Query of the DNA barcodes of the query taxon (the barcode of NHMUK013698469 is identical to that of NHMUK013698468 and differs by a C as opposed to a T in position 118 of the holotype, NHMUK013698467) on BOLD placed the taxon as more than 9.06–9.22% by p-distance from any other micromoth. However, as an exception, a single gelechiid came up as the top hit (Ephysteris diminutella (Zeller, 1847), Process ID LON7008-18, GenBank accession MN805721, which is 8.99% by p-distance). Of its 618 bp, 62 are ambiguously coded. When these are ignored, it is 9.0% pairwise divergent (Bioedit), whereas a clean (658 bp) sequence adjacent by its Process ID code for the same species, LON7007-18 (MN805536) is 13.07% divergent to the query taxon. This sequence should ideally be removed from consideration by BOLD. All the other ten top hits were unidentified Lepidoptera (probably lecithocerids) or identified as Lecithoceridae. The nearest identified lecithocerid species at 9.2–9.38% p-distance for two different specimens was identified as ‘Thubana exaema’ (i.e., Thubana exoema (Meyrick, 1911)) from Sri Lanka (see Discussion). The two haplotypes of the query taxon were 9.55–9.71% to the nearest two sequences identified only as a lecithocerid, and 9.68–9.82% to the nearest ones identified as a Torodora (BOLD:AAH3804 from Australia and BOLD:ABY1674 from Vietnam). In the corresponding NJ tree, the query taxon linked, albeit with long branches, to eight nearest terminals representing five BINs all from SE Asia, two of which were identified as Lecithoceridae and one of which as Torodora Meyrick, 1894, whereas it fell relatively remotely from T. exoema among the top 99 hits.

As another distance approach, using a local BLAST in Bioedit of the 429 Lecithoceridae sequences representing different BINs downloaded from BOLD, the sequences of the query taxon were 9.57–9.73% pairwise divergent to a species of Lecithoceridae (BOLD:ACT7825, LNAUT3910-15, Malaysia), 9.8–9.95% to another Lecithoceridae (BOLD:ADV1376, GMPBS211-18, Pakistan), 10.18–10.33% to Halolaguna sublaxata Gozmány, 1978 (BOLD:ABA2899, KF523781) and 10.1–10.35% to Torodora aenoptera Gozmány, 1978 (BOLD:AEG4946, MN852952), these representing the four top “hits” by local BLAST.

As a third approach, the top hit using nucleotide BLAST on Genbank on 29/01/2023 was Halolaguna sublaxata (KF523779.1) at 9.86–9.71% divergence (depending on the haplotype of the query taxon). However, equally divergent was a member of Xyloryctidae, Xylorycta cirrhophragma Meyrick, 1921 (KF404885.1), with H. sublaxata (KF523781.1) at 10.02%, showing weak signal to noise for this approach, which does not benefit from the DNA barcode dataset comprehensiveness of BOLD.

We then examined the amino acid translation of the DNA barcode region of the alignment, first examining the 429 BINs attributed to Lecithoceridae. The 161^st complete codon has the state Asparagine (N), as opposed to Serine (S), which is more usual in Gelechioidea. This is the state in NHMUK013698467–9 and is particularly prevalent in Lecithoceridae. Among the Lecithoceridae BINs, the following states (using standard amino acid abbreviations) are represented: N = 91.8%, S = 7%, K = 0.9% and D = 0.2%. By contrast, among the 221 Autostichidae BINs the proportions of states were N = 0.9%, S = 94.6%, others = 0.45% (F, L, M, T, Y), whereas among the 810 BINs of Xyloryctidae, N = 2.5%, S = 71.7%, F = 9.5% and other states 16.3% (Y, M, A, T, Q, L, K, I). An ‘N’ (generally the triplet AAT although sometime AAC) was a feature of all the sequences identified to at least a Torodorinae genus, except that an apparent reversal (see Fig. 1b) to S (the triplet AGT) occurred locally in two of four Sisyrodonta BINs (BOLD:AAN9227, BOLD:AAO6600), and among those that belong to Lecithocerinae, a reversal to S was seen in the Lecithocera nigrana (Duponchel, 1835) complex (comprising one BIN and six closely related BINs from Greece, Madagascar, and Pakistan), and in two BINs of Sarisophora Meyrick, 1904 from Australia. We found no such clear synapomorphies at subfamily level for single nucleotide or codon positions, although some group-specific patterns in complete codon 94.

