosis: A. minimum superficially resembles other black and white Apatema species, e. g. A. sallyae and A. mediopallidum.It can be distinguished from these by the very small size, and the two regular and distinct creamy white forewing fasciae; in A. sallyae the inner fascia is oval, clearly not reaching dorsum and the outer fascia is indistinct; in A. mediopallidum the inner fascia is irregular with black dots and outer fascia is absent.In the male genitalia the short labis, the shape of the cornuti in group I and the few short cornuti in group III are characteristic.In the female genitalia the anteriorly membranous part of ductus bursae is characteristic.
Biology: Unknown.The specimens were collected at light during late October and November at altitudes ranging from sea level to 600 m.
Distribution: Known only from the islands of Fuerteventura and Lanzarote, Spain.Etymology: The species is named after its small size, from the Latin adjective minimus meaning smallest.
Remarks: KLIMESCH (1985: 137) mentions two specimens of A. mediopallidum from Lanzarote, Haria, 2 11, 3-5-III-1967, leg.F. Kasy, with a wingspan 7 mm.These two specimens most likely belong to A. minimum sp.n.Forewing ground colour black with white markings; base white, at costa with a tiny white spot before an irregular oblique white fascia at 1/3, narrow at costa, widening towards dorsum; costal spot white, reaching half way towards tiny white tornal spot; diffuse white markings at dorsum between fascia and tornal spot, and one diffuse mark apically; fringe grey.Hindwing pale-grey, with grey fringe.
DNA barcodes (Figs 64-66, Tables 1-2): Two specimens were barcoded with some internal variation in COI (uncorrected p = 0.0049).The two specimens are placed sister to A. helleri.A. lapalmae is clearly differentiated from all other species with an uncorrected p distance to other species ranging from 0.0563 (A.helleri) to 0.1264 (A.minimum).Barcode Index Number: ADY6666.
Diagnosis: A. lapalmae resembles A. helleri.It can be distinguished by the darker head and neck; the white fascia is narrow at costa, widening towards dorsum (broader at costa in A. helleri).In the male genitalia the relatively narrow plate with spines in cornuti group I and the on average difference in length of needle-shaped cornuti between cornuti group II and III -longest in group III are characteristic.In the female genitalia the longitudinal fold of the anterior half of ductus bursae is characteristic for A. lapalmae and A. helleri, and it is not possible to separate the two species based on the female genitalia.
Biology: The specimens were reared from a piece of dead wood overgrown with fungus, it was found in a dark part of a Laurisilva forest.
Distribution: Only known from the island of La Palma, Spain.Etymology: The species is named after its place of occurrence, the island of La Palma.The name is an adjective.
Description Adult: Wingspan 10-12.5 mm.Labial palp slender, upturned, segment 2 white, laterally black at base, segment 3 with black basal half and white apical half.Antenna black, with indistinct dark-grey rings in female; male antenna uniformly dark-grey.Head, neck and thorax white, tegula white, basally black.Forewing black with white markings; base white, at costa with a tiny white spot and an irregular oblique white fascia at 1/3 of wing; costal spot white, extending half way towards a tiny white tornal spot; diffuse white markings at dorsum between fascia and tornal spot and one diffuse mark in apical area; fringe grey.Hindwing pale-grey, with pale-grey fringe.
DNA barcodes (Figs 64-66, Tables 1-2): Two specimens were barcoded with some internal variation in COI (uncorrected p = 0.0031).The two specimens are placed on a long branch as sister to A. lapalmae.A. helleri is clearly differentiated from all other species with uncorrected p distance to other species ranging from 0.0563 (A.lapalmae) to 0.1316 (A.minimum).Barcode Index Number: ADS6251.
Diagnosis: A. helleri resembles A. lapalmae.It can be distinguished by the pure white head and thorax; the white fascia is as wide at costa as in the middle of the wing (more narrow at costa in A. lapalmae).In the male genitalia cornuti group I and II with needle-shaped spines of the same length and approximately the same number of spines is characteristic.In the female genitalia the longitudinal fold of the anterior half of ductus bursae is characteristic for A. helleri and A. lapalmae, and it is not possible to separate the two species based on the female genitalia.
Biology: Early stages unknown.The specimens were attracted to light during March and June-November at altitudes ranging from sea level to 500 m.
