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Research Article
Monopis jussii, a new species (Lepidoptera, Tineidae) inhabiting nests of the Boreal owl (Aegolius funereus)
expand article infoMarko Mutanen, Peter Huemer§, Jonna Autto|, Ole Karsholt, Lauri Kaila#
‡ University of Oulu, Oulu, Finland
§ Tiroler Landesmuseen-Betriebsgesellschaft m.b.H., Innsbruck, Austria
| Unaffiliated, Rovaniemi, Finland
¶ Natural History Museum of Denmark, Copenhagen, Denmark
# University of Helsinki, Helsinki, Finland
Open Access

Abstract

Monopis jussii Kaila, Mutanen, Huemer, Karsholt & Autto, sp. nov. (Lepidoptera, Tineidae) is described as a new species. It is closely related to the widespread and common M. laevigella ([Denis & Schiffermüller], 1775), but differs in its distinct COI DNA barcode sequences, four examined nuclear loci as well as details in forewing coloration and pattern. Most reared specimens of M. jussii have emerged from the nest remnants of the Boreal owl (Aegolius funereus (Linnaeus, 1758)), but also nests of the Ural owl (Strix uralensis Pallas, 1771) and the Great tit (Parus major Linnaeus, 1758) have been observed as suitable habitats. Based on the present knowledge, the new species has a boreo-montane distribution as it is recorded only from northern Europe and the Alps. Several extensive rearing experiments from Strix spp. nest remnants from southern Finland did not produce any M. jussii, but thousands of M. laevigella, suggesting that the species is lacking in the area or, more unlikely, that the nest of these owl species do not serve as good habitat for the new species. This unexpected species discovery highlights, once again, the usefulness of DNA barcoding in revealing the cryptic layers of biodiversity. To serve stability we select a neotype for Tinea laevigella [Denis & Schiffermüller], 1775, and discuss the complicated synonymy and nomenclature of this species.

Keywords

boreo-montane, cryptic diversity, DNA barcoding, nuclear marker

Introduction

The lepidopteran fauna of Central and North Europe has been investigated for a longer time and more intensively than that of any other region in the world. Consequently, discoveries of species new to the region are nowadays uncommon and usually involve expansive or invasive species. Large-scale efforts to build taxonomically comprehensive regional DNA barcode reference libraries have, however, resulted in a boost in discoveries of overlooked species during the last 15 years, as demonstrated by the increase of new species descriptions e.g. in the family Gelechiidae by Huemer et al. (2020). Characteristic to the new discoveries is that they often concern unexpected cases of cryptic diversity among well-known and often widespread species. Examples of such recent findings, originally detected as deep intraspecific splits in DNA barcode sequences, include Leptidea reali Reissinger, 1990 (Dinca et al. 2011), Olethreutes subtilana (Falkovitsh, 1959) (Segerer et al. 2010), Phalonidia udana (Guenée, 1845) (Mutanen et al. 2012a), Epinotia cinereana (Haworth, 1811) (Mutanen et al. 2012b), Nemophora scopolii Kozlov, Mutanen, Lee & Huemer, 2016 (Kozlov et al. 2017), several Elachista spp. (Mutanen et al. 2013) and Hoplodrina alsinides (Costantini, 1922) (Huemer et al. 2020).

There are many more additional cases of potential cryptic diversity in European Lepidoptera, as dozens of species show high levels of genetic polymorphism in their mitochondrial DNA (Mutanen et al. 2016, Huemer et al. 2020). While polymorphism in the mitochondrial DNA may result from multiple other phenomena, including mitochondrial introgression and retained ancestral polymorphism, many of those cases are likely to result from cryptic diversity.

An intraspecific split of the mitochondrial DNA being reflected in the nuclear genome in sexually reproducing species and in sympatry would strongly suggest the presence of cryptic diversity, because, unlike mitochondrial DNA, nuclear DNA is subject to genetic recombination. From this starting point, we sequenced four nuclear markers of Monopis laevigella ([Denis & Schiffermüller], 1775), a widespread and common species of tineid moths, showing a deep sympatric genetic split in its DNA barcode region in Europe (Gaedike 2019). Despite the limited number of analyzed specimens, the results provided unequivocal genetic support for the presence of two biologically distinct species. Subsequent morphological examination revealed consistent differences in the adult wing patterns, providing additional support for the overlooked cryptic diversity. Additionally, based on the presently available data, the two species show overlapping, but different ranges and based on the present knowledge, also a different ecology. Based on these grounds, we here describe one of the taxa as new to science.

Material and methods

The material examined was acquired from the following collections:

ITJ Research collection of Juhani Itämies

MUT Research collection of Marko & Tomi Mutanen

MZH Finnish Museum of Natural History, Helsinki, Finland

TLMF Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria

ZMUO Zoological Museum, University of Oulu, Finland

ZSM Zoologische Staatssammlung München, Germany

Terminology of genitalia follows Robinson and Nielsen (1993) and Gaedike (2019).