Based on analysis of 17 terminals including nuclear data or 19 terminals including the two haplotypes of the query taxon, IQ-TREE automatically implemented the following partitioned model selection: COI: GTR+F+I+G4; EF-1a: Tim3e + G4; Wingless: K2P + G4; RpS5: TIM2e + G4; CAD: TIM2 + F + G4; GAPDH: TIM2u + F + I; MDH: TIM2e + G4. Although the edge-unlinked partitioned analysis in IQ-TREE provided a slightly higher log likelihood than the corresponding edge-linked analysis (-23644 to -23770; 17 terminals and -23805 +/343 to -23939 +/- 346; 19 terminals) the standard errors overlapped and the lower Bayesian Information Criterion for the edge-linked analysis was therefore preferred (48548 vs 49261). In this analysis, when 17 taxa were run (i.e., only taxa with nuclear data), the Ceuthomadarinae node showed a value of pp = 1 /bootstrap = 100%, with the Lecithocerinae + Torodorinae + Crocanthinae node with 1/64%, the Crocanthinae + Tororodinae node with 1/100%, and the Torodorinae node (Thubana + Torodora) not achieving support thresholds (0.877/64%). When the two haplotypes of the query taxon were added (Fig. 1a) and 19 terminals were run, the respective nodal values became 1/100% (Ceuthomadarinae), 0.998/93% (Ceuthomadarus + Synersaga), 0.996/80% (latter including also Nosphistica), 1/100% (Lecithocerinae), 0.996/80% (Lecithocerinae + Crocanthinae + Torodorinae), 1/99% (Crocanthinae + Torodorinae), but unsupported (0.337/35%) for the existing Torodorinae, and no support either (0.529/59%) for a sister relationship between the new taxon and Thubana cf. deltaspis Meyrick, 1935 (Fig. 1a), although the branch of the DNA barcode of the query taxon fell closest to the torodorine rather than crocanthine branch.

We attempted various phylogenetic analyses of available sequences using Phyml 3.0, an instance of which is shown (Fig. 1b). We based the last analysis on a matrix of 51 taxa and the 5408 nucleotide positions, including the 17 terminals from GenBank. This dataset is contiguous for all taxa across 483 positions of COI-5P (with all but 10 exemplars contiguous over 658 bp), using the GTR + G + I model that was selected by BIC. For these 17 taxa, dispersed across the subfamilies, COI-3P and 3–7 nuclear genes were added from the GenBank dataset referred to in Materials and Methods in order to provide a general framework for rooting and for the subfamilies (number of genes shown in Fig. 1b, otherwise just COI-5P).

The Phyml analyses never placed the new taxon within the Ceuthomadarinae nor the Lecithocerinae. These groups each appear to be monophyletic according to Wang and Li 2020 and our analysis (see Park et al. 2022 regarding the relationship of “Torodora babeana” to Homaloxestis Meyrick, 1910). Rather, they consistently placed the DNA barcode of the new taxon within a distal group of Lecithoceridae, i.e., the subfamilies Torodorinae + Crocanthinae (in the last case comprising representatives of the genera Crocanthes, Aprosoesta Turner, 1919 and Pacificulla Park, 2013). Consistently the DNA barcode of the query taxon fell in a group or clade (pp = 0.99 in Fig. 1b with Sisyrodonta ‘ochrosidera’—a complex of BINs, three BINs within Protolychnis, and Halolaguna sublaxata. BOLD places Sisyrodonta in Lecithocerinae, while Common 1990 has it in the Lecithoceridae, but our analyses always placed it in this grouping of Torodorinae. However, our analyses always failed to provide a significant support level for a sister genus or the sister group of the branch of the query taxon.