Distribution: Only known from the northern part of the island of Gran Canaria, Spain.Remarks: REBEL (1910) described A. helleri from a single specimen collected by K. M. Heller in Teror on the northern part of Gran Canaria on the 25-V-1907.We have not been able to trace the type specimen and it is apparently lost.The species was placed in Blastobasis, but Rebel already then had his doubt "Mit Rücksicht darauf, daß mir nur ein einziges, einer fremden Sammlung angehöriges

Apatema sallyae
Description Adult: Wingspan 10-14 mm.Labial palp slender, upturned, segment 2 creamy white, laterally dark-grey at base, segment 3 creamy white, dorsally greyish.Antenna dark-grey, with indistinct pale-grey rings in female.Head and neck creamy white, thorax creamy white, dark-grey towards neck; tegula creamy white, dark-grey towards base.Forewing ground colour uniform blackish grey with distinct creamy white markings; base at dorsum creamy white, at costa with a tiny spot followed by an almost oval irregular fascia at 1/3 not reaching dorsum; apical costal spot creamy white, narrowing, almost reaching dorsum; fringe grey.Hindwing pale-grey, with pale-grey fringe.
DNA barcodes.(Figs 64-66, Tables 1-2): Two specimens were barcoded with no internal variation in COI.The two specimens appear rather isolated on a long branch without clear affiliation to any other species.A. sallyae is clearly differentiated from all other species with uncorrected p distance to other species ranging from ca 0.074 (A.helleri, A. lapalmae and A. skulei) to 0.1208 (A.minimum).Barcode Index Number: ADS4523.
Diagnosis: A. sallyae resembles no other Apatema species known from the Canary Islands.However, it resembles A. mediopallidum Walsingham, 1900, with which it has been confused in the past.It can be distinguished by the creamy white head, neck and thorax and the large oval creamy white spot in the middle of the forewing.In the male genitalia the broad-based triangular spines in group I, and the short needle-shaped cornuti in group II are characteristic.In the female genitalia the anteriorly widening and evenly sclerotized ductus bursae are characteristic.
Biology: Early stages unknown.All the specimens were attracted to light during March-June at altitudes ranging from sea level to 1100 m.
Distribution: Only known from the southern part of the island of Gran Canaria.Etymology: The species name (a noun in the genitive case) is dedicated to the first author`s oldest daughter Sally.
Remarks: A. sallyae is figured by KLIMESCH (1985: 147, figs. 15, 17) under the name Apatema mediopallidum Walsingham, 1901, with  Description Adult: Wingspan 10 mm.Labial palp slender, upturned, segment 2 white, laterally dark-grey at base, segment 3 white, with few dark-grey scales ventrally.Antenna dark-grey, with indistinct pale-grey rings.Head, neck and thorax creamy white, mottled with brown; tegula creamy white, brownish towards base.Forewing ground colour white, mottled with dark-grey and brown, especially in basal and apical areas; centrally with a very indistinct, oblique light grey fascia, bordered by four indistinct black spots; fringe grey.Hindwing light grey with grey fringe.
Female genitalia: Unknown.DNA barcodes (Figs 64-66, Tables 1-2): Two specimens were barcoded with some variation in COI (uncorrected p = 0.0030).The two specimens appear isolated on a long branch without clear affiliation to other species.A. lanzarotae is clearly differentiated from all other species with uncorrected p distance to other species ranging from 0.0801 (A.lapalmae) to 0.1316 (A.minimum).Barcode Index Number: AEC2600.
Diagnosis: A. lanzarotae resembles no other Apatema species known from the Canary Islands, although it may be difficult to distinguish from old specimens of A. fasciata.It can be distinguished by the indistinct black spots, in A. fasciata the spots are distinct.In the male genitalia the needle-shaped spines distally in cornuti group I are characteristic.
Biology: Unknown.The specimens were collected in late autumn at light.Distribution: Known only from a few localities in the northern part of the island Lanzarote, Spain.Etymology: The species is named after its place of occurrence, the island of Lanzarote.The name is an adjective.Description Adult: Wingspan 9.5-13 mm.Labial palp slender, upturned, segment 2 creamy white, laterally dark-grey at base and towards apex, medially with a dark-grey spot near apex, segment 3 black, dorsally and apically creamy white.Antenna black, with indistinct dark-grey rings in female, uniformly dark-grey in male.Head, neck and thorax creamy white mottled with dark brown; tegula creamy white, dark brown towards base.Forewing blackish brown mottled with light brown, cream and white; base black; centrally with an irregular oblique whitish brown fascia bordered by four black spots; apical spot at costa white, extending half way towards a tiny white tornal spot; fringe grey.Hindwing grey with grey fringe.

