Research Article |
Corresponding author: Jadranka Rota ( jadranka.rota@gmail.com ) Academic editor: Erik J. van Nieukerken
© 2014 Jadranka Rota, Antonio Aguiar, Ole Karsholt.
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:
Rota J, Aguiar A, Karsholt O (2014) Choreutidae of Madeira: review of the known species and description of the male of Anthophila threnodes (Walsingham, 1910) (Lepidoptera). Nota Lepidopterologica 37(1): 91-103. https://doi.org/10.3897/nl.37.7928
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We review and illustrate the four species of Choreutidae recorded from Madeira – Anthophila threnodes (Walsingham), A. fabriciana (Linnaeus), Choreutis nemorana (Hübner), and Tebenna micalis (Mann) – and describe and illustrate for the first time the male of A. threnodes, as well as the biology of this Madeiran endemic. We provide brief notes on each of the species and give short diagnoses for correctly identifying them. Finally, we discuss previous misidentifications of Madeiran choreutids and the occurrence of choreutids on other oceanic islands.
The Lepidoptera fauna of the Madeira Islands consists of only 331 species (
Madeiran lepidopterology began with Thomas Vernon Wollaston who, for health reasons, stayed in Madeira for long periods between 1847 and 1855 and also made later visits to the island. His results, including descriptions of many new species, were published by himself (
Below we treat the small family Choreutidae. Currently, 413 species of choreutids are described, and most of them are found in the tropics (Rota unpublished database). They are usually diurnal and often brightly coloured. So far four species have been found in Madeira. We describe and illustrate the unknown male and the biology of the endemic Anthophila threnodes (Walsingham), and provide images and brief notes on the other three species: Anthophila fabriciana (Linnaeus), Choreutis nemorana (Hübner), and Tebenna micalis (Mann). At the end we discuss earlier misidentifications of Madeiran choreutids.
Genitalia dissections and terminology follow
We conducted standard DNA extraction from abdomens of two specimens of Anthophila threnodes (voucher codes noted below) and attempted standard PCR amplification of the DNA barcode (cytochrome oxidase subunit I) using primers LCO and HCO (
Primer pairs and sequence |
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GAP1F (AARGCTGGRGCTGAATATGT) / GAP2R (TAACTTTGCCRACAGCYTT) |
GAP3F (GTGCCCARCARAACATCAT) / GAP4R (CGGCTGGAGTARCCATAYTC) |
Data from the type material are cited literally whereas data from the other material are given in a standardized format.
AMFA – Collection of Antonio M. F. Aguiar, Funchal, Madeira
BMNH – The Natural History Museum, London, U.K.
ICLAM – Insect Collection Laboratório de Qualidade Agrícola, Madeira, Portugal
SIP – Collection of Leo Sippola, Pirkkala, Finland
ZMUC – Zoological Museum, University of Copenhagen, Denmark
Hemerophila threnodes Walsingham, 1910: 257.
Holotype. 1♀, ‘B.M. Genitalia Slide 20115 A. Diak’ | ‘Type’ ’Madeira, V. 1886, Leech, 62296’ | ‘Walsingham Collection 1910-427’ | ‘Hemerophila threnodes Wals., ♀, Ent. Monthly Mag. 46, p. 257 (1910). Type ♀ (1/1) descr. 62296’ (BMNH). 1♂, 1♀, ‘Madeira: 1858. Wollaston. BM 1858-21.’ | ‘Simaethis fabriciana L. teste Stn. Ann-Mag. N. H. (3. s) III. 210, no 4 (1859), Topotype Madeira.’ | ‘V’ [May] (BMNH); 1♂, Funchal, 20–31.iii.1995, leg. L. Sippola, genitalia slide Rota JR2013-05, DNA voucher for extraction An_th2 (ZMUC); 1♂, Cabo Girão, Cruz da Caldeira, 8.vii.1999, leg. A. M. F. Aguiar, genitalia slide Rota JR2013-04, DNA voucher for exraction An_th1 (AMFA); 1♀, Vereda da Entrosa, Arco de São Jorge, 225 m, 12.ii.2000, leg. A. M. F. Aguiar (AMFA); 3♂, 2♀, same data but la. 14.ii.2001, Urtica sp., leg. A. M. F. Aguiar & J. Jesus (AMFA, ICLAM); 2♂, 3♀, same data but 200 m, la. 29.xi.2001, Urtica sp., leg. A. M. F. Aguiar & O. Karsholt, genitalia slides Hendriksen 5240, Karsholt 5236 (AMFA, ZMUC); 2♀, same data but, 15.v.2003, leg. A. M. F. Aguiar & J. Jesus (AMFA); 1♀, AM 5089, Levada Grande, Boaventura, 285m, 4.iii.2000, larva on Urtica membranacea, leg. A. M. F. Aguiar (AMFA); 1♀, Monte, 20.iii.2002, leg. L. Sippola (SIP).