The Phyml 3.0 analysis only showed separation of Crocanthinae and Torodorinae in cases when taxon/BIN sampling was relatively dense (instance shown in Fig. 1b). More often with fewer terminals (especially among Crocanthes, which include several multi-BIN species on BOLD), resulting trees had Crocanthinae and Torodorinae partially intermingled, occasionally with the query taxon grouping with C. diula and C. prasinopis. When the sampling of identified taxa from BOLD among Torodorinae plus Crocanthinae was improved, Torodorinae formed a separate grouping as did Crocanthinae, albeit without support. The topology of Lecithocerinae was similar to that shown by Wang and Li (2020: 5, S2, S3) except for the position of Homaloxestis. The query taxon fell in a supported grouping (p = 0.99) with the three Protolychnis, three Sisyrodonta and Halolaguna sublaxata.

The MrBayes analyses on the 51-terminal 5408 bp dataset using a paratype sequence of the query taxon (NHMUK013698469, OQ339153) provided a 50 percent majority rule consensus tree (shown in Suppl. material 2, unlinked and Suppl. material 3, linked). These analyses show support for Ceuthomadarinae (pp = 1), but for Ceuthomadarus + Synersaga only in S3 (pp = 0.96). Also, they show support for Lecithocerinae (pp = 0.99 and 1 respectively), but a lack of resolution for Torodorinae + Crocanthinae. Within this “bush”, there were a number of unsupported groupings that include the query taxon (+ Halolaguna + three each of Sisyrodonta and Protolychnis with pp = 0.97, Suppl. material 3 or without support, Suppl. material 2), (Athymoris Meyrick, 1935 + Cophomantella Fletcher, 1940 + Protolychnis trigonias, pp = 0.95, Suppl. material 3 or without support, Suppl. material 2), in each case with unsupported groupings of Crocanthes and Aprosoesta (‘pancala’).

Park et al. (2022: 12, 14) note the following [syn-]apomorphies for Lecithoceridae: gnathos fused to tegumen, antennae usually longer than forewing and mesial process of gnathos of the male genitalia always downturned and laterally compressed. In the new taxon the gnathos is fused to the tegumen, the antennae are the same length as the forewing (Park et al. (2022: 15) note that a recent study of the Afrotropical fauna has confirmed that the antenna is not always longer than the forewing in several genera, including Protolychnis) but the gnathos of the new taxon is much reduced (Figs 5, 8). However, the subfamily Crocanthinae is defined as having the autapomorphic character with the gnathos always absent or reduced in the male genitalia (Park 2015: 252; Park et al. 2022: 16). At a family level, the reduced gnathos in the new taxon is therefore not inconsistent with a diagnosis of Lecithoceridae. The wing venation, including the stalking of R3, R4 and R5 and the stalking of CuA1 and CuA2 in the forewing and the absence of M2 in the hindwing is typical of a number of lecithocerid genera, as is the presence of tergal spines in the pre-genital abdomen. In the male genitalia, the narrow thorn like uncus, thin strongly sclerotised vinculum and the vestigial saccus are also often found in Lecithoceridae. The rings of outwardly pointing yellow scales arising from each flagellomere on the male antenna are also potentially synapomorphic with Protolychnis and Sisyrodonta, if those two taxa are found to belong to a clade not including Halolaguna (but see Fig. 1b).