Apatema mixtum
Variation: The species varies in size and appearance of the four black spots which are often well separated but may be confluent in pairs.The costal and tornal spots may be connected by a thin, angulated fascia.
DNA barcodes (Figs 64-66, Tables 1-2): Six specimens were barcoded with very low variation in COI (uncorrected p = 0.0010).The specimens are placed on a long branch and are sister to A. fasciata.A. mixtum is differentiated from all other species with uncorrected p distance to other species ranging from 0.0676 (A.lapalmae) to 0.1247 (A.minimum).Barcode Index Number: ADR5557.
Diagnosis: Apatema mixtum closely resembles A. fasciata and adults cannot be distinguished with certainty.In the male genitalia the few, short, sometimes diminutive spines in cornuti group II are characteristic, in A. fasciata cornuti group II is missing.In the female genitalia the double rotated ductus bursae is characteristic.
Biology: Early stages unknown.The specimens were attracted to light during January-March, August and November-December at altitudes ranging from sea level to 1050 m.
Distribution: Known from several localities on the island of Tenerife, Spain.
Etymology: The name refers to the Latin adjective mixtus meaning mixed, in this case mixed with A. fasciata.
Remarks: Despite the missing details in the genitalia drawings by KLIMESCH (1985: 148, figs. 19, 20, 21)  Description Adult: Wingspan 10-11 mm.Labial palp slender, upturned, segment 2 on outer surface black with a creamy coloured band, on inner surface cream coloured with black apex, segment 3 black with cream coloured tip.Antenna black, with indistinct dark-grey rings.Head dark-grey, paler towards face and neck; thorax and tegula black mottled with yellowish grey.Forewing black (especially at base and in apical area) mottled with light brown, cream and white; with an irregular whitish brown fascia in middle of wing bordered by four black spots; costal spot white, extending half way towards tiny white tornal spot; fringe grey.Hindwing grey, with grey fringe.
Variation: The species varies in size and appearance of the four black spots which are often well separated but may be confluent in pairs.The costal and tornal spots may be connected by a thin, angulated fascia.
DNA barcodes (Figs 64-66, Tables 1-2): Eight specimens were barcoded with reasonably high variation in COI (uncorrected p = 0.0165).The specimens are sister to A. mixtum.Despite high internal variation A. fasciata is differentiated from all other species with uncorrected p distance to other species ranging from ca 0.08 (A.stadeli and A. brunneum) to 0.1272 (A.minimum).Specimens from Gran Canaria (6 specimens) were clearly separate from specimens from Fuerteventura (2) and Lanzarote (1) with uncorrected p distances between them of 0.0269 and 0.0335 respectively.Similarly, the uncorrected p distance between the single specimens from Fuerteventura and Lanzarote was 0.0282.Barcode Index Numbers: ADV0186 (Gran Canaria), ADZ8617 (Fuerteventura), AEC2601 (Lanzarote).
Diagnosis: A. fasciata closely resembles A. mixtum, A. grancanariae and A. transversum, for separation see below under A. grancanariae.In the male genitalia the triangular cornuti in cornuti group I and the missing cornuti in group III are characteristic.In the female genitalia the weakly sclerotized and rotated ductus bursae is characteristic.
Biology: Early stages unknown.The specimens were attracted to light during January-December at altitudes ranging from sea level to 895 m.
Distribution: Known from Portugal: Madeira, Deserta Grande and Porto Santo; Spain: Fuerteventura, Gran Canaria and Lanzarote.
Remarks: Gelechia fasciata was described from an unstated number of specimens collected by T. V. Wollaston on the island of Deserta Grande in the Madeira archipelago.Records of A. fasciata from outside of Madeira and the Canary Islands (e. g.GOZMÁNY, 1955: 316)  Description Adult: Wingspan 7.5-10.5 mm.Labial palp slender, upturned, segment 2 creamy white, laterally black at base and towards apex, medially with a dark-grey spot near apex, segment 3 black, dorsally and apically creamy white.Antenna black, with indistinct dark-grey rings in female, uniformly dark-grey in male.Head and neck light brown mottled with brown; thorax dark brown; tegula creamy white, dark brown towards base.Forewing ground colour black mottled with brown; base black with a small pale reddish-brown spot towards base; with a small pale reddish-brown spot at costa near base; centrally with an irregular, oblique reddish-brown fascia bordered by four black spots; apical spot at costa creamy white, extending half way towards the tiny reddish-brown tornal spot, occasionally forming an indistinct outer fascia; fringe grey.Hindwing grey with grey fringe.
