Revision of the West-Mediterranean geometrid genus Ekboarmia, with description of a new species from Portugal (Lepidoptera, Geometridae, Ennominae)

The West-Mediterranean geometrid moth genus Ekboarmia Wehrli, 1943 (Lepidoptera: Geometridae, Ennominae) is revised based on morphology, life history, and DNA barcodes. It was found that wing patterns allow reliable identification of species, whereas the genitalia are rather uniform in shape and less informative, and the genetic divergence (in the COI gene) between species is considerably lower than in the Geometridae on average, ranging 1.5–3.9%. Four species are considered as valid: E. atlanticaria (Staudinger, 1859), with one putative subspecies in North Africa, E. fascinataria (Staudinger, 1900), E. sagnesi Dufay, 1979, and E. miniaria sp. n. from Portugal. Boarmia atlanticaria gracilis Albers & Warnecke, 1941 is downgraded to junior synonym of Boarmia(?) atlanticaria atlanticaria Staudinger, 1859, syn. n. E. fascinataria is removed from the European checklist due to lack of confirmed records. Larvae, where information exists, are external feeders on Juniperus needles (Cupressaceae), and adults are nocturnal, inhabiting various habitats up to 1400 metres above sea level, having apparently at least two generations per year. Adults, male and female genitalia, distribution map, and diagnostic characters are illustrated for all species. Larva and pupa of E. atlanticaria and larva of E. sagnesi are illustrated, and results of DNA barcode analysis are presented for most taxa studied.


Introduction
The concept of the geometrid genus Ekboarmia is based on Boarmia atlanticaria Staudinger, 1859, a West-Mediterranean species that occurs in the Iberian Peninsula, the Balearic Islands, Sardinia, and in North Africa. This nocturnal species is small, with the wingspan ranging 19-27 mm, and the adult is characterised by grey-brown wings and forewing postmedial line, which is slightly to strongly dentate, with a deep, z-shaped incision near the forewing costa. The larva of E. atlanticaria is green, a needle-mimic that matches (see Fig. 6) the branches of its foodplant Juniperus phoenicea (Cupressaceae) (Corley 2004;Flamigni et al. 2016). Externally, the unrelated Menophra harterti (Rothschild, 1912) and Ecleora solieraria (Rambur, 1834) are very similar to E. atlanticaria (see Flamigni et al. (2007) for diagnostic characters).
The genus Ekboarmia included for a long time two externally similar yet diagnostic species, E. atlanticaria and E. fascinataria (Staudinger, 1900), the latter occurring in North Africa from Morocco to Algeria. The few articles that deal with the genus Ekboarmia can be summarised as follows. Albers and Warnecke (1941) illustrated type specimens of both species as photographs and the male genitalia as line drawings. They provided new faunistic information, for instance a record of E. atlanticaria from Sardinia, where the species was found for the first time by Bytinski-Salz (1937), and described a new subspecies of E. atlanticaria from Morocco. In 1979 a new Ekboarmia species was described, namely E. sagnesi from the French Alps (Dufay 1979). It differs externally from the above-mentioned species by having more uniform pale brown ground colour and forewing postmedial line is less angled inwards near costa. At the time of description, only males were known. Twenty-six years later Colomb (2005) described the life history and illustrated the female of E. sagnesi, reporting that in captivity larvae of E. sagnesi feed on Juniperus as was hypothesised by Dufay (1979). Colomb (2005) also illustrated a superficial line drawing of the corpus bursae of the female genitalia, having an unusual curved structure, not reported in other Ekboarmia species. More recently, a new subspecies of E. sagnesi was described from south-east Spain, Jaén province (Expósito Hermosa 2007), on the basis of a single male. Leraut (2009, p. 185) synonymised the taxon with a note 'general appearance and genitalia in all aspects correspond to the holotype of sagnesi Dufay'. The two Spanish species (E. atlanticaria and E. sagnesi) have been treated in detail by Redondo et al. (2007), the only Italian species (E. atlanticaria) by Flamigni et al. (2007Flamigni et al. ( , 2016 and the three known Ekboarmia species (E. atlanticaria, E. sagnesi and E. fascinataria) by Leraut (2009).
