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The genus Apatema Walsingham, 1900 in the Canary Islands and Madeira, with description of 13 new species (Lepidoptera: Autostichidae, Oegoconiinae)

El género Apatema Walsingham, 1900 en las Islas Canarias y Madeira, con descripción de 13 nuevas especies (Lepidoptera: Autostichidae, Oegoconiinae)

P. Falck
Dinamarca
O. Karsholt
Zoological Museum Natural History Museum of Denmark Universitetsparken, Dinamarca
T. J. Simonsen
Natural History Museum Aarhus, Dinamarca

The genus Apatema Walsingham, 1900 in the Canary Islands and Madeira, with description of 13 new species (Lepidoptera: Autostichidae, Oegoconiinae)

SHILAP Revista de lepidopterología, vol. 49, núm. 194, pp. 273-318, 2021

Sociedad Hispano-Luso-Americana de Lepidopterología

Received: 28 December 2020

Accepted: 08 February 2021

Published: 30 June 2021

Abstract: The genus Apatema Walsingham, 1900 in the Canary Islands and Madeira is revised based on external morphology, genitalia and DNA barcodes. We recognize 18 species, 13 of which are described as new: Apatema confluellum Falck & Karsholt, sp. n. (Spain: Tenerife), Apatema minimum Falck & Karsholt, sp. n. (Spain: Lanzarote and Fuerteventura), Apatema lapalmae Falck & Karsholt, sp. n. (Spain: La Palma), Apatema sallyae Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema lanzarotae Falck & Karsholt, sp. n. (Spain: Lanzarote), Apatema mixtum Falck & Karsholt, sp. n. (Spain: Tenerife), Apatema pseudolucidum Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema griseum Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema grancanariae Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema brunneum Falck & Karsholt, sp. n. (Spain: Tenerife), Apatema transversum Falck & Karsholt, sp. n. (Spain: Gran Canaria), Apatema stadeli Falck & Karsholt, sp. n. (Spain: Tenerife) and Apatema skulei Falck & Karsholt, sp. n. (Spain: Gran Canaria). The monotypic Dysallomima Gozmány, 2008, syn. n. is treated as a new synonym of Apatema and its type species Lampros coarctella Rebel, 1896 is transferred as Apatema coarctella (Rebel, 1896) comb. n. Apatema helleri (Rebel, 1910), comb. n. is transferred from the Blastobasidae. A. mediopallidum Walsingham, 1900 is removed from the list of Lepidoptera of the Canary Island. Morphology of all species is described and figured in detail. All species occurs in the Canary Islands and only one, A. fasciata (Stainton, 1859) occurs both in these islands and in the Madeiran archipelago. Records of A. mediopallidum Walsingham, 1900 from the Canary Islands are based on misidentifications. Analyses of DNA barcodes support the identification distinctiveness of the species as each species appear monophyletic, well-supported and genetically isolated in the analyses. The only exception is A. coarctella, which could not be rendered monophyletic with respect to a single A. mediopallidum specimen, when sequences from non-Canarian Apatema were included in the analysis. We argue that this result is an artefact.

Keywords: Lepidoptera, Autostichidae, Apatema, new species, DNA barcoding, Canary Islands, Spain, Madeira, Portugal.