DNA barcodes (Figs 64-66, Tables 1-3): Fifteen specimens were barcoded with very high variation in COI (uncorrected p = 0.0334).The specimens are sister to a clade comprising A. junnilaineni, A. griseum, A. stadeli, A. transversum, A. skulei, A. brunneum, A. lucidum and A. grancanariae.Despite high internal variation A. coarctella is differentiated from all other species with uncorrected p distance to other species ranging from 0.0597 (A.lapalmae) to 0.1255 (A.minimum).A. coarctella is divided into three well-separated sub-groups comprised by specimens from Tenerife (6 specimens), La Palma (2), Lanzarote (2), Gran Canaria and Fuerteventura (5), respectively.While within group variation is very low, the four sub-groups are very divergent with the uncorrected p distance between Gran Canaria + Fuerteventura, and La Palma being 0.0534, the distance between Gran Canaria + Fuerteventura, and Tenerife being 0.0463, the distance between Tenerife and La Palma being 0.0295, the distance between Tenerife and Lanzarote being 0.0447, and the distance between La Palma and Lanzarote being 0.054.The distance between Gran Canaria + Fuerteventura, and Lanzarote is low 0.0022, and the two groups do not appear to be isolated.Barcode Index Numbers: ADR9887 (Tenerife), AEA1295 (La Palma), ADY6487 (Fuerteventura, Gran Canaria and Lanzarote).Diagnosis: A. coarctella resembles small specimens of A. fasciata and A. mixtum.It can be distinguished by an on average smaller size and the reddish-brown markings on the forewing.In the male genitalia the forked distal part of sacculus is characteristic, it separates it from all other known Apatema species.In the female genitalia the dilated, membraneous ductus with lateral sclerotisations is characteristic.
Biology: Unknown.Most of the specimens are collected at light during January-April, June, July and October-December at altitudes ranging from sea level to 1200 m.Distribution: A. coarctella often occurs in large numbers and it is known from Fuerteventura, La Gomera (KLIMESCH, 1985: 138), Gran Canaria, Lanzarote and Tenerife.
Description Adult: Wingspan 13-16 mm.Labial palp slender, upturned, segment 2 yellowish white, laterally dark brown at base and apically, segment 3 yellowish white.Antenna yellowish brown, in female with indistinct brown rings.Head, thorax and tegula pale yellowish brown, neck pale brown.Forewing ground colour pale yellowish brown mottled with brown especially distally at costa and in apically; base with a black dot at costa and dorsum; four black spots, three in a row along cell at 1/3, 1/2 and distally, the latter often comma-shaped and one spot near dorsum at 2/5; costal spot yellowish and indistinct; fringe grey.Hindwing light grey with grey fringe.
Variation: The two outer pairs of spots are sometimes confluent.Females are often heavily mottled with dark brown giving a dark brown appearance, but the black spots are still visible.
DNA barcodes (Figs 64-66, Tables 1-2): Four specimens were barcoded with some internal variation in COI (uncorrected p = 0.0031).The specimens are sister to A. grancanariae.A. lucidum is clearly differentiated from all other species with uncorrected p distance to other species ranging from 0.0504 (A.grancanariae) to 0.1309 (A.minimum).Barcode Index Number: ADY7243.
Diagnosis: A. lucidum resembles A. pseudolucidum.It can be distinguished by the pale yellowish brown colour of the forewing and by having only four black and less distinct spots (six small black, very distinct spots in A. pseudolucidum).In the male genitalia the lack of cornuti group II and rather few long needle-shaped cornuti in cornuti group III are characteristic.In the female genitalia the dilated membranous ductus bursae and the rotation angel are characteristic.
Biology: Early stages unknown.The adults were flying actively in the evening sunshine in mixed forest and attracted to light during April-June at altitudes ranging from 400 m to 1700 m.
Distribution: Only known from the island of Tenerife, Spain.
Remarks: The adults figured by GOZMÁNY (2008: 483, plate 115, figs. 10, 10a) do not correspond to A. lucidum, fig. 10 is most likely A. fasciata and fig.10a A. mixta.The male and female genitalia are figured by GOZMÁNY (2008: 370, plate 4, fig.10;438, plate 72, fig. 10)  Description Adult: Wingspan 13.5-15 mm.Labial palp slender, upturned, segment 2 white, laterally black at base and apically, segment 3 black with white tip.Antenna black, in female with indistinct grey rings.Head and neck pale brown mottled with dark brown; thorax dark brown, paler brown towards abdomen; tegula brown, darker brown basally.Forewing ground colour dark-grey mottled with black; base black with a creamy white spot medially; centrally with an irregular oblique creamy white fascia mottled with grey and brown, the fascia is bordered by four black spots; apical spot at costa creamy white, extending half way towards tiny white tornal spot, sometimes forming an indistinct outer fascia; fringe grey.Hindwing grey with grey fringe.