Preparation of genitalia generally follows the method outlined by Robinson (1976). Male genitalia were mounted in dorso-ventral position as it was considered to best show shapes of diagnostic structures, even if the shape of the gnathos is not optimally expressed. Male genitalia were stained using Eosin, female genitalia as well as abdominal pelts of both sexes using Chlorazol black. Structures were embedded in Euparal. Images were edited using Corel PHOTO-PAINT (2019).

Species of Tineidae have been systematically sequenced for the standard barcode region of the mitochondrial COI (cytochrome c oxidase subunit 1) in the connection of ongoing regional or national DNA barcoding projects in the Alps (Lepidoptera of the Alps campaign) and Finland (FinBOL). DNA barcode sequencing was conducted at the Canadian Centre for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph) using standard Sanger protocols as explained in deWaard et al. (2008). We successfully sequenced 87 specimens of Monopis representing twelve species, the newly described species included. Five European species of Monopis (M. luteocostalis Gaedike, 2006, M. henderickxi Gaedike & Karsholt, 2001, M. christophi Petersen, 1957, M. pallidella Zagulajev, 1955 and M. barbarosi (Koçak, 1981)) were not included in this sampling. Each of them is morphologically clearly distinct from M. jussii sp. nov. (Gaedike 2019). Full collection and taxonomic data as well as voucher photographs, DNA sequences and GenBank accession numbers of all these specimens are available in the Barcode of Life Data Systems (BOLD; Ratnasingham and Hebert 2007) in the public dataset DS-MONOJUS at https://dx.doi.org/10.5883/DS-MONOJUS. Collection data of the specimens are also given in Table 1. Some of the COI sequences used in this study were previously published in Mutanen et al. (2016), the others are novel.

Table 1.

Summary of the collection data of barcoded specimens of Monopis used in this study. For more details, see the public BOLD dataset at https://dx.doi.org/10.5883/DS-MONOJUS.