Resumen: Se revisa el género Apatema Walsingham, 1900 en las Islas Canarias y Madeira basado sobre la morfología externa, genitalia t ADN código de barras. Reconocemos 18 especies, 13 de las cuales se describen como nuevas: Apatema confluellum Falck & Karsholt, sp. n. (España: Tenerife), Apatema minimum Falck & Karsholt, sp. n. (España: Lanzarote y Fuerteventura), Apatema lapalmae Falck & Karsholt, sp. n. (España: La Palma), Apatema sallyae Falck & Karsholt, sp. n. (España: Gran Canaria), Apatema lanzarotae Falck & Karsholt, sp. n. (España: Lanzarote), Apatema mixtum Falck & Karsholt, sp. n. (España: Tenerife), Apatema pseudolucidum Falck & Karsholt, sp. n. (España: Gran Canaria), Apatema griseum Falck & Karsholt, sp. n. (España: Gran Canaria), Apatema grancanariae Falck & Karsholt, sp. n. (España: Gran Canaria), Apatema brunneum Falck & Karsholt, sp. n. (España: Tenerife), Apatema transversum Falck & Karsholt, sp. n. (España: Gran Canaria), Apatema stadeli Falck & Karsholt, sp. n. (España: Tenerife) and Apatema skulei Falck & Karsholt, sp. n. (España: Gran Canaria). El monotípico Dysallomima Gozmány, 2008, syn. n. es tratado como una nueva sinonimia de Apatema y la especie tipo Lampros coarctella Rebel, 1896 es transferida como Apatema coarctella (Rebel, 1896) comb. n. Apatema helleri (Rebel, 1910), comb. n. es transferida desde los Blastobasidae. A. mediopallidum Walsingham, 1900 es removida de la lista de Lepidoptera de las Islas Canarias. Morfológicamente se describen y figuran en detalle, todas las especies. Todas las especies ocurren en las Islas Canarias y solo una, A. fasciata (Stainton, 1859) ocurre ambas en estas islas y en el Archipiélago de Madeira. Los registros de A. mediopallidum Walsingham, 1900 de las Islas Canarias están basados en malas identificaciones. El análisis del AND código de barras soporta la identificación, separadamente, de cada una de las especies que aparecen monofiléticamente, bien soportadas y genéticamente aisladas en el análisis. La sola excepción es A. coarctella, la cual no puede ser tratada como monofilética con respecto a una sola muestra de un espécimen de A. mediopallidum, cuando las secuencias de los Apatema no canarios, fueron incluidos en el análisis. Argumentamos que este resultado es un artefacto.

Palabras clave: Lepidoptera, Autostichidae, Apatema, nuevas especies, ADN código de barras, Islas Canarias, España, Madeira, Portugal.

Introduction

Oegoconiinae are a small group of Gelechioidea, most of which are distributed in the Western Palearctic region. They were included by GOZMÁNY (2008) in his revision of the Palearctic Symmocidae. According to recent molecular studies, e. g. HEIKKILÄ et al. (2013) Oegoconiinae should be regarded as a subfamily of Autostichidae.

WALSINGHAM (1900: 220) erected the genus Apatema for a new species, A. mediopallidum, which he described from France: Corse. In a treatment of Microlepidoptera of the Canary Islands he subsequently transferred Gelechia fasciata Stainton, 1859 and Lampros coarctella Rebel, 1896 (as a synonym of fasciata) to Apatema and described a new species, A. lucidum (Walsinghm, 1908: 945). During most of the 20th century most specimens from outside of the Macaronesia Islands were referred to A. mediopallidum. Apatema species are rather small Gelechioidea having blackish forewings with whitish or yellowish markings, their genitalia are rather simple and similar between the species, and it has only slowly become evident that the genus is more diverse that hitherto suspected.

The Lepidoptera of the Canary Islands are considered well-known. Similar to other oceanic islands the fauna of the Canary Islands is less diverse than that of the adjacent continental countries (Morocco, Portugal), but includes a number of endemic species. Recent field work by the first author (e. g. FALCK et al., 2019; FALCK & KARSHOLT, 2019) has revealed a number of hitherto unrecorded or even undescribed species of Lepidoptera occurring in these islands. In the present paper we revise the genus Apatema in the Canary Island and the Madeiran Archipelago.

Many Canarian Apatema species are quite easily identified based on adult habitus (e. g. A. sallyae Falck & Karsholt, sp. n. and A. lucidum Walsingham), while others are difficult to identify without genitalia dissections (e. g. A. mixtum Falck & Karsholt, sp. n. and A. fasciata (Stainton)). When dissecting the male genitalia it is very important to separate the cornuti groups (Fig. 28) to ensure correct identification. In the female genitalia (Fig. 48) some species (e. g. A. stadeli Falck & Karsholt, sp. n.) have a slightly twisted ductus bursae. This gives the impression of a longitudinal fold of the anterior part of ductus bursae in the genital-slide, despite being an artefact, which we will refer to as “a longitudinal fold”.

Here we revise the Canarian Apatema species based on adult morphology and DNA barcodes (a 658 bp fragment of the mitochondrial COI gene). We provide detailed descriptions and diagnoses for all species and discuss their molecular diversity and potential phylogenetic relationships.