Between 1995 and 2011, 11 specimens of a small Geometridae, represented by both sexes, were found in Portugal, which, due to their unusual appearance could not be placed at first in any known European genus. Later the second author was able to solve this problem, through study of the large ZFMK collection. In this paper we demonstrate that these specimens belong to genus Ekboarmia and describe them as a species new to science, revise the entire genus, provide diagnostic characters for all taxa, and new faunistic and life history information on these moths.
Distribution map is based on examination of authentic specimens, and was created by compiling label data from collection specimens. These data were supplemented by additional information from recent publications with precise distribution data including Spain and Portugal (Redondo et al. 2009) and Italy (Flamigni et al. 2007(Flamigni et al. , 2016. The taxa were delimited on the basis of combining data from various sources including morphology, biology and DNA barcodes, and applying the phylogenetic species concept of Nixon and Wheeler (1990). In the taxonomy section, the diagnostic characters are in bold italic font style.
The genitalia and the abdomen were prepared following methods described by Hardwick (1950). The phallus was photographed during dissection, and afterwards the vesica was everted from the same sample. The vesica was everted via the caecum that was cut open, by placing the phallus inside a hypodermic syringe (Sihvonen 2001). The wing venation slides were prepared by removing one pair of wings either by gentle downwards pressure or by lifting with fine forceps. The wings were then placed in a dish containing 99.5% ethanol and scales removed from both surfaces with delicate brushes. They were placed on a slide in a drop of ethanol, which was replaced by a drop of euparal, and placed under a coverslip. The preparations were left unstained.
For the DNA analyses, one or two legs were removed from each dried specimen and stored in an individual tube, which in most cases contained ethanol. DNA extraction, amplification, and sequencing of the barcode region of the mitochondrial cytochrome c oxidase I (COI) gene (658 base pairs) were carried out in the Canadian Centre for DNA Barcoding, Ontario, Canada, using standard high-throughput protocols (Ivanova et al. 2006;de Waard et al. 2008), which are described at CCDB (2013). Sequence divergence within and between species was calculated using the Kimura 2-parameter model (Kimura 1980) and the neighbour-joining algorithm (Saitou and Nei 1987), as implemented in BOLD (Ratnasingham and Hebert 2007;http://www.boldsystems.org/)

Results and discussion
The genus Ekboarmia is a uniform group in regard to the structure of the male and female genitalia and differences between the taxa are minute. This concerns both the external and internal genitalia structures. In the male genitalia the most variable interspecific features, and thus diagnostic in separating the taxa, are the shape of juxta, the angle at which the vesica opens, width of the uncus and the setose ridge in the medial part of valva. In the female genitalia the most variable structures are shape of the lamella antevaginalis, shape of the signum, and presence or absence of a signum.
With regard to the external characters, the group is less uniform. E. atlanticaria and E. fascinataria form a closely related yet distinguishable species pair, E. sagnesi has somewhat similar medial lines on wings but the overall appearance is more uniform. E. miniaria sp. n. is the most distinct in its small size, dark brown wing colour, the different course of the postmedial line in the male and most notably, the female is almost uniform dark brown, without markings. Such sexual dimorphism is not apparent in other Ekboarmia species. External differences between recognised species are conspicuous and diagnostic, allowing reliable and easy identification.
DNA barcodes were available for the majority of named taxa (Fig. 26). These proved useful and were in agreement with morphological characters. Interspecific distances were found to range from 1.5% to 3.5% in the three examined species of the genus, thus being considerably smaller than in other geometrid genera (on average within genus 10%, see Hausmann et al. 2011). Mean intraspecific variation ranged from 0.15% to 1.35% (average 0.60%). We comment on the genetic data of each species under relevant sections below. The most notable DNA barcode absence is that of E. fascinataria and despite extensive efforts, we have failed to get fresh material from North Africa for the genetic work. The most recent E. fascinataria specimens in European col-lections, both public and private, that we managed to locate are from 1979 (in coll. Sommerer). The absence of recent material may be an artefact, and not a true indication of disappearance of E. fascinataria, because we simply may not have contacted the right sources. In addition, it is known that collecting activities in many areas in Algeria and Morocco are few and geographically biased to certain locations.
The larvae of the taxa whose biology is known feed exclusively on needles of Juniper trees. Larva of E. atlanticaria has been found on Juniperus phoenicea in Portugal (Corley 2004), Spain (Staudinger 1859), and Sardinia (Flamigni et al. 2016), and is perhaps monophagous on it.