Species Sample ID Sequence length Collector(s) Collection date Country Province Site Latitude / Longitude
Monopis burmanni TLMF Lep 18816 658 Huemer P. 13-Jun-2006 Austria Tyrol Nordtirol, Kranebitter Innaue 47.265, 11.323
Monopis burmanni TLMF Lep 18234 658 Huemer P. 05-Jun-2015 Austria Tyrol Nordtirol, Ellbachtal, unterer Kaiserboden 47.539, 11.926
Monopis crocicapitella TLMF Lep 06512 658 O. Rist 23-Sep-2005 Austria Vienna Wien Stadlau 48.217, 16.467
Monopis crocicapitella TLMF Lep 03882 658 Huemer P. 21-May-2004 Spain Comunidad Valenciana Valencia, El Saler, Albufera 39.3255 -0.312972
Monopis fenestratella MM18616 658 Marko Mutanen 1997 Finland N Mäntsälä 60.688, 25.168
Monopis fenestratella MM18615 658 Marko Mutanen 1997 Finland N Mäntsälä 60.688, 25.168
Monopis fenestratella MM08511 658 Marko Mutanen larva 1997-1998 Finland Ta Pälkäne
Monopis fenestratella MM08510 552 Marko Mutanen larva 1997-1998 Finland Ta Pälkäne
Monopis imella TLMF Lep 19836 658 Buchner P. 29-Aug-2014 Austria Niederoesterreich, Sollenau 47.905, 16.266
Monopis imella TLMF Lep 25734 639 Huemer P. 07-Sep-2016 Austria Burgenland, Jois SW, Hackelsberg 47.9539, 16.7747
Monopis imella TLMF Lep 25735 638 Huemer P. 07-Sep-2016 Austria Burgenland, Jois SW, Hackelsberg 47.9539, 16.7747
Monopis imella TLMF Lep 23122 658 Huemer P. 26-May-2017 Austria Burgenland, Hackelsberg 47.9528, 16.7733
Monopis imella TLMF Lep 19838 658 Buchner P. 17-Aug-2014 Austria Niederoesterreich, Sollenau 47.905, 16.266
Monopis imella MM18899 658 Kari Vaalamo, Bo Wikström 13-Jul-2002-19-Jul-2002 Finland Al Kökar 59.9031, 20.74
Monopis imella MM18898 658 Pekka Sundell, M. Varesvuo, L. Jalonen, Kalle Lundsten 25-Aug-2004-10-Sep-2004 Finland Al Kökar 59.92, 20.898
Monopis imella MM26020 658 Huotari, Laasonen 08-Jul-2014 Hungary Tokaj Tarcal 48.0512, 21.1811
Monopis imella MM26021 658 Huotari, Laasonen 08-Jul-2014 Hungary Tokaj, Tarcal 48.0512, 21.1811
Monopis jussii MM17525 658 Marko Mutanen 2001 Finland Oba Ylikiiminki 64.984, 26.153
Monopis jussii MM18626 658 Panu Välimäki & Marko Mutanen 2006 Finland Oba Oulu 64.9768, 25.3056
Monopis jussii MM15526 658 Marko Mutanen larva 2001 Finland Oba Ylikiiminki
Monopis jussii TLMF Lep 09795 658 Huemer P. 23-Jun-2006 Italy South Tyrol Suedtirol, Tiers E, Plafetscher Wald 46.472, 11.596
Monopis laevigella TLMF Lep 09306 658 Huemer P. 19-Jun-2012 Austria Tyrol Nordtirol, Oberpettnau, Platten 47.301, 11.126
Monopis laevigella TLMF Lep 10365 658 Huemer P. 16-Jun-2013 Austria Tyrol Nordtirol, Tiefenbachklamm/ Brandenberg 47.484, 11.864
Monopis laevigella TLMF Lep 10441 658 Huemer P. 16-May-2013 Austria Tyrol Nordtirol, Tiefenbachklamm/ Brandenberg 47.484, 11.864
Monopis laevigella TLMF Lep 07389 658 Huemer P. 25-May-2008 Austria Tyrol Nordtirol, Telfs/ Moritzen SW, Innau 47.299, 11.05
Monopis laevigella TLMF Lep 10354 658 Huemer P. 16-Jun-2013 Austria Tyrol Nordtirol, Tiefenbachklamm/ Brandenberg 47.484, 11.864
Monopis laevigella TLMF Lep 07970 658 Huemer P. 25-May-2012 Austria Vorarlberg Umg.Zwischenwasser, Ueble Schlucht, Eingang 47.267, 9.667
Monopis laevigella MM19355 658 O. Martin larva 14-Oct-2004 Denmark Sjaelland Nez, Bognaes, Egehoved
Monopis laevigella MM17303 658 Tomi Mutanen 09-Jun-2010 Finland Ab Salo 60.335, 23.088
Monopis laevigella MM17522 658 Henrik Bruun 01-Apr-2007 Finland Ab Nauvo 60.225, 21.945
Monopis laevigella MM21029 658 Ali Karhu 27-Jun-2008-29-Jun-2008 Finland Ka Liperi 62.552, 29.167
Monopis laevigella MM21028 658 Ali Karhu 1-Jun-2010-25-Jul-2010 Finland Ka Liperi 62.551, 29.226
Monopis laevigella MM21026 658 Ali Karhu 03-Jul-2007 Finland Ka Liperi 62.563, 29.013
Monopis laevigella MM21025 658 Ali Karhu 2005 Finland Ka Liperi 62.511, 29.475
Monopis laevigella MM17524 606 Marko Mutanen 30-Jun-1997 Finland Oba Hailuoto 64.968, 24.671
Monopis laevigella MM15527 658 Marko Mutanen 30-Jun-2001 Finland Oba Oulu 64.977, 25.306
Monopis laevigella MM10119 658 Marko Mutanen, Nestori Mutanen, Anttoni Mutanen 12-Jul-2008 Finland Oba Kiiminki 65.071, 25.725
Monopis laevigella MM18625 658 Panu Välimäki 21-Jun-2000 Finland St Luvia 61.29, 21.587
Monopis laevigella MM17526 658 Juhani Itaemies 14-Feb-2005 Finland St Eurajoki 61.193, 21.417
Monopis laevigella TLMF Lep 27537 658 Huemer P. 29-Jun-2019 Italy Piedmont Fenestrelle, ca. 0,7 km NE Pequerel 45.0517, 7.07111
Monopis laevigella TLMF Lep 12113 658 Huemer P. 17-Jul-2013 Italy South Tyrol Suedtirol, N Zwischenwasser/ St. Lorenzen 46.739, 11.873
Monopis laevigella TLMF Lep 11818 658 Huemer P. 25-Jul-2013 Italy South Tyrol Suedtirol, Franzenshoehe / Stilfserjoch 46.534, 10.486
Monopis laevigella TLMF Lep 02066 658 Huemer P. 01-Jul-2010 Italy South Tyrol Suedtirol, Ritten/ Obergruenwald 46.597, 11.439
Monopis laevigella TLMF Lep 05368 658 Huemer P., Tarmann G. M. 01-Aug-2011 Macedonia Mavrovo NP, Radika valley, around bridge, 10 km NNW Sveta Voda 41.789, 20.547