Hemerophila threnodes was originally described from one female collected in Madeira by J. H. Leech in May 1886. No exact locality was stated. Based on the material collected more recently, we provide a description of the male. We also illustrate female genitalia.
A. threnodes is characterized by its dark, almost black wings and a scattering of light bluish scales on the thoracic dorsum and forewing upperside (
Male genitalia (
Female genitalia (
Urtica membranacea Poir. ex Savigny and probably other Urtica spp. (Urticaceae).
Larvae have been found in March, May, and November, and adults have been collected in February, March, May, and July, indicating at least two broods. The adult flies during the day. It occurs in open landscapes at low altitudes.
(
A. threnodes is characterized by its blackish wings, and should be relatively easy to differentiate from the other Lepidoptera in Madeira. It resembles A. fabriciana but the wings of that species are much lighter greyish brown. Male genitalia are very similar to those of A. fabriciana, but they differ in having a much shorter spine on the phallus. Female genitalia are similar to those of other species of Anthophila, but they differ in only having a hint of spiralization in the basal third of the ductus bursae, unlike those of e.g. A. fabriciana (L.), A. abhasica Danilevsky, and A. armata Danilevsky, where the spiralization is apparent. We cannot find differences between the larvae of A. threnodes and A. fabriciana in their general appearance (the differentiation between the two would likely be possible based on the ultrastructure as seen with the scanning electron microscope, but this has not been done).
Phalaena (Tortrix) fabriciana Linnaeus, 1767: 880.
1♀, Vereda da Entrosa, Arco de São Jorge, 225 m, 14.ii.2001, leg. A. M. F. Aguiar (ICLAM).
Urtica sp. (Urticaceae).
This is a new record for Madeira. The single specimen was collected at the same place as several A. threnodes. We examined photographs of this specimen deposited in the BMNH and are of the opinion that it is correctly identified.
A. fabriciana resembles A. threnodes, but differs by its greyish brown wings. It has the cream-white spots apically at costa and dorsum connected by a light zigzag-line, and has a white streak near the margin in the hindwings (
Tortrix nemorana Hübner [1799]: pl. 1, fig. 3.
Serra d’Água, Pousada dos Vinhaticos, 660 m, 10♂, 13♀, 12–13.ix.1975, leg. O. Lomholdt & N. L. Wolff; São Vicente, sea level, 1♀, 16.vi.1993; 5♂, 3♀, same data, but larva on Ficus carica, leg. O. Karsholt (ZMUC); São Vicente, Ribeira do Inferno, 79m, 3♂, 12.ix.1996, leg. F. Aguiar & J. Jesus (AMFA, ICLAM); Ponta do Pargo, Porto do Pesqueiro, 311 m, 1♂, 1♀, 23.iv.1994, leg. A. M. F. Aguiar (AMFA); Fajã da Nogueira, 600–1000 m, 1♀, 8.x.1994, leg. O. Karsholt (ZMUC); Curral das Freiras, 850 m, 1♂, 20.ix.1997, leg. O. Karsholt (ZMUC); same data, but 597 m, 3♂, 2♀, la. 16.iv.1998, Ficus carica, leg. F. Aguiar & J. Jesus; 3♂, 1♀, same data, but 7.v.1998 (ICLAM); Estreito de Câmara de Lobos, Levada do Norte, Garachico, 1♂, 1♀, 538 m, 26.vii.2001, J. Jesus (AMFA, ICLAM).
Ficus carica L. (Moraceae).
This species is common wherever there are Ficus carica trees. Larvae, hiding under a thin web, skeletonise mainly young leaves (
The almost square, brown forewings, and the black and yellow hindwings make C. nemorana unmistakeable among Madeiran Lepidoptera (
Choreutis micalis Mann, 1857: 181.