To conclude, four correlating characters sets were useful in delimiting the species (not all character sets were available for all species): external features, genitalia structures (including both the external and internal genitalia), DNA barcodes, and details of life history. Three of these characters sets were available for E. miniaria sp. n., supporting its placement in Ekboarmia.
Four species are presently considered valid: E. atlanticaria (Staudinger, 1859), with one putative subspecies in North Africa, E. fascinataria (Staudinger, 1900), E. sagnesi Dufay, 1979, and E. miniaria sp. n. from Portugal. Diversity and distribution. Four species in the West-Mediterranean area, including the French Alps, Iberian Peninsula, the Balearic Islands, Sardinia in Italy, and Morocco, Algeria, and Tunisia in North Africa (Fig. 17). External characters and abdomen. Medium-sized, wingspan 18-29 mm. Wings grey-brown or different shades of brown, markings blackish. Postmedial line prominent, on forewing it is rather smooth, dentate or even zigzag-shaped, turning inwards near costa, outer margin bordered with whitish. Fovea absent. Venation (Figs 1-2); Sc free, R1 and R2 in forewing completely coincident, often approaching the stem of R3+R4 near its base, rarely anastomosing with it. Common stalk (stem) of R3-R5 remote from upper corner of cell (= origin of M1). R3 and R5 branching off rather basally (proximally). Hindwing with large frenulum. Veins Rs and CuA1 arising remote from upper and lower corner of cell. Space between M1 and M3 large. 3A present, weak, reaching posterior margin just behind middle. Male antennae bipectinate almost to the tip (5-8 apical segments unpectinated), rami narrow, dorsally unscaled, arising proximally on ventral side from rather long flagellomeres. Female antennae filiform. Frons flat, rather broad, smooth-scaled, palps porrect or slightly curved upwards, rough-scaled, reaching well beyond frons. Haustellum moderately developed. Chaetosemata round type, near eye margin, well developed. Male hindtibia with weak hair pencil (if not everted, a weak groove visible only), hindtibia of both sexes with 2+2 spurs (index of spurs 0-2-4). Setal comb (oval field) of minute setae present on abdominal sternite 3 (Figs 3-5), weak sterno-tympanal process laterally on sternite 1+2, with short, free, distally rounded end.  Salema, 19.iv.2007, slide Sihvonen 2156. 1. Forewing fovea is often present in the Boarmiini, but absent in Ekboarmia. 2. Hindwing anal margin was partly folded during slide preparation, hence vein A3 appears at the margin of the wing (=dissection artefact). Veins are partly filled with air (white sections), particularly in forewing. The presence of setal comb correlates with presence of sterno-tympanal processes laterally on sternite 1+2, with short, free end. Probably its function is the eversion of the hind leg hair-pencil. Species with a strongly developed hair-pencil always have a strong setal comb and a long sterno-tympanal process.
Biology. Probably all species feed on Juniper needles. Larva of E. atlanticaria has been recorded on (Culot 1919) and found on Juniperus phoenicea (Zangheri 1968;Corley 2004) and that of E. sagnesi has been reared on Juniperus communis (Colomb 2005). Moths are found in a variety of habitat types from coastal dunes to open pine forests to mountain slopes with scattered trees and bushes, from sea-level to 1400 metres.
Immature stages. Larvae of E. sagnesi (see Colomb 2005) and E. atlanticaria (see Fig. 6 and Spuler 1904) are green needle-mimics that closely match the needles of their Juniperus foodplants. Larvae of both species have a row of red dots dorsally, interspersed with yellow and white markings both dorsally and laterally. Pupa typical (Fig. 7), with D2 setae in the cremaster fused and elongated.