Monopis monachella TLMF Lep 08436 658 Huemer P. 25-Jul-2012 Austria Vorarlberg Lustenau, Schweizer Ried, AZE Haeusle S 47.446, 9.69
Monopis monachella TLMF Lep 19839 658 Buchner P. 07-Jun-2014 Austria Niederoesterreich, Sollenau 47.905, 16.266
Monopis monachella MM13366 658 Marko Mutanen, Panu Välimäki 2008 Finland Ab Dragsfjärd 60.011, 22.498
Monopis monachella MM11934 658 Marko Mutanen, Panu Välimäki 2007 Finland N Hanko 59.836, 23.236
Monopis monachella MM17249 658 Lauri Kaila 21-Aug-2005 Finland N Tammisaari 59.829, 23.612
Monopis monachella MM12377 658 Marko Mutanen, Panu Välimäki 2007 Finland Sa Imatra 61.108, 28.799
Monopis neglecta TLMF Lep 07250 658 Sumpich J. 10-Jun-2010 Austria Lower Austria Hardegg Umgebung/ Thaya Haenge 48.854, 15.858
Monopis neglecta TLMF Lep 17583 658 Deutsch H. 30-Aug-2002 Austria Tyrol Osttirol, Lengberg 46.801, 12.891
Monopis neglecta TLMF Lep 06608 658 Rist O. 11-Jun-2010 Austria Vienna Wien Mauer 48.15, 16.25
Monopis nigricantella TLMF Lep 03881 658 Huemer P. 07-Sep-2005 Spain Comunidad Valenciana Valencia, El Saler, Albufera 39.3255, -0.312972
Monopis nigricantella TLMF Lep 03879 658 Huemer P. 18-May-2004 Spain Comunidad Valenciana Valencia, El Saler, Albufera 39.3255, -0.312972
Monopis nigricantella TLMF Lep 03878 658 Huemer P. 22-May-2004 Spain Comunidad Valenciana Valencia, Santa Pola, Playa del Pinet 38.1583, -0.625278
Monopis nigricantella TLMF Lep 03880 658 Huemer P. 08-Sep-2005 Spain Comunidad Valenciana Valencia, El Saler, Albufera 39.3255, -0.312972
Monopis obviella TLMF Lep 15096 636 Huemer P. 19-Jun-2014 Austria Tyrol Nordtirol, Baumkirchen W 47.296, 11.552
Monopis obviella TLMF Lep 09367 658 Huemer P. 02-Jun-2012 Austria Tyrol Nordtirol, Flaurling NW, Innau 47.302, 11.121
Monopis obviella TLMF Lep 08054 658 Huemer P. 15-Jun-2012 Austria Vorarlberg Bludesch, Bludescher Magerrasen E, Umg. Jordan 47.203, 9.747
Monopis obviella TLMF Lep 09962 658 19-Jun-13 Austria Vorarlberg Umg.Langenegg, Langenegg-Leiten, Fohren 47.467, 9.883
Monopis obviella TLMF Lep 25739 658 Huemer P. 07-Sep-2016 Austria Burgenland, Jois SW, Hackelsberg 47.9539, 16.7747
Monopis obviella TLMF Lep 19832 658 Buchner P. 29-Aug-2014 Austria Niederoesterreich, Sollenau 47.905, 16.266
Monopis obviella MM18928 658 Kari Vaalamo, Bo Wikström 19-Jul-2008-23-Jul-2008 Finland Al Lemland 59.9564, 20.0116
Monopis obviella MM06790 658 Marko Mutanen 13-Jul-2007 Finland Al Lemland 60.026, 19.961
Monopis obviella MM21130 658 Marko Mutanen, Tomi Mutanen, Anttoni Mutanen, Nestori Mutanen 16-Jul-2011 Finland N Hanko 59.834, 23.013
Monopis obviella TLMF Lep 27604 658 Huemer P. 28-Jun-2019 Italy Piedmont Fenestrelle, ca. 1 km WNW Pequerel 45.0497, 7.05139
Monopis obviella TLMF Lep 27794 630 Huemer P. 23-Jul-2019 Italy Piedmont Fenestrelle, ca. 0,7 km NE Pequerel 45.0517, 7.07111
Monopis obviella TLMF Lep 10292 658 Huemer P. 25-Jun-2013 Italy South Tyrol Suedtirol, Margreid/ Fennerschlucht 46.288, 11.201
Monopis obviella TLMF Lep 02169 658 Huemer P. 04-Jun-2010 Italy South Tyrol Suedtirol, Montiggl/ Kleiner Priol 46.428 11.03
Monopis obviella TLMF Lep 12282 658 Huemer P. 05-Jul-2013 Italy South Tyrol Suedtirol, Schleiser Leiten 46.698, 10.517
Monopis spilotella MM04157 658 Marko Mutanen Finland Le Enontekiö 68.997, 20.744
Monopis spilotella MM24137 658 Marko Mutanen, Anttoni Mutanen, Nestori Mutanen 05-Jul-2014 Finland Lkoc Muonio 67.9178, 23.7466
Monopis spilotella MM03158 658 Marko Mutanen 2006 Finland Oba Kiiminki 65.071, 25.725
Monopis spilotella MM02304 658 Marko Mutanen, Panu Välimäki 2006 Finland Sa Imatra 61.108, 28.799
Monopis weaverella TLMF Lep 15166 658 Huemer P. 09-Jun-2014 Austria Tyrol Nordtirol, Ellbachtal, unterer Kaiserboden 47.539, 11.926
Monopis weaverella TLMF Lep 15178 658 Huemer P. 09-Jun-2014 Austria Tyrol Nordtirol, Ellbachtal, unterer Kaiserboden 47.539, 11.926
Monopis weaverella TLMF Lep 18561 658 Huemer P. 20-Jul-2005 Austria Tyrol Nordtirol, Umg. Innsbruck, Samertal, Jaegerkar 47.34, 11.382
Monopis weaverella TLMF Lep 07388 658 Huemer P. 25-May-2008 Austria Tyrol Nordtirol, Telfs/ Moritzen SW, Innau 47.299, 11.05
Monopis weaverella TLMF Lep 09220 658 Huemer P. 06-Jun-2010 Austria Tyrol Nordtirol, Walchsee/ Schwemm N 47.661, 12.298
Monopis weaverella MM21138 658 Marko Mutanen ,Tomi Mutanen 18-Jun-2011 Finland Ab Nauvo 60.192, 21.923
Monopis weaverella MM13581 658 Marko Mutanen, Panu Välimäki 2008 Finland Ab Dragsfjärd 60.011, 22.498
Monopis weaverella MM21027 658 Ali Karhu 21-Jun-2004-23-Jun-2004 Finland Ka Liperi 62.552, 29.167
Monopis weaverella MM04159 658 Marko Mutanen Finland Le Enontekiö 68.997, 20.744
Monopis weaverella MM04158 658 Marko Mutanen Finland Le Enontekiö 68.997, 20.744
Monopis weaverella MM02600 639 Marko Mutanen, Panu Välimäki 2006 Finland Sa Imatra 61.108, 28.799
Monopis weaverella TLMF Lep 22008 658 Schaefer W. 07-Aug-2015 Germany Kefenrod 50.35, 9.21667

