Corresponding author: Martin F.V. Corley ( email@example.com )
Academic editor: Lauri Kaila
© 2017 Martin F.V. Corley, Sónia Ferreira, Alexander L. Lvovsky, Jorge Rosete.
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: Corley MFV, Ferreira S, Lvovsky AL, Rosete J (2017) Borkhausenia crimnodes Meyrick, 1912 (Lepidoptera, Oecophoridae), a southern hemisphere species resident in Portugal. Nota Lepidopterologica 40(1): 15-24. https://doi.org/10.3897/nl.40.10938
Borkhausenia crimnodes Meyrick, 1912, a species described from Argentina, has been found resident in Beira Litoral, Portugal, constituting its first records in Europe. Borkhausenia intumescens Meyrick, 1921, described from South Africa, is shown to be synonymous with B. crimnodes, described from Argentina. COI barcode sequencing has shown that a Portuguese specimen has 100% similarity with specimens collected in South Africa. The origin of the Portuguese population remains unclear but it is likely to be connected with timber importation for the paper industry. Male and female genitalia of B. crimnodes type and the Portuguese specimens are illustrated and described.
Nine specimens of an oecophorid moth have been taken by Jorge Rosete on separate occasions from five different localities in central Portugal between July 2012 and July 2016. The first specimen, a female which appeared to belong to Oecophoridae, was examined in 2013 by Martin Corley. The following year, dissection of a male evidently belonging to the same species confirmed the family placement but could not be recognised as any known European member of that family. The possibility that it was an adventive from another continent was considered more likely than that it was a native of Portugal.
Central Portugal is dominated in many areas by plantations of eucalyptus. This, together with the fact that there are more Oecophoridae species in Australia than in any other continent, gave rise to the consideration that the species might have an Australian origin. Other Australian Oecophoridae have arrived in Europe as adventives, such as Tachystola acroxantha (Meyrick, 1885) (
In 2016, a DNA sample was taken for barcoding. This gave a 100% match on BOLD with three unidentified specimens that had been captured in South Africa. This new information opened up new possibilities. Images of the moth and the genitalia of both sexes were sent to Alexander Lvovsky (Russia), who has a wide knowledge of Old World Oecophoridae. He was able to determine that the Portuguese specimens belong to Borkhausenia intumescens Meyrick, 1921, described from Port Elizabeth in South Africa. At the suggestion of AL, MFVC studied the type of the Argentinian B. crimnodes Meyrick, 1912 at BMNH. From this it became evident that Borkhausenia intumescens is a junior synonym of B. crimnodes.
Nine specimens of B. crimnodes have so far been captured in Beira Litoral, Portugal by JR (Table
Borkhausenia crimnodes Meyrick, 1912. Holotype: Argentina: Parana. R. `06  J.F.G. Clarke male gen. prep. 4765. (BMNH), seen by MFVC.
Borkhausenia intumescens Meyrick, 1921. Two specimens. Lectotype in
Distribution of B. crimnodes in Portugal (green squares). Red stars indicate the location of paper mills.
Borkhausenia crimnodes specimens captured in Portugal. For each we list the Municipality, locality, the UTM coordinates, date of collection, genitalia preparation code when applicable and collection where the specimens are deposited.
|Pombal||Louriçal||29TNE2228||27.vii.2012||§ Corley gen. prep. 4109||J. Rosete|
|Pombal||Louriçal: Casais do Porto||29TNE2329||11.vi.2015||J. Rosete|
|Pombal||Carriço: Mata do Urso, Lagoa de São José||29TNE1128||17.viii.2012||J. Rosete|
|Ansião||2 km east of Ansião||29TNE5019||13.iv.2014||$ Corley gen. prep. 4172||J. Rosete|
|Soure||Paúl da Madriz||29TNE3142||15.vii.2016||J. Rosete|
Genomic DNA was extracted from leg tissue (Table
Both PCR reactions had 10 μL of final volume, containing 5 μL of Multiplex PCR Master Mix (QIAGEN), 0.4µM of each primer, and 1-2μL of DNA. PCR amplification was carried out on a T100 Thermal Cycler (BioRad) using the following conditions: initial denaturation at 95 °C for 15 min; 5 cycles at 95 °C for 30 s, 47 °C for 45 s, 72 °C for 45 s; then 40 cycles at 95 °C for 30 s, 51 °C for 45 s, 72 °C for 45 s; and a final elongation step at 60 °C for 10 min. The barcodes were sequenced in an Illumina Miseq platform, following the approach described by
We used OBITools (https://git.metabarcoding.org/obitools/obitools) for general sequence processing. Geneious v.6.1.5 (http://www.geneious.com/) was used for final sequence assembly. The sequence obtained was blasted against GenBank and BOLD databases. The average divergence (uncorrected p-distance) between the sequence of Portuguese specimen and Borkhausenia sp. COI sequences available in GenBank and BOLD was calculated in MEGA v.5.2.1 (
BMNH Natural History Museum, London
South Africa, Port Elizabeth. Justification for the synonymy is elaborated in the section Synonymy below.