The results from the DNA barcode query and the analyses considering distance and Neighbor Joining support a diagnosis of Lecithoceridae for the new taxon. Also, from our search of the sequences for synapomorphies, we consider the state Asparagine (N) in the 161^st complete codon of the DNA barcode to represent a ground plan synapomorphy of Lecithoceridae, which is reversed in a few genera and species (for example distally in the genus Sisyrodonta and in Lecithocera nigrana and its widely dispersed species complex). The new taxon exhibits this synapomorphy. This is a groundplan feature of all lineages of Lecithoceridae sensu Wang and Li (2020), apart from their outgroup, Martyringa (USA-SE Asia), which exhibits either a Phenylalanine (F) or Serine (S) for this position, whereas Ceuthomadarus, Synersaga and Nosphistica all show an Asparagine in the homologous position.

According to Park (2022) the subfamily Ceuthomadarinae can be distinguished from the other subfamilies based on the absence of a proboscis and the subfamily Lecithocerinae can be defined and distinguished from Torodorinae by the presence of a bridge-like structure connecting the tegumen and the valval costa of the male genitalia. The new taxon has a well developed proboscis and does not have the bridge-like structure in the male genitalia used to distinguish Lecithocerinae from Torodorinae. The new taxon is therefore the first record of a new subfamily of Lecithoceridae for Europe. However, placement, on morphology, between the current concepts of Crocanthinae and Torodorinae is not as straightforward. On the basis of Park’s definition of Crocanthinae, it would appear that the new taxon should be placed in Crocanthinae because it has a reduced gnathos. However, apart from the reduced gnathos, there is little in the adult morphology to connect the new taxon to Crocanthinae.

The suggested synapomorphies of Crocanthinae, apart from the state of the gnathos, include relatively bright coloured wings and hindwing often with similar markings like those of the forewing (Park et al. 2022: 17). The new taxon has neither wing synapomorphy. Park (2015) redefines Crocanthes (the type genus of Crocanthinae) with the following morphological characters: labial palpus with dimorphism, male with second segment long, thickened with rough scales, and third segment absent, aborted or shortened, but female with normal slender third segment. Forewing normally elongate, with usually well-developed postmedian fascia; costa gently curved beyond 2/3; apex acute or normally produced; termen usually concave medially or slightly convex; venation with R2 usually free or sometimes short-stalked with R3 and R4; R5 absent; M2 absent; CuA1 and CuA2 short-stalked; cell opened. Hindwing usually unicolorous, as wide as forewing, apex acute; termen slightly concave or strongly oblique; venation with M2 absent; M3 and CuA1 stalked basally; CuA2 arising from the ½ length of the wing. Apart from the configuration of the hindwing venation, the new taxon displays few of these characters. The labial palps are not dimorphic, in the male the second segment is not thickened with rough scales and the third segment is not absent, aborted or shortened, the forewings are not elongate and do not have a postmedian fascia and the apex is rounded. In the forewing venation, R3 is on a common stalk with R4 and R5, R5 is present and stalked with R4. M2 is present, the stalk of CuA1 and CuA2 is substantial and the cell is closed. In addition, the antennae in Crocanthes are not (with the exception of C. diula) thickened with outwardly projecting rings of scales. In the male genitalia of Crocanthes, the vinculum is U shaped, the juxta usually has caudal projections, the sacculus is developed and the cornuti in the aedeagus are complex.

In addition, the general appearance (externally and in the male and female genitalia) of the new taxon is substantially different from any species currently described within Crocanthinae, which are almost exclusively only known from the island of New Guinea and Australia, although a few genera reach other parts of Indonesia and the Philippines.

Only three existing genera of Crocanthinae are represented here (Fig. 1b) of which Aprosoesta tended to be the earliest diverging, and Crocanthes is also represented by its type species. The new taxon fell away from the sampled members of Crocanthinae when taxon sampling of identified terminals from BOLD was enhanced (as in Fig. 1b).

