Corresponding author: Pasi Sihvonen ( email@example.com )
Academic editor: Sven Erlacher
© 2017 Peder Skou, Dieter Stüning, Pasi Sihvonen.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Skou P, Stüning D, Sihvonen P (2017) Revision of the West-Mediterranean geometrid genus Ekboarmia, with description of a new species from Portugal (Lepidoptera, Geometridae, Ennominae). Nota Lepidopterologica 40(1): 39-63. https://doi.org/10.3897/nl.40.10440
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.
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.
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.
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.
The study is based on material housed in the following collections: Private collection of Bernd Müller, Berlin, Germany (Müller); Muséum National d’Histoire Naturelle, Paris, France (
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 (
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
The genitalia and the abdomen were prepared following methods described by
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 (
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.
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 (
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.
Ekboarmia Wehrli, 1943, Gross-Schmetterlinge der Erde 4 (Supplement.): 517. Type species: Boarmia atlanticaria Staudinger, 1859 [replacement name].
Rhoptria Gumppenberg, 1892, Nova acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum 58: 299. Type species: Boarmia atlanticaria Staudinger, 1859. [Junior homonym of Rhoptria Guenée, 1857 (Geometridae: Ennominae)].
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.
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
Wing venation of E. atlanticaria (Staudinger, 1859) male (Portugal: Province Algarve, Salema, 19.iv.2007, slide Sihvonen 2156, coll. Skou). 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.
Male abdominal sternite A3 has a small setal comb in Ekboarmia, shown here in E. sagnesi Dufay, 1979. 3–4. Illustrated from a specimen from Spain: Leon, Valle de Luna, 1200 m, 2.vii.2011 (slide Sihvonen 2157, coll. Skou). 5. The setae are detached easily during preparation, revealing an oval field (specimen from France: Dept. Isère, Valle de la Romanche, 1200 m, 22.-23.vi.1999, slide Sihvonen 2095, coll. Skou). 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.
Uncus triangular, with slightly elongated tip, apex rounded. Socii absent. Gnathos arms fused ventrally, upturned, plate-shaped. Valva evenly narrowing towards apex, costa sclerotised, apical part of valva with narrow, setose ridge. Juxta sclerotised, with two large arms, inner and/or distal margins serrate or dentate. Saccus elongated, upturned medially. Phallus deeply cleft distally, with dorsal and ventral sclerotised process, latter shorter, gradually narrowing to pointed tip, dorsal one longer, spatulate, with narrow base which is bent by everted vesica. Caecum with sclerotised band on ventral side only, dorsal side membranous. Vesica slightly enlarged at base, without cornuti.
Papillae anales rather short, oval, setose. Lamella antevaginalis sclerotised, loosely V-shaped transverse band. Lamella postvaginalis weakly sclerotised, rounded plate. Ductus bursae short, fluted, membranous. Colliculum elongate, sclerotised. Corpus bursae elongated, oval, membranous, without sclerotisations. Signum stellate, of medium size, small or absent.
Probably all species feed on Juniper needles. Larva of E. atlanticaria has been recorded on (
Larvae of E. sagnesi (see
Ekboarmia has recently been classified in Ennominae tribe Boarmiini (
atlanticaria atlanticaria Staudinger, 1859, Stettiner Entomologische Zeitung 20 (7–9): 218, (Boarmia?). Syntypes 3 ♂♂, 3 ♀♀ (MNHU), 1 ♂ (ZFMK; genitalia dissected by E. Wehrli, prep. number 5199). Spain: Andalusia, near Chiclana (examined externally, illustrated also in Albers & Warnecke (1941), plate II, figs. 9, 12 (on p. 332).
= atlanticariagracilis Albers & Warnecke, 1941, Mitteilungen der Münchner Entomologische Gesellschaft 31 (1): 115, pl. 4, fig. 40, (Boarmia). Syntypes 4♂♂, 5 ♀♀ (2 syntype ♂♂ in Universität Hamburg, Centrum für Naturkunde, Germany), [Morocco]: Mehedya (examined externally, based on a photograph in the original description). Downgraded from subspecific rank of Ekboarmia atlanticaria gracilis (for instance
atlanticaria holli Oberthür, 1909, Ètudes de Lépidoptérologie comparée 3: 412, pl. 27, fig. 152, (Hemerophila). Holotype male (ZFMK), Algeria: Alger [Originally as sp.] (examined, including genitalia, dissected by E. Wehrli, prep. number 5193).
Infrasubspecific names, probably unavailable: pallidior Lucas, 1956 (f.).
139 specimens (100♂♂, 39♀♀; data provided in a Suppl. material
Diagnostic external characters (indicated) of Ekboarmia species. 9. E. atlanticaria male (Staudinger, 1859), Portugal: Salema, 40 m, 19.iv.2007 (coll. Skou). 10. E. fascinataria male (Staudinger, 1900), Algeria: Hammam Righa, April 1928 (coll.
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.
. The holotype is large, greyish-brown (Fig.
Ekboarmia adults. 13a. E. atlanticaria ♂, syntype, Spain: Andalusia, near Chiclana (coll. MNHU); 13b. E. atlanticaria ♂, Italy: Sardinia, Teulada, 8-11.xi.1934 (coll.
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.
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 (
Larva monophagous. It has been found on Juniperus phoenicea (
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
BIN: BOLD: AAD7906 (n=11 from Morocco, Portugal and Spain including the Balearics belonging to nominal subspecies) (Fig.