DNA barcodes (Figs 64-66, Tables 1-2): Six specimens were barcoded with very high variation in COI (uncorrected p = 0.0233).The specimens are sister to A. lucidum.Despite high internal variation A. grancanariae is differentiated from all other species with uncorrected p distance to other species ranging from 0.0504 (A.lucidum) to 0.1352 (A.minimum).Barcode Index Numbers: AEA2797, AEA0771 and ADY6665.
Diagnosis: A. grancanariae resembles other Apatema species with similar wing pattern, especially A. fasciata, A. mixtum and A. transversum.It can be distinguished from A. fasciata and A. mixtum by an on average larger size, but it is not always possible to separate adults confidently.It can be distinguished from A. transversum by the paler and more oblique fascia in the middle of the forewing.In the male genitalia the short sclerite, the relatively long spines in cornuti group I, the missing cornuti group III and the lack of microspines in the vesica are characteristic (in A. fasciata the cornuti in group I are triangular, in A. mixtum cornuti group II and III are present with few minute cornuti in group II, in A. transversum cornuti group II and III are present with a number of shorter spines in group II).In the female genitalia the dilated, membranous ductus bursae with a 90 rotation just before corpus bursae is characteristic.
Biology: Unknown.The specimens were attracted to light in March at altitudes ranging from 460 m to 1400 m.
Distribution: Known only from a few scattered localities in the southern part of the island of Gran Canaria, Spain.

Apatema brunneum
Description: Adult.Wingspan 10.5-13 mm.Labial palp slender, upturned, segment 2 creamy white, laterally and basally dark-grey, with dark-grey rings distally, segment 3 black, dorsally and apically creamy white.Antenna black, with indistinct dark-grey rings in female.Head, neck and thorax yellowish brown, head in particular mottled with dark brown; tegula yellowish brown, dark brown towards base.Forewing dark brown mottled with yellowish brown, apically black with a golden shine; base black, center of wing an irregular oblique yellowish brown fascia bordered by four black spots; costal spot yellowish, mottled with brown, extending to tornal spot, forming an angulated fascia; fringe grey.Hindwing grey with grey fringe.
Variation: The species is rather uniform in appearance, but the four black spots display some variation; they are often well separated, but the outer pair may be confluent.
DNA barcodes (Figs 64-66, Tables 1-2): Four specimens were barcoded with very low internal variation in COI (uncorrected p = 0.0015).The specimens are placed as sister to group comprising A. transversum, A. stadeli and A. skulei.A. brunneum is clearly differentiated from all other species with uncorrected p distance to other species ranging from ca 0.06 (A.lucidum and A. stadeli) to 0.1106 (A.confluellum).Barcode Index Number: AEC2839.
Diagnosis: A. brunneum resembles several other Apatema species, especially A. mixtum and A. fasciata.It can be distinguished by the yellowish brown fascia and the overall golden shine.In the male genitalia the relatively large saccus and the large cornuti group I, turned about the longitudinal axis, giving an impression of the spines being pointing in different directions, are characteristic.In the THE GENUS APATEMA WALSINGHAM, 1900 IN THE CANARY ISLANDS AND MADEIRA female genitalia the dilated and membranous part of ductus bursae, the twisted ductus seminalis and the rotated corpus bursae are characteristic.
Biology: Early stages unknown.The adults were flying actively in the evening sunshine, and later attracted to artificial light, in mixed forest at altitudes from 750 m to 1050 m.
Distribution: Only known from a few forest-localities in the island of Tenerife, Spain.Etymology: The species is named after the brown colour of the forewing, from the Latin adjective brunneus (= brown).
Remarks: The female genitalia is figured by GOZMÀNY (2008: 341 and 438  Description adult: Wingspan 10-12.5 mm.Labial palp slender, upturned, segment 2 creamy white, base laterally and towards apex dark-grey, segment 3 black, dorsally and apically creamy white.Antenna black, with indistinct dark-grey rings in female, uniformly dark-grey in male.Head creamy white, neck and thorax creamy white mottled with dark brown; tegula creamy white, dark brown towards base.Forewing grey-brown mottled with yellowish brown, dark-grey and black, especially towards apex; base black; centrally an irregular, slightly oblique yellowish brown fascia bordered by four black spots, the colour of the wing laterally of the fascia only slightly darker; costal spot yellowish white, extending to tiny white tornal spot, forming a diffuse fascia; fringe grey.Hindwing grey with grey fringe.