The subfamily Torodorinae is defined (Park et al. 2022) by the absence of a bridge-like structure connecting the tegumen and the valva and by the uncus usually thorn-like, directed caudally in the male genitalia. The present taxon has both these synapomorphies (although these are also present in most Crocanthinae). A comparison of the features of the new taxon with Park et al. (2022) shows that, although it has an unusual combination of features, it is the case that taxa with similarity to one or more characters among palps, antennae, wingshape, wing pattern, forewing venation and male genitalia, can be found within Torodorinae. In our view it is not inconsistent for a lecithocerid moth with a reduced gnathos to fall within Torodorinae. We also note that Yu et al. (2022) have recently noted 12 species of Torodora (the type genus of the subfamily) in which the gnathos is without a mesial process.

Moving to the molecular data, in interpreting the IQ-TREE analysis on the 19-terminal dataset (Fig. 1a) and based on the Phyml (Fig. 1b) and MrBayes analyses (Suppl. materials 2, 3) of the full datasets, parts of the trees with good coverage of nuclear data (Lecithocerinae in particular with 10 taxa) show relatively good resolution, whereas other parts predominated by COI-5P (in which only Thubana cf. deltaspis, Torodora aenoptera and Crocanthes prasinopis included nuclear data) show relatively weak phylogenetic signal.

The IQ-TREE analysis (Fig. 1a) provides support for placement of the new taxon within a combined clade of Torodorinae + Crocanthinae (1/99), and no support for its placement in Lecithocerinae or Ceuthomadarinae nor for a clade of Xenotorodor plus Crocanthinae (the last grouping occurred in the unlinked analysis, but without support). The analyses of the full dataset (Fig. 1b, Suppl. materials 2, 3) similarly provide no support for placement of the new taxon in Lecithocerinae or Ceuthomadarinae and, although the supporting data is almost all COI-5P, provide posterior support for the placement of the new taxon in a subclade of Torodorinae rather than Crocanthinae.

However, as noted in Park (2022), the subfamilial relationship of Lecithoceridae still reveals some problems. We confirm monophyly of Ceuthomadarinae (although inclusion of Synersaga or even Nosphistica is equivocal) and Lecithocerinae, although the relationship of these subfamilies varied among analyses and were sensitive to parameterisation, but linked analyses produced relationships consistent with that shown by Wang and Li (2020), with Lecithocerinae sister to Torodorinae + Crocanthinae. However, further work needs to be done to demonstrate that Torodorinae really constitutes a monophylum without the inclusion of all or part of Crocanthinae (i.e., whether proximal, distal, or sister to Crocanthinae in trees). Both the 17- and 19-terminal edge-unlinked IQ-TREE analysis showed support for a Crocanthinae + Torodorinae clade but the Torodorinae-only clade was unsupported by bootstrap in either analysis (or in Fig. 1b).

The morphological differences between Xenotorodor and Crocanthes are dealt with above. Also, the new taxon is morphologically divergent from all existing genera within Torodorinae and is genetically divergent from those eight traditional torodorine genera (plus Sisyrodonta) whose DNA barcode sequences were available for analysis. It displays some distinctive characters shared with some genera (including Sisyrodonta and Cophomantella which are not currently placed in either Torodorinae or Crocanthinae), but to place it in an existing genus would involve polyphyletic expansion of the concept of that genus. The genera with which the present taxon displays some potential synapomorphies (and the reason for rejecting placement in those genera) are:

Protolychnis Meyrick, 1925 (type species Lecithocera maculata Walsingham, 1881, from South Africa). The antenna of the type species was described by Lord Walsingham (Walsingham 1881) as thick yellowish ochreous. This character is present in most if not all species within the genus. The palps of the members of this genus are yellow and the antenna shorter than the forewing (Park et al. 2022: 189). The forewing pattern of P. trigonias (Meyrick, 1904) and P. chlorotoma (Meyrick, 1914) have some similarities to the present taxon (although the hindwings are pale in P. trigonias) and the forewing and hindwing venation are similar. However, in the male genitalia, the gnathos is developed with a large mesial process, and in the type species and most other species the saccus is developed, the juxta has caudal projections, the aedeagus is slender, usually longer than the valva and the female genitalia has a horseshoe shaped signum (Park et al. 2022: 190). The female genitalia also lack an appendix bursae. Further, although the venation is similar, M2 and M3 are approximated in the forewing in Protolychnis whereas in the present taxa they are almost parallel (see Park and De Prins (2019), Park and Koo (2020, 2022) and Park et al. (2022) for further information). DNA barcodes of Protolychnis show a phylogenetically close, but not sister relationship with the new taxon (Fig. 1b). A DNA barcode has been identified as the type species (P. maculata; BOLD:AAK9877; HM860216.1), and there are at least two closely related taxa widespread in Africa on BOLD (BOLD:ADT8222, not shown, is 4.1% pairwise divergent from the P. maculata sequence). Nevertheless, the sequence of the new taxon OQ339151 (NHMUK013698467) exhibits a 11.4% pairwise divergence to HM860216.1. This seems a considerable distance for a potential congeneric. It is greater than for the highly allopatric P. lychnocentra from Australia (which is clearly a Protolychnis according to Park et al. 2022) and which differs from P. maculata by 10.3% and X. stygioxanthus by 11.52%, and a likely member of the genus from China (BOLD:ACD5139; Fig. 1b) which differs from P. lychnocentra by 6.5% and P. maculata by 9.6% (Fig. 1b). P. trigonias (Meyrick, 1904) (BOLD:AAO4343; also Australia) did not group with the two other identified Protolychnis in trees; this taxon is 12.64% pairwise divergent from X. stygioxanthus; 610 bp) and 13.1% pairwise divergent from P. maculata; rather, it groups with Athymoris martialis Meyrick, 1935 and a species of “Cophomantella” (BOLD:AAL6994) (Fig. 1b, Suppl. material 2). It therefore seems doubtful that P. trigonias represents a true Protolychnis, although the missing abdomen of the type (Styloceros trigonias Meyrick, 1904) has made morphological assessment difficult (Park et al. 2022: 196).

Sisyrodonta Meyrick, 1922 (type species Sisyrodonta ochrosidera Meyrick, 1922, from Australia). This is currently a monotypic genus known from Western Australia, although DNA barcodes indicate (e.g. Fig. 1b) that there is a species complex widely spread through Australia (BOLD:AAJ6848, BOLD:ACK2022; identified as a Crocanthes, BOLD:AAN9227, BOLD:AAO6600). The forewing pattern of S. ochrosidera is black (with a purplish sheen) and yellow, the antenna and labial palps are yellow and each flagellomere of the male antenna is ringed with large yellow scales. The antenna is more lamellate than in the present taxon and Meyrick considered it to be unique (Meyrick 1922) but in our view the antennae are similar to those of the new taxon. However, the hindwings are pale in both the type species and undescribed members of the complex. More importantly, CuA1 is absent in the forewing in the genus and CuA2 and M3 are stalked in the type species. In the male genitalia, the uncus of S. ochrosidera is broad at base and strongly sclerotised throughout, the gnathos has a large, strongly sclerotised medial section which is strongly beaked and strongly projected posteriorly and in the specimen of Sisyrodonta sp. from Western Australia (NHMUK013698527; Slide no. NHMUK014331161) which we have examined, the aedeagus is thin and elongate and without a cornutus. Sisyrodonta was included in the ‘Gelechiadae’ by Fletcher (1929: 204). It was transferred to the Lecithoceridae by Sattler (1973: 250). The genus was transferred to the Xyloryctidae from the Lecithoceridae by Gozmány (1978: 263). However, Common (1990: 264) considered that the wing venation, the presence of the supplementary wing-coupling setae, and the behaviour of the adults leaves little doubt that it belongs in the Lecithoceridae. Sisyrodonta was also treated in the Lecithoceridae by Nye and Fletcher (1991: 277). We consider that Sisyrodonta is correctly placed in the Lecithoceridae although the strongly sclerotised medial section of the gnathos is unusual for the family. We place the genus, which in the full dataset analyses (see Fig. 1b, Suppl. material 3) fell in a small grouping that included Protolychnis and Halolaguna, as well as the new taxon, in the Torodorinae.