Remarks. Ekboarmia has recently been classified in Ennominae tribe Boarmiini (Vives Moreno 1994; Leraut 1997;Hausmann et al. 2011 (Figs 8, 9, 13). Wingspan 19-27 mm, medium-sized species in the genus. Wings grey-brown, forewing medial area darker brown, general appearance somewhat smooth, slightly glossy. Forewing with antemedial line inclined, deeply V-shaped and acutely angled back towards base but often this part very narrow and only weakly visible. Forewing medial line narrow, often strongest and approaching postmedial line near inner margin. Forewing postmedial line dark-brown to blackish, inclined, weakly dentate on veins, shallowly curved inwards between veins, dark brown, deeply V-shaped and angled towards base (along vein M1) near apex, angled again towards apex before reaching costa. Forewing postmedial line not angled outwards at inner margin and medial area narrow. Dark brown to blackish streak in forewing subapical area, often appearing as an extension of postmedial line. Outer margin bordered with whitish. Subterminal line mostly present, but indistinct, dark grey or brown, more or less parallel and close to termen. Terminal line blackish, narrow, often discontinuous, stronger at vein endings. Fringes uniform grey-brown. Hindwing medial line weakly curved, strongest near inner margin, often absent towards costa. Area between medial and postmedial lines often darker than remaining wing surface. Hindwing postmedial line weakly dentate, more strongly curved outwards than medial line, outer margin whitish. Hindwing subterminal line grey or brown, diffuse. Forewing discal spots blackish, small, distinct. Hindwing discal spots smaller. Wings below grey-brown, medial lines diffuse. Postmedial line discontinuous, with blackish dentation, discal spots and terminal line blackish. Course of postmedial line differs from that of upperside, antemedial line hardly visible. Frons, collar, thorax and abdomen concolorous with wings. Antennae concolorous with wings dorsally. Morphological structure of antennae, hindtibia, tympanal organs, tergites and sternites of abdomen as mentioned in generic description.

External characters and abdomen
Variation. Spring generation much larger. Wings vary from grey-brown to different shades of brown. Medial area sometimes completely suffused with dark grey or brown. Terminal line can be continuous or discontinuous. Very rarely specimens occur with stronger dentation on postmedial line and/or a shallow incision near inner margin, producing a somewhat fascinataria-like appearance. We consider such specimens to fall within intraspecific variation of atlanticaria. The male syntype of atlanticaria from Andalusia in coll. ZFMK shows this feature.
Female genitalia (Fig. 22). Papillae anales slightly elongated, setose. Apophyses posteriores longer (about 1.6×) than apophyses anteriores.  Wehrli (1939Wehrli ( -1954 Wehrli (1939Wehrli ( -1954 Wehrli (1939Wehrli ( -1954) also]; 14f. E. fascinataria ♂, Algeria: Hammam Righa, April 1928 (underside of 14a, coll. ZFMK) [illustrated in Wehrli (1939Wehrli ( -1954  Subspecies holli. The holotype is large, greyish-brown (Fig. 13i). In ZFMK is a series of distinctly smaller specimens, collected later in the year, probably a similar phenomenon of seasonal variation as for taxon gracilis, see Remarks. The specimens from Tunisia (Fig. 13j, k) are reared and it is therefore difficult to form a judgement upon them. DNA barcodes of Tunisian specimens are distinct from other E. atlanticaria specimens (see under Genetic data). No specimens from Algeria were available for DNA barcoding. Historically the subspecies has been delimited to occur in Tunisia and Algeria. We do not propose formal changes to the current classification because more research is needed.
Distribution and abundance (Fig. 17). West-Mediterranean. In Europe only in southwestern Iberian Peninsula, the Balearic Islands, and Sardinia. There is also a record of a single specimen from southern France, undoubtedly a migrant from outside the known distribution area, of which the closest is the Balearic Islands. Outside Europe, in Morocco, Algeria, and Tunisia. In Europe a somewhat local but not rare species, which can be found in limited numbers in its localities.
Phenology. Multivoltine. Recorded from early March to early November with a few additional records in late January, February, and December. Larvae have been found in February (Staudinger 1859), in March, and mid October (M. Corley, pers. comm.). Hibernation probably takes place as egg, larva, and pupa, or a combination of these. The moths are nocturnal and both sexes come readily to light.
Habitat. In a variety of habitat types where the foodplant occurs, including coastal dunes as well as non-sandy inland localities. From sea level up to 300 m; according to Redondo et al. (2009) up to 500 m.
Genetic data. BIN: BOLD: AAD7906 (n=11 from Morocco, Portugal and Spain including the Balearics belonging to nominal subspecies) (Fig. 26). In Tunisia (putative subsp. holli) with two separate, sympatrically occurring BINs (BOLD: ABZ3161, n=3) diverging by 1.0% and (BOLD: ABZ6474, n=2) and by 2.2% from the Iberian and Moroccan populations. E. sagnesi samples were nested within E. atlanticaria groups (only short sequences available for the former, see E. sagnesi). Distance from E. miniaria 3.5% and from E. sagnesi 1.5%. Similar species. Forewing postmedial line weakly dentate and not angled outwards towards posterior margin in E. atlanticaria (strongly dentate or zigzag-shaped and sharply angled outwards towards posterior margin in E. fascinataria). Forewing medial area narrow at posterior margin in E. atlanticaria (wide in E. fascinataria). The not closely related Menophra harterti (Rothschild, 1912) has striated hindwings and forewing postmedial line is continuous, reaching wing margin below apex. The medial area of the not closely related Ecleora solieraria (Rambur, 1834) is not dark and hindwing postmedial line is rather straight (see Flamigni et al. 2007 for illustrations).