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
Taxon gracilis from Morocco has been considered valid at subspecies level (e.g.,
fascinataria Staudinger, 1900, Deutsche entomologische Zeitschrift Iris 12: 395, pl. 6, fig. 3, (Boarmia). Holotype female (MNHU), [Algeria]: Teniet el Had (examined externally, illustrated also in Albers & Warnecke (1941), plate II, fig. 10 (on p. 332).
65 specimens (22♂♂, 43♀♀; data provided in Suppl. material
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
Male genitalia of Ekboarmia species, diagnostic characters are indicated and explained. Scale (where shown) is 1 mm. 18a. E. atlanticaria genitalia, Portugal: Prov. Algarve, Salema, 40 m, 19.iv.2007, slide PS2053 (coll. Skou); 18b. E. atlanticaria phallus, Portugal: Prov. Algarve, Salema, 40 m, 19.iv.2007, slide PS2053 (coll. Skou); 18c. E. atlanticaria vesica, Portugal: Prov. Algarve, Salema, 40 m, 19.iv.2007, slide PS2053 (coll. Skou); 18d. E. atlanticaria juxta, Portugal: Prov. Algarve, Salema, 40 m, 19.iv.2007, slide PS2053 (coll. Skou); 19a. E. fascinataria genitalia, Algeria: Hammam Righa, May 1928, slide 2307-DS (coll.
Probably bivoltine. Only rather few and imprecise records available: February, March, April, May, June, October.
No data available.
E. atlanticaria, see text above.
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 (
E. fascinataria has been recorded from France (Llauro, Department Pyrénées-Orientales by
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
herrerai Expósito Hermosa, 2007: SHILAP Revista de Lepidopterologia 35 (138): 269, fig. 1–2, (Ekboarmia). Holotype male (coll. A. Expósito Hermosa, Móstoles, Madrid, Spain), Spain, Andalusia: Jaén, Hornillos, Puente de Guada (genitalia examined). Junior synonym proposed by
13 specimens (6♂♂, 7♀♀; data provided in a Suppl. material
Forewing medial area concolorous with wings or darker, particularly near inner margin. Forewing postmedial line can be rather smooth or distinctly dentate.
Female genitalia of Ekboarmia species, diagnostic characters are indicated and explained. Scale (where shown) is 1 mm. 22a. E. atlanticaria genitalia, Spain: Prov. Huelva, ESE Mazagon, 20 m, 21.ix.2009, slide 2309-DS (coll.
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.
Larva monophagous on Juniperus. Reared on Juniperus communis (C. Tautel pers. comm.,
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.).
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.
Neighbour-joining tree of Ekboarmia taxa, with the position of E. miniaria sp. n. from Portugal highlighted. Tree was generated under the K2P nucleotide substitution model of 20 barcode sequences as implemented on BOLD. The position of E. sagnesi within E. atlanticaria is likely to be an artefact, resulting from short barcode sequences. The taxonomic status of E. atlanticaria subspecies holli from Tunisia needs further research.
There are no similar species in Europe. Rather uniform greyish brown colour and small angle in forewing postmedial line near costa are diagnostic.
Holotype male, pinned, with genitalia on a separate slide. Original labels: HOLOTYPE/ Ekboarmia/ miniaria [red rectangle label]; Portugal Grandola/ Ameiras de B.[aixo] [38°14.29’N; 8°32.42’W]/ 20.vi.2011/ A. & Z. Laštůvka lgt.; Prep. number 2050/ Pasi Sihvonen (in coll. Skou, to be deposited at Zoological Museum, University of Copenhagen, Denmark).
Paratypes 5 ♂♂ and 5 ♀♀, all with same type label: PARATYPE/ Ekboarmia/ miniaria [red rectangle label]. One female will be deposited at
June specimens are dark brown; the specimen taken in October is light brown, except terminal area, which is distinctly dark brown (see Figs
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.
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.
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
Habitat of Ekboarmia miniaria sp. n., showing the location where the first specimen was found in 1995 by Brian Elliott. Portugal: Lagoa de Santo André, 24.ix.2009. Photo by Peder Skou.
Habitat of Ekboarmia miniaria sp. n., showing the location where majority of specimens have been found. The dominant pine is Pinus pinaster. In the undergrowth Halimium atriplicifolium, Stauracanthus genistoides and Cistus psilosepalus are among the dominant species. Portugal: Grandola, Ameiras de Baixo. Photo taken on 16.vi.2009 by Zdenek Laštůvka.
Habitat of Ekboarmia miniaria sp. n., showing the location where majority of specimens have been found. The dominant pine is Pinus pinaster. In the undergrowth Halimium atriplicifolium, Stauracanthus genistoides and Cistus psilosepalus are among the dominant species. Portugal: Grandola, Ameiras de Baixo. Photo taken on 21.vi.2011 by Zdenek Laštůvka.
There are no similar species in Europe.
BIN: BOLD: AAZ6253 (n=2 from Portugal) (Fig.
Brian Elliot, Barry Goater, and Ales and Zdenek Lastuvka are thanked for donating E. miniaria specimens to Peder Skou, which at the time were unidentified. Photographs, taxonomic and life history information have been provided by Claude Tautel (Paris, France), Michael Leipnitz and Hossein Rajaei (Stuttgart, Germany), Manfred Sommerer (Munich, Germany), Dirk Stadie (Eisleben, Germany), Tomas Molina (Lejona, Spain) and Javier Gaston (Bilbao, Spain). Claude Tautel, in addition, sent specimens of E. sagnesi on loan to check their genitalia and donated a male to the collection of