Four nuclear genes, carbamoylphosphate synthase domain protein (CAD), elongation factor 1 alpha (EF-1a), cytosolic malate dehydrogenase (MDH) and wingless, were sequenced at the University of Oulu, Finland. These genes were chosen primarily based on the high amplification success rate in other Tineidae, but also based on our previous experience on their general good functionality to provide useful taxonomic information between closely related species. In these analyses, three specimens of M. laevigella and two specimens of M. jussii, all collected from Finland, were included. Legs of dry and pinned adult specimens were used for extraction of genomic DNA with DNeasy Blood & Tissue Kit (Qiagen). We largely followed the sequencing protocol by Wahlberg and Wheat (2008), but PCR clean-up was carried out with ExoSAP-IT (Affymetrix, Santa Clara, CA, USA) and Sephadex columns (Sigma-Aldrich, St. Louis, MO, USA). Additionally, sequencing was performed using an ABI 3730 DNA Analyzer (Applied Biosystems, Foster City, CA, USA). Sequences were checked and edited using BioEdit software (Hall 1999). The sequences were uploaded to a VoSeq database (Peña and Malm 2012). The same dataset was used to generate fasta files for Neighbor-Joining analyses.

Minimum genetic p-distance barcode divergence between M. laevigella and M. jussii was calculated using analytical tools in BOLD Systems v. 4.0 (http://www.boldsystems.org). Neighbor-joining trees for the barcode region for all included Monopis species and specimens as well as four nuclear genes for five analyzed specimens of M. laevigella and M. jussii were constructed under p-distance model using Mega 7.0 (Kumar et al. 2016). The trees were stylized using CorelDraw v. 20.0.0.633.

Results

DNA sequencing resulted in a barcode of 552 bp or longer for 81 specimens. All except seven specimens yielded a full-length (654 bp) barcode. BOLD’s barcode gap analysis showed that all included species have highly species-specific DNA barcodes with the mean of minimum divergences (p-distance model) to the nearest species being 10.01% (range 4.43–17.58%) (Figure 1). The minimum divergence between M. laevigella and M. jussii is 4.43%.

Figure 1. 

A compressed Neighbor-Joining tree DNA barcode region of European Monopis with most European species represented. The depth of the triangle is proportional to the intraspecific genetic variability within species and the height to sampling intensity.

For each nuclear gene, data of only a single specimen of two analyzed M. jussii specimens were retrieved. Informative (i.e. data from both species available) sequence lengths by genes were as follows: CAD: 336 bp, EF-1a: 410 bp, MDH: 334 bp, wingless: 307 bp. Genetic p-distances between the two species were: CAD: 2.1%, EF-1a: 2.2%, MDH: 1.5%, and wingless: 4.1%. As a rule, the specimen of M. jussii formed a sister to the two or three specimens of M. laevigella (Figure 2).

Figure 2. 

Comparison of genetic variability in four nuclear genes, CAD, EF-1a, MDH and wingless, between Monopis laevigella and M. jussii sp. nov.

Monopis jussii Kaila, Mutanen, Huemer, Karsholt & Autto, sp. nov.