Cophomantella Fletcher, 1940 (type species Onebala elaphopis Meyrick, 1910, from India). Cophomantella was established as an objective replacement name for Cophomantis, Meyrick, 1925, a junior homonym of a frog, Cophomantis Peters, 1870 (Nye and Fletcher 1991). As a result of various combinations, “Cophomantella” applies to a few remaining species and Park et al. (2022: 24) recommend that the type species should be transferred to a genus in Gelechiidae. We have examined the type of C. elaphopis, including the wing preparation and male genitalia contained on the type slide (JFGC 8911), and apart from the reduced gnathos and somewhat reduced venation, it exhibits many of the typical features of a torodorine including the tergal spines on the abdomen, the thorn-like uncus directed caudally, the shape and orientation of the valva and the large, complex cornuti in the aedeagus. We therefore consider Cophomantella to be a further example of a genus within Torodorinae with a reduced gnathos, and we transfer it back to Lecithoceridae. The new taxon is not, however, within Cophomantella. The palps of C. elaphopis are dark, the scaling on the antennae of the male is not projected outwards, the forewing markings are different and in the forewing venation, R3 and R4 are stalked, R5 is absent and M3 is absent. Also, in the male of C. elaphopis there is an expansible pencil of hairs from the base on the forewing ventral surface along the costa (Meyrick 1910). This is not present in the new taxon. In the male genitalia of C. elaphopis, the vinculum is U shaped and the tegumen and valvae are also a different shape from those of the new taxon. The one South East Asian BIN on BOLD attributed to the genus (BOLD:AAL6994) that we analysed fell with posterior support together with Protolychnis trigonias from Australia (Fig. 1b; but see also Suppl. material 3).

Torodora Meyrick, 1894 (type species Torodora characteris Meyrick, 1894, from Myanmar). This is a genus which comprises more than 200 described species (Park et al. 2022: 271) and seems very likely to be polyphyletic. It is considered here on the basis of the resemblance of several features in the male genitalia, including the shape of the uncus, juxta, vinculum, saccus and valva and the rough texture of the tegumen in T. meifengensis Park, 2015; T. octavana (Meyrick, 1911); and T. umbriella Park & Heppner, 2022 (Fig. 5, Park et al. 2022 plates 113C, 118E, 132A). The genus is generally defined by the wing venation as follows: Forewing with R3, R4 and R5 usually on a common stalk, CuA1 and CuA2 stalked, and M2 present in both wings (Park et al. 2022, 271). The present taxon satisfies these criteria except that M2 is absent in the hindwing, although Park et al. (2022: 272) notes that venation is variable and that generic assignment should be based on a combination of characters. According to Park, the male genitalia of Torodora are characterised by the hooked gnathos, the foot shaped or variously elongated valva and the strong spinous zones on the tergites of the abdomen (Park et al. 2022: 272), although Yu et al. (2022) give mesial process of the gnathos as present or absent as a generic character and identify 12 species of Torodora in which it is absent. The latter two characters are present in the new taxon (the second occurs in the ground plan of all Lecithoceridae, Autostichidae and Xyloryctidae) but the new taxon lacks the hooked mesial process of the gnathos (which is present in the three species of Torodora referred to above). In addition, the wing pattern and thickened antenna in the male would be unusual for Torodora. The one identified species we analysed, T. aenoptera, fell in an analysis (Fig. 1b) within Torodorinae, next to a species from Australia (BOLD:AAH3804), and a species of Parkiana from Madagascar (see Park et al. 2020), however, with no support for its placement.