Remarks. Taxon gracilis from Morocco has been considered valid at subspecies level (e.g., Scoble 1999; Leraut 2009). The description of taxon gracilis was based on a series of nine males and females, collected in August and November, and the specimens are very small and dark greyish-brown. In our opinion type specimens belong to a summer or autumn generation, probably developed under dry conditions. Wet conditions in winter and spring yield larger specimens (see Fig. 13g). Same hypothesis has been presented by Wehrli (1943in Wehrli 1939-1954. One specimen from Moroc- Figure 17. Confirmed records of Ekboarmia species. All records of E. fascinataria from Europe have proved to be E. atlanticaria, therefore this species is removed from the European fauna until a reliable record will be provided. According to current knowledge E. fascinataria has a scattered distribution in North Africa, but this may be an artefact of low collecting activity biased towards few sampled areas. An uncertain record of E. sagnesi is marked with a question mark, see text for details. co was available for barcoding and it nested within E. atlanticaria subsp. atlanticaria specimens from the Iberian Peninsula. Due to similarity in barcodes, adjunct geographical distribution on the opposite sides of Straits of Gibraltar, and presumably seasonal variation of external appearance, we synonymize Boarmia atlanticaria gracilis Albers & Warnecke, 1941 with Boarmia(?) atlanticaria atlanticaria Staudinger, 1859.

External characters and abdomen
Variation. Wingspan varies quite a lot, the holotype female (collected in May/June) with 29 mm wingspan is the biggest specimen encountered so far. Specimens in the ZFMK collection, collected from March to May in Algeria, Hammam Righa are almost of the same size, while a large series from June consists of distinctly smaller specimens (figured as 1 st and 2 nd generation in Wehrli 1939Wehrli -1954. Wing colour varies from grey-brown to grey; males of the Algerian 2 nd generation are considerably darker than the females. In addition, a small number of specimens are found in which the forewing postmedial line is not distinctly angled outwards at inner margin. Such specimens, in which this character is somewhat transitional between E. fascinataria and E. atlanticaria, are nevertheless identified as E. fascinataria, as all other relevant characters are typical for this species. Male genitalia (Fig. 19). Generally somewhat larger and more robust than E. atlanticaria (see above), but without any noticeable differences. Base of the juxta tends to be a little longer and narrower, the incision between the furca-arms narrower. The spatulate process at apex of phallus seems to be more elongated and parallel-sided (elongated-oval in E. atlanticaria). Large differences as recently published by Leraut (2009, fig. 96) were not observed.
Female genitalia (Fig. 23). Generally as E. atlanticaria (see above), but little more robust, with slight, rather gradual differences. Apophyses anteriores and posteriores, and lamella antevaginalis are longer and stronger, lamella postvaginalis is a larger, more equally rounded plate, colliculum longer, but narrower, corpus bursae with a shallow and a more strongly rounded side. Signum a little larger, with fewer but larger marginal teeth. However, all or some of these characters may be due to variation and should be verified by study of more material. Distribution and abundance (Fig. 17). Maghrebian. In Algeria and Morocco. The abundance of the species is unknown. Apparently it is some decades since it was last collected. No confirmed records from Europe, see Remarks.
Phenology. Probably bivoltine. Only rather few and imprecise records available: February, March, April, May, June, October. Rungs (1981) gives February to June and September to November. Wehrli (1939-1954 mentions 'several generations' and figures specimens of a 1 st and 2 nd generation. Larval time and hibernation are unknown. It is unknown when the adults fly, but probably at night.
Biology. Unknown. Habitat. Unknown. Genetic data. No data available. Similar species. E. atlanticaria, see text above.