Figures 3, 4, 5, 6, 7, 8, 9

Type material

Holotype ♂ (Figure 3): FINLAND, PPe Yli-Kiiminki, larva 2001, ex nest of Aegolius funereus, M. Mutanen leg. R. Gaedike prep. 8607. (ZMUO).

Figure 3. 

The holotype male of Monopis jussii sp. nov. PPe Yli-Kiiminki, larva 2001, ex nest of Aegolius funereus, M. Mutanen leg., R. Gaedike prep. 8607. (Coll. ZMUO).

Figure 4. 

Comparison of habitus between Monopis laevigella and M. jussii sp. nov. A–C M. laevigella female D–F M. laevigella male G–I M. jussii paratype, females J–L M. jussii paratype, males.

Paratypes. FINLAND • 7 ♂ 16 ♀, PPs Kiiminki, 65.1163°N, 25.8291°E, Larva 1995, ex nest of Aegolius funereus, L. Kaila prep. 6317, 6325, 6326, M. Mutanen leg. (ZMUO); Finland: 10 ♂, 16 ♀, PPe Yli-Kiiminki, larva 2001, ex nest of Aegolius funereus, L. Kaila prep. 6314, 6315, 6316, 6322, 6323, 6324, R. Gaedike prep. 8606, 8607, 8698, DNA samples MM15526, MM17525, M. Mutanen leg. (ZMUO); • 2 ♀, Oba Utajärvi, Pälli, 64.8363°N, 26.21°E, larva 1980 ex nest of Aegolius funereus, J. Itämies leg. (ITJ); • 3 ♂ 3 ♀, Kn Puolanka, Piltunkijärvi, 64.7618°N, 27.3151°E, larva 18.6.1976 ex nest of Aegolius funereus (1974), M. Rikkonen leg. (ZMUO); • 2 ♂, Kn Vaala, Otermajärvi, 64.6724°N, 27.1047°E, larva 12 Jun 1976 ex nest of Aegolius funereus (1974), M. Rikkonen leg. (ZMUO); • 1 ♀, Kn Kajaani, 64.2263°N, 27.7932°E, VYÖ 1210 ad luc 15. –21 Jun 2006, DNA sample MM 17523, R. Leinonen leg. (ZMUO). ITALY • 1 ♀, Südtirol, Tiers E, Plafetscher Wald, 1600–1650 m, 46.472°N, 11.596°E, 23 Jun 2006, leg. Huemer, DNA sample TLMF Lep 09795 (TLMF).

Other material

FINLAND • 7 ♂ 4 ♀, Ta Valkeakoski, Sääksmäki, 61.2326°N, 24.1137°E, ex larva (host unknown); 1992, S. Karhula leg. (MZH); • 2 ♀, Kn Kajaani, Karankalahti, 64.2222°N, 27.721°E, ex larva 2016 from nest of Strix uralensis, Itämies & Kyrki leg. (ZMUO); 1 ♀, PPe: Oulu, Oinaansuo, 65.0249°N, 25.6209°E, larva 28 Apr 1992 in nest of Parus major, J. Itämies leg. (ZMUO); • 1 ♀, EP Jurva, 62.7002°N, 22.0153°E, ex larva 2006, H. Vuorinen leg. (ZMUO); 2 ♀, Ks Kuusamo, 66.2565°N, 29.2807°E, ex larva 1975, J. Viramo leg. (ZMUO); • 1 ♂ 1 ♀, Ks Salla, Värriö, R1 & R3, 30 Jun 1989 & 21 Jul 1987, Erkki Pulliainen leg. (ZMUO); • 1 ♀, Li Inari, Kivijoki, 68.6125°N, 28.3509°E, 15 Jul 1993, E. & L. Laasonen leg. (ZMUO); • 1 ♂, Ks Kuusamo, Autiotalo, 66.3591°N, 29.6029°E, 28 Jun 1995, E. & L. Laasonen leg. (ZMUO); • 1 ♂, PPn Rovaniemi, 66.5509°N, 25.7619°E, 17 Jun 1992, T. Mutanen leg. (ZMUO); • 1 ♀, EnL Enontekiö, Saana, 69.0456°N, 20.8554°E, 11 Jul 2016, Marko, Nestori & Anttoni Mutanen leg. (ZMUO); • 1 ♀, Pedersöre, 8 Jul 1939, Sjöholm leg. (ZMUO); • 1 ♀, Om Jakobstad, 63.7098°N, 22.6489°E, 21 Jun 1936, E. Sjöholm leg. (ZMUO); 2 ♂, KP Haapajärvi, Harjunniemi, 63.7434°N, 25.3292°E, ad luc. 3 Jul 1975 & 6 Jul 1975, A. Kosonen leg. (ZMUO); NORWAY • Finnmark Alta, Mattisfossen-Sakkopadne, 5 Jul 1973, J. Kyrki leg. (ZMUO); SWEDEN • Härjedalen, Vemdalen, 3 Jul 1947, Henrik Bruun leg. (ZMUO).