Thubana (Type species Thubana bisignatella Walker, 1864, from Borneo). Although it is 9.2–9.38% pairwise divergent, T. exoema (Meyrick, 1911) is the nearest identified sequenced species of Lecithoceridae to the present taxon in the NJ tree resulting from the BOLD search, while the one Thubana species analysed (Figs 1a, 1b) also shows no sister relationship to it. According to Park et al. (2022: 212), the genus is characterised by the following synapomorphic characters: the forewing commonly having a large creamy-white or light-orange triangular costal patch, with a few exceptions, and the venation with M3, CuA1, and CuA2 on a common stalk; R3 stalked with R4 and R5; R5 absent or often present; the hindwing with M2 present and closely approximated to the stalk of M3 and CuA1; the abdomen with dense spinous zones on tergites. The present taxon does not have the triangular costal patch, R5 is present in the forewing and M2 is absent in the hindwing. The tergal spines are moderate rather than dense. The male genitalia of Thubana are significantly different. The gnathos is strongly developed with a large downward mesial projection, the vinculum is U shaped and projects well beyond the base of the valva, the juxta has caudal projections and the cornuti in the aedeagus are complex and not a single spike. In the female genitalia the ductus bursae is long and usually with sclerotisation or scobination and the corpus bursae has a signum.

Halolaguna (Type species Halolaguna sublaxata Gozmany, 1978, from China). This is one of the genera which appears in a molecular clade of Torodorinae with the present taxon (along with typical Protolychnis and Sisyrodonta, Fig. 1b, pp = 0.99, albeit without support for its sister taxon, and in a clade that has posterior support (pp = 0.97) in the linked rather than unlinked MrBayes analysis, see Suppl. material 3). Morphologically there is little similarity with this genus. In Halolaguna the apex of the forewing projects and the wing pattern, antennae and colour of the labial palps are different. M2 is also present in the hindwing. In the male genitalia the gnathos is developed with a strong downward mesial projection, the vinculum projects beyond the base of the valva, the saccus is developed, the juxta has caudal projections and the cornuti in the aedeagus are complex with minute spines. In the female genitalia the ductus bursae is long and the corpus bursae has a signum.

Crocanthes (Type species Crocanthes prasinopis Meyrick, 1886: 277, from Australasia) and Aprosoesta Turner, 1919 (type species A. pancala Turner, 1919). See above for discussion of Crocanthes. Aprosoesta is mentioned separately but only because of the relative proximity of the A. pancala sequences to those of the new taxon in Fig. 1b. Aprosoesta was previously treated as a junior synonym of Crocanthes but was re-established in Park 2015 on the basis of the similar maculations of the forewing and hindwing, and characters in the forewing venation and the third segment of the labial palps in the male. In terms of morphology, Aprosoesta is as divergent as Crocanthes from the new taxon (A. ‘pancala’ BOLD: AAO6447 is about 11.5% divergent by nucleotides).

The origin of the population of this taxon in Southern Spain is not clear. It could be a previously undiscovered but long resident population. Although the Microlepidoptera of Spain seem reasonably well studied, there must be many species still awaiting discovery. However, this is a distinctive taxon, evidently common where it occurs, which is not likely to be overlooked and the habitat in which it occurs is not uncommon in Southern Spain so there is no obvious reason why it would not occur elsewhere. A second possibility is that it is a previously unknown species originating from North Africa which is now spreading into Southern Europe (although it is clearly distinct morphologically and genetically from Ceuthomadarus from NW Africa). A number of species have been found around Tarifa which are taking this route, most likely as a result of climate change making conditions hotter and harsher in North Africa, causing resident species to expand northwards. DG has recorded in this area since 2017 but did not see this taxon until May 2020. Considering the increasing numbers of X. stygioxanthus at the locations around CIMA and the discovery of specimens at Huerta Grande, arrival from Africa may explain the Spanish population. A third possibility is that this taxon is an invasive species that has come, possibly from Australia, as a result of plantation of Eucalyptus spp. or other imported plants in the area. However, the taxon does not seem to be closely related to any known or previously DNA barcoded Australian genus of lecithocerid moth, and the DNA barcoding campaign on ANIC collections was more extensive than for any comparable tropical region.