Remarks. Although the genitalia of E. atlanticaria and E. fascinataria are similar, without distinct diagnostic characters, the validity at species level is supported by distinct external appearance and perhaps sympatric occurrence in Morocco and Algeria. This is not unique, as there are examples among the Geometridae and more widely in other Lepidoptera in which external appearance is distinct but the genitalia are rather homogenous (European examples include for instance Macaria notata -M. signaria, Isturgia limbaria -I. roraria, Isturgia sparsaria -I. messapiaria (Skou and Sihvonen 2015), Nemophora degeerella (Kozlov et al. 2016)), and DNA barcodes are para-or polyphyletic (Mutanen et al. 2016 pinpoint numerous Geometridae examples), and yet these taxa are considered valid at species level. The currently available data do not support synonymisation of E. atlanticaria and E. fascinataria. DNA barcode and life history of E. fascinataria are not known at the moment. These, when available, are likely to shed more light on the taxonomy.
E. fascinataria has been recorded from France (Llauro, Department Pyrénées-Orientales by Bérard 1995) and it has been included in the French list of Lepidoptera (Leraut 1997) and in Fauna Europaea (Hausmann et al. 2011). In our opinion the specimen in Bérard (1995) is E. atlanticaria and also Leraut (2009) states in his more recent publication that the species is not yet known from Europe. Expósito Hermosa (2007, p. 270) explains the identity of a male specimen from the Island of Formentera in the Baleares, recorded on 24 August, 1968. The conclusion is vague (our translation from Spanish): "male genitalia concur with the picture of E. fascinataria from Morocco as illustrated in Albers and Warnecke (1941, p. 137, fig. 1); however, characters of E. atlanticaria and E. fascinataria share a sufficient number of correlations and an important number of coincidences, so specimens of E. atlanticaria and E. fascinataria share the same identity in Spain." Due to lack of confirmed records, we propose removal of E. fascinataria from the European list until there is a reliable record. Dufay, 1979 sagnesi Dufay, 1979, Bulletin de la Société entomologique de France 84: 129, figs 1-3, (Ekboarmia). Holotype male (MNHN), France: Hautes-Alpes, high valley of the Romanche (examined externally, illustrated in Leraut (2009), plate 47, fig. 18, paratype male genitalia illustrated in Dufay (1979) Examined non-type material. 13 specimens (6♂♂, 7♀♀; data provided in a Suppl. material 1). (Figs 11, 15). Wingspan 25-29 mm, large species in the genus. Wings dark grey, sometimes with chocolate-brown tinge, rather uniform in colour. Forewing antemedial line black, narrow, deeply angled inwards below costa. Medial line narrow, barely visible and rather straight, strongest near posterior (inner) margin. Postmedial line distinct, particularly at inner margin, black, dentate, bordered white outside, weakly V-shaped and angled towards base subapically, angled again towards costa. Postmedial line not angled outwards near inner margin and medial area rather narrow. Hindwing medial line almost straight, not reaching costa. Hindwing postmedial line black, weakly dentate, outer margin bordered with white, weak or absent near costa. Subterminal line absent or very faint. Terminal line faint, continuous. Fringes concolorous with wings, uniform. Discal spots weak or absent. Wings below uniform pale greyish brown, postmedial line and discal spots most visible, antemedial line of forewing absent. Course of postmedial line in forewing does not conform to upperside. Frons, collar, thorax, and abdomen concolorous with wings. Other structures as in E. atlanticaria, see above.

External characters and abdomen
Variation. Forewing medial area concolorous with wings or darker, particularly near inner margin. Forewing postmedial line can be rather smooth or distinctly dentate.
Male genitalia (Fig. 20). Generally as E. atlanticaria (see above). Setose ridge extends into medial part of valva in E. sagnesi (medial ridge absent in other Ekboarmia species). Vesica opens at 90 degree angle (at 135 degree angle in E. atlanticaria and E. fascinataria). Juxta arms broader distally than basally, dentate or smooth along inner margin, always dentate distally, base with roundish lobe in E. sagnesi (juxta arms gradually tapered towards apex, margin dentate at apex only in E. atlanticaria and E. fascinataria, base with elongated lobe in E. miniaria).