Diagnosis

Monopis jussii sp. nov. is externally close to M. laevigella, but the forewing appears darker, as it is less mottled with pale scales, especially along the margins (Figures 4, 5). Fringes are yellow and with a clear fringe line in M. laevigella but grey and without the fringe line in M. jussii. Besides the genetic markers, the forewing colour is indeed the best clue to separate these species. There is nevertheless some variation, especially in M. laevigella. Both male and female genitalia vary considerably, as do those of M. laevigella. The variation in all characters of genitalia overlaps between these species, and, apparently, they cannot be identified by genital characters. For variation of M. laevigella see also Gaedike (2019). Moreover, M. weaverella (Scott, 1858) and M. neglecta Šumpich & Liška, 2011 may occasionally fall within the morphological variation of these two species, especially in females. The males of M. weaverella and M. neglecta can however be distinguished from M. laevigella and M. jussii by the shape of gnathos, best decipherable in lateral view (see Gaedike 2019): gnathos arms are straight, triangular in M. weaverella and M. neglecta, angled particularly in anterior margin in M. laevigella and M. jussii.

Figure 5. 

Comparison of forewing patterns of Monopis laevigella (A) and M. jussii sp. nov. (B). The arrows indicate differences in fringe colour (yellow/grey), fringe line (present/absent; chequered/non-chequered) and forewing costa (many white scales between the costa and the dorsal spot/few white scales between the costa and the dorsal spot).

Description

Forewing length 5.8–8.5 mm (n = 8 ♂ and 8 ♀) (note that the specimens are reared which may have affected their size). Maxillary palpus, labial palpus and head ochreous yellow; outer side of labial palpus with dark grey scales, second segment distally bristled. Scape of antenna ochre with pecten formed of bristle-shaped scales, pedicel and flagellum dark brown. Thorax dark grey, dorsomedially variably intermixed or entirely with pale ochre scales; tegula dark grey, apically often paler grey or ochre. Fore and mid leg inwardly ochre, outwardly leaden grey, apex of tibia and tarsal segments ochre. Hind leg inwardly pale, outwardly ochre, intermixed with grey scales; spurs and apex of tibia and tarsal articles ochre. Forewing dark grey, variably mottled with pale grey scales; costa narrowly and variably sometimes ochre; basal scales of termen with alternating pale ochre and grey scales, distal scales of termen unicolorous grey, contrast between distally paler basal scales and darker distal scales giving an impression of faint fringe line; silvery grey spot somewhat basal of middle of wing length at fold. Hind wing bluish grey with somewhat darker grey veins; fringe basally narrowly ochre, otherwise grey. Underside of wings grey with ochre margin; underside of hindwing dark grey along costal margin. Abdomen leaden grey, basal segments ventrally more or less ochre.

Male genitalia (Figure 6). Uncus elongate, triangular, laterally with long, hair-like scales, distally pointed, bifid. Gnathos arms angled in the middle, tapered toward hook-shaped apex. Basal and distal margins of tegumen reinforced, U-shaped, anterior margin more deeply. Shape of valva highly variable, gradually varying from ovoid and basally broadest to somewhat elongate and medially widest; distally round. Every aspect of saccus variable; straight or somewhat undulate, apically little or very much widened; length also very variable. Phallus straight and nearly parallel-sided, slightly widened at basal 1/3; length compared to that of saccus impossible to establish due to variation in length of saccus. Phallus distally inserted in cylindrical, internally spinose anellus. Vesica distally densely spinose, devoid of cornuti.

Figure 6. 

Overview of male genitalia of Monopis jussii sp. nov. A paratype, Finland, Kiiminki, M. Mutanen leg., L. Kaila prep. 6317 B Finland, Yli-Kiiminki, M. Mutanen leg., L. Kaila prep. 6315.

Female genitalia (Figures 79). Papilla analis membranous, elongate, distally round, with a few setae. Apophysis posterioris as long as segments 7+8, posteriorly starting as continuation of papilla analis, slender, anteriorly slightly widened, apex cut. Apophysis anterioris 1/3 length of and slightly stouter than apophysis posterioris, twice as long as 8th segment, distally not widened. Ovipositor telescopic, with two retractile nodes; with a few stout setae. Ventral pseudapodemes (sensu Davis and Robinson 1999) not decipherable. Tergum 8 posteriorly somewhat sclerotized. Ostium a widely U-shaped opening, laterally bordered as posteriorly curved rim, laterad shallowly emarginated in posterior direction, emargination with a few long setae; devoid of microtrichia but minutely granulose. Length of antrum variable, narrowed toward colliculum; colliculum tubular, length variable, 2–4 times as long as wide, usually narrowed in the middle. Ductus bursae between colliculum and corpus bursae membranous, as long as apophysis anterioris. Corpus bursae oval, 3 times as long as wide; in approximately the middle to posterior 1/3 ca. 12 elongate, sharply spicular or dentate signa forming transverse band.