Material and methods

Most of the specimens were attracted to an 8 watt super actinic light. Some of the specimens especially from forest localities were obtained by netting in the afternoon sunshine and a few specimens were disturbed from the vegetation during daytime. Genitalia were dissected following Robinson (1976). Whole specimens were photographed with a Canon EOS 700D camera equipped with a Canon EF 100 mm objective. The genitalia slides were photographed using a Soptop CX40T Trinocular microscope in conjunction with a Toup Tek P10500A-E3 / E3ISPM05000KPA-E3 / 5.0MP USB3 camera.

The so-called DNA barcode fragment (a 658 bp fragment of the mitochondrial COI gene) was sequenced from all known Apatema species from the Canary Islands to detect genetically distinct clusters, and to obtain molecular data for new species. DNA samples (dried legs) were prepared according to the accepted standards and were processed at the Canadian Centre for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph). Genetic clusters are presented with their barcode index number (BIN; cf. RATMNASINGHAM & HERBERT, 2013).

To assess overall DNA diversity and relationships within the Canary Island Apatema we analysed the DNA barcodes under a Bayesian framework in MrBayes 3.2. (RONQUIST et al., 2011). The analysis was done utilising the model-jumping option (nst=mixed) allowing MrBayes to assess the best evolutionary model for the dataset. The rates were set to GAMMA, all other priors were left as default. The analysis was run for 10 million generations with sampling every 1000 generation, and the first 25% were used as burnin. The run files were subsequently assessed in Tracer 1.7.1 (part of the BEAST package, BOUCKAERT etal., 2019) to confirm that the two parallel analyses had converged and reached stationarity. The majority rule consensus tree was visualised in FigTree 1.4.4. (RAMBAUT, 2014). We analysed the dataset in BEAST in conjunction with BEAUti (BOUCKAERT et al., 2019) under the GTR model for molecular evolution (the model found by MrBayes) with rates set to GAMMA and all other priors left as default. The analysis was run for 10 million generations with sampling every 5000 generations. The resultant trees were then imported into DensiTree (BOUCKAERT & HELED, 2014), with the first 25% used as burnin, to visualise potential conflict areas in the data. To test whether the Apatema species found on the Canary Island and Madeira potentially comprise a monophyletic group, we augmented our DNA barcode dataset with publicly available barcode sequences from four additional species of Apatema (16 sequences) and three species of the closely related genus Oegoconia (six sequences). The new dataset was analysed under a Bayesian framework in MrBayes 3.2. utilising the model-jumping option with rates set to GAMMA. The analysis was run for 20 million generations with sampling every 1000 generations, and the first 25% were used as burnin. The run files were subsequently assessed in Tracer 1.7.1, and the majority rule consensus tree was visualised in FigTree 1.4.4. In all Bayesian analyses a single specimen of each of the three species Epanastatis sophroniella (Rebel, 1894), E. canariensis (Rebel, 1906) and Ambloma klimeschi Gozmány, 1975 were used as outgroups. All specimens used in the molecular analyses are listed in Table 1. We explored the general genetic diversity in Apatema from the Canary Island by calculating uncorrected p distances in Mega X (KUMAR et al., 2018). We calculated the average uncorrected p distance within as well as between each putative species (as listed in Table 1 and the Taxonomic section); the values are listed in Table 2. We further divided A. coarctella into populations from Fuerteventura, Gran Canaria, Lanzarote, La Palma and Tenerife and separately calculated the average uncorrected p distance within and between the four populations; the values are listed in Table 3. Similarly, we divided A. fasciata into populations from Gran Canaria, Fuerteventura and Lanzarote and separately calculated the average uncorrected p distance within and between the three populations; the values are listed in Table 4.

Table 1.
Species affiliation, voucher and BOLD ID, and collection localities for all species included in the molecular analyses as described in the text.
Species affiliation, voucher and BOLD ID, and collection localities for all species included in the molecular analyses as described in the text.

Table 1.
Species affiliation, voucher and BOLD ID (Cont...)
Species affiliation, voucher and BOLD ID (Cont...)

Table 1.
Species affiliation, voucher and BOLD ID (Cont...)