Distribution and abundance (Fig. 17). Southwest European. Only few and isolated populations are known: In Val de la Romanche, Dept. Hautes-Alpes, France, from Valle de Luna and Velilla de Rio Carrión (Javier Gaston, pers. comm.), both in Leon Province and Sierra de Cazorla, Jaén Province, Spain. The species is unknown outside Europe. An apparently extremely local species that appears as single specimens or in limited numbers. Weiss (1920) listed E. atlanticaria from Spain, Sierra de Albarracin. We have not been able to trace the Weiss Collection, and since E. atlanticaria has not been reported since from that area, which is unlikely to contain habitats suitable for this species, Weiss's record is surely based on a misidentification. Redondo et al. (2009) mention Weiss's record from Sierra de Albarracin also, suggesting it is possibly E. sagnesi because potentially suitable habitats are present in the area (this record is shown as a question mark in the map). E. sagnesi should certainly be looked for in Sierra de Albarracin.
Phenology. Uni-or bivoltine. In Spain from early June to mid-July, in France from early May to late June and in August. Larval period and method of hibernation are unknown. The moths are nocturnal and come to light.
Habitat. Mountain slopes with scattered trees and bushes. In France from 1150 to 1400 m, in Spain known from around 1200 to 1400 m. In Spain, Leon found on slopes with Juniperus sabina (Tomas Molina, pers. comm.).
Genetic data. E. sagnesi specimens (n=3 from France and Spain, including the holotype of taxon herrerai), grouped within E. atlanticaria, as the sister-group to E. atlanticaria specimens from the Iberian Peninsula, the Balearic Islands and Morocco (Fig. 26). Distances from E. atlanticaria 1.5% (from its Tunisian populations 2.0%) and 3.9% from E. miniaria. The barcodes of E. sagnesi were incomplete, only 127-273 bp in length, so the results must be considered tentative.
Similar species. There are no similar species in Europe. Rather uniform greyish brown colour and small angle in forewing postmedial line near costa are diagnostic.
Remarks. Colomb (2005) illustrated a superficial, hand-drawn picture of the female genitalia, which shows an unusual curved, probably sclerotised structure on the corpus bursae. We have not observed such structure in any of the E. sagnesi specimens examined, and the signum (which Colomb apparently did not illustrate) of E. sagnesi is weakly stellate, not resembling such curved structure. Second author was able, through the courtesy of Claude Tautel, to re-examine the badly damaged genitalia, preserved in a tube with glycerol pinned under the moth. Remaining sclerotised parts clearly showed the identity with E. sagnesi, membranous parts were largely lost, even though attempts were made to make them visible by staining. The curved, probably sclerotised structure was loosely floating around in the tube and not attached to a membrane. It probably was lying on the bursa only accidentally when the original drawing was made.

External characters and abdomen
Variation. June specimens are dark brown; the specimen taken in October is light brown, except terminal area, which is distinctly dark brown (see Figs 16c, h). It is not clear whether this specimen shows the normal appearance of the second generation or whether it is a strongly aberrant individual.
Male genitalia (Fig. 21). Generally as in E. atlanticaria, but smaller. Uncus wide in E. miniaria (similar, but a little less wide in E. sagnesi, narrow in E. atlanticaria and E. fascinataria). Phallus apex does not bend when vesica is everted (bends when vesica is everted in E. atlanticaria, E. fascinataria and E. sagnesi). Vesica opens at about 90 degree angle (at about 135 degree angle in E. atlanticaria and E. fascinataria). Juxta arms broader apically than basally, dentate along inner margin, base with elongated lobe (juxta arms tapering apically, dentate at apex only, base solid in E. atlanticaria and E. fascinataria; juxta arms broader apically, dentate (or smooth) along inner margin, base with roundish lobe in E. sagnesi).
Etymology. The species name miniaria (word stem based on the Latin "minima", an adjective in the nominative singular), refers to the small size of the species. The wingspan and genitalia of both sexes are smaller than in any other Ekboarmia species.
Distribution and abundance (Fig. 17). Endemic to Portugal, where it is presently known from only two localities in the southern part of the country. Seven specimens were found in one night (20.vi.2011).
Phenology. Possibly bivoltine. So far, the species has been recorded in mid-June and once between 16 September and 10 October. Nothing is known about larval time and hibernation. The moths are nocturnal and come to light.
Habitat. Open pine forests (Pinus pinaster) on sandy soil with undergrowth of herbs and scattered bushes, including Halimium atriplicifolium, Stauracanthus genistoides, and Cistus psilosepalus among the dominant species. One of the two known localities had suffered a bushfire, probably 8-10 years earlier. Found close to sea level and at 90 m (Figs 27-29).