Borkhausenia intumescens Meyrick, 1921, South Africa, Port Elizabeth. 4. Lectotype male (
Male and female similar (Figs
Forewing with R4 and R5 stalked; R5 to costa near apex; Cu1 and Cu2 separately from the cell. Hindwing with Rs and M1 separately from the cell; M3 and Cu1 from one point; Cu1 and Cu2 separately from the cell.
The male genitalia illustrated in this paper (Figs
In spite of these differences there are very good reasons for considering the Portuguese specimens and the types of B. intumescens and B. crimnodes to belong to a single species. The moths are of similar size and have the same habitus (Figs
The larval food of B. crimnodes is unknown, but is likely to be some sort of dry plant matter or debris, perhaps affected by mould or other fungi. Three specimens were taken from inside the building where JR lives, adjacent to garages. The prevalence of the species in this building is unexplained. One was taken beneath a street light, one from a porch light and three were taken at 160-watt blended mercury vapour light. Individuals have been captured in April, June, July, August, September and October, suggesting that the species is multivoltine in Portugal.
The BLAST search in BOLD (
Mean (below diagonal) and standard deviation (above diagonal) sequence divergence (uncorrected p-distances) of 658 bp fragment of cytochrome c oxidase I (COI) among pairs of species of Borkhausenia with sequences available on BOLD and GenBank.
|B. crimnodes||B. catochopis||B. fuscescens||B. luridicomella||B. minutella|
At the time of the description of Borkhausenia crimnodes, before the use of genitalia characters as a taxonomic tool had become routine practise, many genera of Microlepidoptera were understood in a much wider sense than is now the case. Borkhausenia is an example of a genus that once included large numbers of species. More than 268 species were described as Borkhausenia species (
cornutus in the phallus (male), and the long telescopic ovipositor (female). One distinct feature of B. crimnodes that is not shared by the European species is the presence of a signum, which is furthermore of a form that is unusual in the family. Likewise, the cornutus is quite different from the cornuti in the European Borkhausenia species.The male genitalia characters (gnathos longer than uncus, divided transtilla and distal process of sacculus crossing the valva) do not of themselves indicate close relationship between these species, particularly as the last two characters can be found in other families of Gelechioidea.
Querying of the BOLD database for Borkhausenia species indicates that the few species which have barcode available on BOLD and GenBank (n=5) are not closely related to one another, exhibiting more than 10% divergence between analysed species. Possibly the genus Borkhausenia, from which many species have already been removed, is still polyphyletic and its taxonomy certainly requires further research. In the absence of a world revision of this and related genera, we retain B. crimnodes in Borkhausenia.
B. crimnodes was described from Paraná in the province of Entre Rios, Argentina. Paraná is by the Paraná River, about 250 kilometres inland from Buenos Aires. B. intumescens was described from Port Elizabeth, Eastern Cape, South Africa. It is also present inland in the mountains. Those Borkhausenia species with known larvae feed on plant detritus, so this is likely to be the case also with B. crimnodes. Species with this larval behaviour are readily transported by human trade across considerable distances and resulting adults can readily find suitable food sources for oviposition. Examples include several species of Blastobasis originally from Madeira and now well established in the British Isles (
Since B. crimnodes is so far recorded from nowhere else in Europe, it would appear to have been accidentally introduced into Portugal, perhaps from South Africa or Argentina, although the possibility that it might be established elsewhere in the world cannot be ruled out. With species that are readily transported long distances by human agency, it can be difficult to establish the true country of origin. While it is puzzling that the known Portuguese sites are in Beira Litoral, remote from the main ports or airports of Lisboa and Porto, there is a smaller port at Figueira da Foz that might have provided a means of entry for Borkhausenia crimnodes. It is known that two paper mills in the region import timber from South America through Figueira da Foz port.
We are most grateful to all those who have helped towards solving the identity of B. crimnodes. David Lees in particular, but also Klaus Sattler and Martin Honey at BMNH gave valuable assistance, Ted Edwards made useful suggestions regarding Australian Oecophoridae, Brian Goodey did an excellent job of photographing MFVC’s genitalia preparations, Ernestino Maravalhas prepared the map, Joana Veríssimo processed the specimens for barcoding. This project has received funding from the European Union’s Horizon 2020 research and innovation program under grant agreement no. 668981. SF is supported by Fundação para a Ciência e Tecnologia through project grant ExpandTree (FCT-ANR/BIA-BIC/0010/2013) co-funded by the European Program COMPETE: FCOMP-01-0124-FEDER-019772 and ALL is supported by Russian state research project no. 01201351189 and the Russian Foundation for Basic Research, grant no. 14-04-00770.