Funchal, Lido, 1♂, 20.viii.1974, leg. E. Traugott-Olsen, genitalia slide Wolff 4298; same locality, but 50 m, 1♀, 18.x.1997, leg. O. Karsholt (ZMUC); Fajã da Nogueira, 1000 m, 4♂, 2♀, 23.viii.1974, leg. E. Traugott-Olsen, genitalia slide Wolff 4294, Rasmussen 4849; same locality, but 600–1000 m, 2♂, 8.x.1994, leg. O. Karsholt, genitalia slide Hendriksen 5765; same locality, but 700 m, 3♂, 2♀, 18.ix.1997, leg. O. Karsholt; same locality, but 1♀, 500 m, 25.xi.2001, leg. O. Karsholt (ZMUC); same locality, but 1♀, 864 m, la. 14.vii.1998, Helichrysum foetidum, leg. F. Aguiar & J. Jesus (ICLAM); Caniço, 2♂, 12–18.ix.1977, leg. O. Lomholdt & N. L. Wolff (ZMUC); Ponta de São Lourenço, sea level, 9♂, 4♀, 24–27.vi.1993, 1♀, 11.vii.1993, 1♂, 17.ix.1997, leg. O. Karsholt (ZMUC); Machico, sea level, 1♂, 27.vi.1993, leg. O. Karsholt (ZMUC); Achadas da Cruz, 725 m, 3♂, 2♀, 8.vii.1993, leg. O. Karsholt (ZMUC); Porto Moniz, sea level, 1♀, 9.x.1994, leg. O. Karsholt (ZMUC); Santo da Serra, 700 m, 8♂, 2♀, 26.x.1994, larva on Helichrysum foetidum, leg. O. Karsholt, genitalia slide Hendriksen 5766 (ZMUC); 1♂, Estreito de Câmara de Lobos, Jardim da Serra, 1130 m, 22.x.1998, leg. F. Aguiar & J. Jesus (ICLAM); 1♀, Porto da Cruz, Chão das Feiteiras, 1251 m, 12.xi.1998, leg. A. M. F. Aguiar (AMFA); 1♂, 5♀, Santana, Pico, Posto Agrario, 411 m, la. 12.xi.1998, Arctium minus; same data, but 1♀ 1.vi.1999, leg. F. Aguiar & J. Jesus (ICLAM); 1♂, Boaventura, Vereda da Entrosa, 130 m, 12.ii.2000; 5♂, 3♀, same data, but 175 m, la. 14.ii.2001, Helichrysum melaleucum, leg. F. Aguiar & J. Jesus (AMFA, ICLAM); 1♂, 1♂, Estreito de Câmara de Lobos, Levada do Norte, Garachico, 538 m, 26.vii.2001 leg. J. Jesus (ICLAM); 1♀, Machico, Funduras, 605 m, 12.vi.2003, leg. J. Jesus (ICLAM); 1♀, Santana, Achada do Gramacho/ Cais de São Jorge, 267 m, 21.vii.2011, leg. J. Jesus (ICLAM); 1♂, Camacha, Levada dos Tornos, direction Camacha – Monte, 788 m, Cirsium vulgare, la. 13.vi.2013, leg. S. Fontinha (ICLAM).
Arctium minus (Hill.) Bernh., Cirsium vulgare (Savi) Ten., Helichrysum foetidum (L.) Cass. and Helichrysum melaleucum Rchb. (Compositae).
Larvae have been found in October, November, February, June, and July, and adults have been collected in all months from June to November, at altitudes from sea level to 1250 m. The adult flies during the day and comes to light. This is a widespread species. In addition to being present on Madeira, it occurs on the Canaries, throughout the Mediterranean, from central and eastern Europe throughout southern Asia to China and Japan, as well as possibly on Marianne Islands, Java, and New Zealand (
With its sub-triangular forewings with black and metallic markings adult T. micalis is unique among Madeiran Lepidoptera (
The successfully amplified A. threnodes COI haplotype is unique and it differs from the A. alpinella haplotype in 14 bases, and from the A. fabriciana haplotype in 12 bases. In the resulting phylogenetic tree, Anthophila threnodes and A. fabriciana are sister species, but without statistical support (PP or posterior probability = 0.81). Together, they are strongly supported as being the sister group to the North American species Anthophila alpinella (PP = 1).