Variation: The species varies in appearance of the four black spots, they are often well separated but may be confluent in pairs.
DNA barcodes (Figs 64-66, Tables 1-2): Four specimens were barcoded with some internal variation in COI (uncorrected p = 0.0055).The specimens are placed in an unresolved trichotomy with A. skulei and A. stadeli.A. transversum is differentiated from all other species with uncorrected p et al. (2003).The threshold is not entirely unproblematic, however, as some species (A.minimum, A. coarctella and A. grancanariae) have within species distance values that are above 0.02.For A. minimum and A. coarctella the average within-species diversity is approximately as high as the average distance between A. griseum and A. pseudolucidum.This clearly indicate that the species requires further investigation.Both MrBayes analyses further support the taxonomic arrangement as all recognised species are monophyletic in the analysis with high support values between 0.98 and 1.The DensiTree rendering of the BEAST analysis (Fig. 66) further illustrates this as all species are rendered as dense and compact, while many deeper relationships between species or species groups are less well defined.The analyses also confirm that A. fasciata and A. coarctella show considerable intraspecific geographical diversity.A. coarctella (Table 3) is divided into three populations comprising specimens from Tenerife, La Palma, and Gran Canaria/Fuerteventura/Lanzarote, respectively.The three populations all display average inter-population uncorrected p distance > 0.02.Similarly, A. fasciata (Table 4) is divided into three populations comprising specimens from Gran Canaria, Fuerteventura, and Lanzarote, respectively (although we were able to include only one specimen from the latter locality).Again the three populations all display average inter-population uncorrected p distance > 0.02.The analysis of all available Apatema sequences (Fig. 65) indicates that the species found on the Canary Island and Madeira do not comprise a monophyletic group as A. confluellum and A. minimum are placed in a well-supported (pp > 0.9) clade with the species A. baixerasi, A. mediopallidum, and A. whalleyi from continental Europe.One A. mediopallidum (PHLSA273-11) from Valencia in mainland Spain groups with A. coarctella and actually renders the latter species non-monophyletic in the majority rule tree, while another A. mediopallidum (LEFIJ5483-16) from Cyprus is also placed deeply within the Canarian Apatema, albeit not clearly associated with any species (Fig. 66).We have not examined either of the A. mediopallidum specimens, but we consider it likely that their placement in the analysis is due to some mitochondrial DNA artefact caused by external factors such as Wolbachia based on similar observations in previous studies (e. g.KONDANDARAMAIAH et al., 2013, SIMONSEN et al., 2019).
The results presented here show that the genus Apatama is very diverse in the Canary Islands.18 of the currently known 28 Apatama species occur in these islands, with the other ten species being found in continental Europe, the Mediterranean islands or North Africa.There is, however, hidden diversity among Apatama species occurring outside of the Macaronesian islands as well as evident from unpublished morphological and genetic studies of the genus (P.Huemer and J. ˇumpich in litt.).We have not found any overlap between species occurring in the Canary Islands and elsewhere.Only one species from continental Europe, A. mediopallidum has been recorded from the Canary Islands (KLIMESCH, 1985: 137), but the records are due to misidentification. A. mediopallidum should be deleted from the list of Canary Island Lepidoptera (VIVES MORENO, 2014: 107).
Table 1.-Species affiliation, voucher and BOLD ID, and collection localities for all species included in the molecular analyses as described in the text.
Fig. 64.-Consensus tree from the 10 million generation analysis in MrBayes with 25% burnin of the DNAbarcodes of the Apatema species occurring in the Canary Island.Numbers below the branches are posterior probability values.

Fig. 65 .
Fig. 65.-Consensus tree from the 20 million generation analysis in MrBayes with 25% burnin of all available DNA-barcodes of Apatema species.Numbers below the branches are posterior probability values.

Fig. 66 .
Fig. 66.-DensiTree rendering of the 10 million generation analysis in BEAST with 25% burnin of the DNAbarcodes of the Apatema species occurring in the Canary Island.

Table 3 .
-Average uncorrected p distance within and between populations of Apatema coarctella as described in the text.

Table 4 .
-Average uncorrected p distance within and between populations of Apatema fasciata as described in the text.