One of the aims of this paper was to provide the necessary information for correctly identifying Madeiran choreutids because misidentifications have been common in the past. For example, A. threnodes was misidentified as A. fabriciana by
Furthermore, Tebenna bjerkandrella (Thunberg), a species similar to T. micalis, has been noted as found in Madeira several times (e.g., by
The biology of the immature stages of A. threnodes is typical for the whole family. Spinning a thin web on or around the young leaves on which the larva feeds is known from a number of other genera (
Finally, we discuss other choreutids that are known to occur on oceanic islands. For example, Brenthia leptocosma Meyrick is currently known only from Mauritius (
As A. threnodes is endemic to Madeira, it would be interesting to firmly establish its sister species so as to try to ascertain the origin of this species. We attempted to do so using molecular data, but were highly limited by the small amount of sequence that we obtained from our material and therefore our finding that A. fabriciana is the sister species of A. threnodes can only be taken as a preliminary result although this also appears to be supported by the close similarity of the two species in their external appearance, as well as their genitalia. Ideally, molecular work employing phylogeographic methods, conducted on freshly collected material of a large number of specimens of A. alpinella, A. fabriciana, and A. threnodes, as well as other potential close relatives of A. threnodes, would provide an answer with more certainty than we can do at this point.
We are grateful to Kevin Tuck, The Natural History Museum, London, U.K. for access to the collections and for information and photographs of specimens; to Jurate De Prins, Royal Museum for Central Africa, Belgium; and Leo Sippola, Pirkkala, Finland for loan of specimens. We greatly appreciate the input on an earlier version of the manuscript provided by Martin Corley and Marko Mutanen. JR was funded by the Finnish Kone Foundation while doing this research. OK´s work on the Lepidoptera of Madeira project was supported by a grant from the Carlsberg Foundation (Denmark).
Species | voucher code | museum (ID No.) | CAD | COI-begin | COI-end | EF1a-begin | EF1a-end | GAPDH | IDH | MDH | RpS5 | wingless |
---|---|---|---|---|---|---|---|---|---|---|---|---|
Prochoreutis inflatella | Pr_in_CT_1 | - | 850 bp | 593 bp | 756 bp | 506 bp | 675 bp | 691 bp | - | 321 bp | 590 bp | 400 bp |
JQ958422 | KJ844047 | HQ533107 | HQ541504 | HQ541504 | JQ958449 | - | KJ844059 | JQ958499 | HQ541579 | |||
Anthophila threnodes | An_th2 | ZMUC | - | 326 bp | - | - | - | 580 bp | - | - | - | - |
- | KJ844048 | - | - | - | KJ844052 | - | - | - | - | |||
Anthophila fabriciana | An_f_BE_2 | - | 412 bp | 584 bp | 774 bp | 517 bp | 675 bp | 661 bp | 402 bp | 405 bp | 596 bp | 365 bp |
JQ958399 | JQ958507 | HQ533054 | HQ541451 | HQ541451 | JQ958431 | KJ844055 | JQ958543 | JQ958477 | HQ541532 | |||
Anthophila alpinella | An_a_CA_2 | - | 412 bp | 588 bp | 774 bp | 495 bp | 645 bp | 690 bp | 697 bp | 406 bp | 596 bp | 400 bp |
JQ958398 | KJ844049 | HQ533053 | HQ541450 | HQ541450 | JQ958430 | JQ958454 | JQ958542 | JQ958476 | HQ541531 | |||
Anthophila sp. (Peru) | An_sp651 | ZMUC | - | 326 bp | - | - | - | 298 bp | - | - | - | - |
- | KJ844050 | - | - | - | KJ844053 | - | - | - | - | |||
Anthophila sp. (Rwanda) | An_JDP1_5570 | RNCA (ENT 5570) | 414 bp | 620 bp | 453 bp | 177 bp | 313 bp | 691 bp | 315 bp | - | 550 bp | 342 bp |
KJ844057 | KJ844051 | KJ844051 | KJ844058 | KJ844058 | KJ844054 | KJ844056 | - | KJ844060 | KJ844061 |