Figure 7. 

Overview of female genitalia of Monopis jussii sp. nov., paratype, Finland, Yli-Kiiminki, M. Mutanen leg., L. Kaila prep. 6324.

Figure 8. 

Details of ostium bursae and colliculum of female genitalia of Monopis jussii sp. nov. A paratype, Finland, Yli-Kiiminki, M. Mutanen leg., L. Kaila prep. 6324 B paratype, Finland, Kiiminki, M. Mutanen leg., L. Kaila prep. 6325 C paratype, Finland, Yli-Kiiminki, M. Mutanen leg., L. Kaila prep. 6322 D paratype, Finland, Kiiminki, M. Mutanen leg., L. Kaila prep. 6326.

Figure 9. 

Signa of corpus bursae of female genitalia of Monopis jussii sp. nov. A paratype, Finland, Yli-Kiiminki, M. Mutanen leg., L. Kaila prep. 6324 B paratype, Finland, Kiiminki, M. Mutanen leg., L. Kaila prep. 6325 C paratype, Finland, Yli-Kiiminki, M. Mutanen leg., L. Kaila prep. 6322 D paratype, Finland, Kiiminki, M. Mutanen leg., L. Kaila prep. 6326.

Genetic characterisation

Clearly distinguishable by its DNA barcode from all other species of Monopis barcoded globally so far (Figure 1). Genetically the closest species with a minimum divergence of 4.43% is M. laevigella. Intraspecific divergence among four barcoded specimens from Finland and Italy is 0.15%. Additionally, the species show 1.5–4.1% interspecific divergence in the nuclear genes of CAD, EF-1a, MDH and wingless (Figure 2).

Etymology

The species is dedicated to Dr Juhani (Jussi) Itämies, a Finnish expert of Lepidoptera who, as far as we know, is the first to have reared this species. He has also spent most of his life on faunistic research of Finnish Lepidoptera and has done incredible work in elucidating the life history of numerous microlepidopteran species.

Distribution

From our available observations M. jussii seems to have a boreo-montane distribution pattern. It is widely distributed in Finland and also recorded from Norway (Finnmark) and Sweden (Härjedalen). Records from the Alps seem rare with a proved, barcode-based locality in the Italian Dolomites and two further unpublished records (ZSM, A. Segerer) in the Bavarian Alps.

Biology

So far reared on five different occasions from the nest bottoms of the Boreal owl (Aegolius funereus). Two specimens in the collection of ZMUO have been reared from the nest of the Ural owl (Strix uralensis) and one specimen from the nest of the Great tit (Parus major). Additionally, three reared specimens of two different rearing events do not state anything about the origin. One specimen has been found in a vacated house. Thirteen specimens in coll. ZMUO and a specimen from the Italian Alps in coll. TLMF have been collected in the wild between 17 June to 21 July, which matches well with the flight time of other Monopis species of these regions.

Taxonomic remarks on Monopis laevigella

Monopis jussii sp. nov. is most closely related to M. laevigella and can easily be confused with that species (see above). We therefore re-evaluate available names in the M. laevigella species group.

Monopis laevigella ([Denis & Schiffermüller], 1775).

Tinea laevigella [Denis & Schiffermüller], 1775: 139.

Misidentifications

Tinea rusticella Hübner, 1796: 61, pl. 3, fig. 17; a junior synonym of Haplotinea insectella (Fabricius, 1794) (Zeller, 1852: 153–154).

Recurvaria rustica Haworth, 1828: 548; unjustified emendation of Tinea rusticella Hübner, 1796.

Tinea saturella Haworth, 1828: 562, unavailable.

Tinea vestianella sensu Stephens, 1835: 344; a misidentification of Phalaena (Tinea) vestianella Linnaeus, 1758.

Blabophanes rusticella ab. semispilotella Strand, 1900: 225; unavailable name, deemed infrasubspecific according to ICZN Art. 45.6.2 from use of the term “ab.”; a misidentification of M. weaverella (Scott, 1858) (Gaedike 2019).

Neotype selection

Tinea laevigella was described from an unspecified number of specimens collected in the area of Vienna, Austria ([Denis & Schiffermüller], 1775). The collection was later deposited in the “Hof-Naturalien-Kabinett” and destroyed by fire during the Vienna Rebellion on 31st of October 1848 (Speta 2003). Since this species can be confused with M. jussii sp. nov. and several other congeneric taxa we designate as neotype a male specimen from Austria to preserve stability (Figure 10). It is labelled “AUSTRIA occ. Nordtirol / Brandenberg / Tiefenbachklamm / 11°51'52"E, 47°29'4"N / 645 m, 16.6.2013 / leg. Huemer” “DNA Barcode / TLMF Lep 10354” (TLMF).

Figure 10. 

Neotype male of Monopis laevigella from Austria, here designated. AUSTRIA occ. Nordtirol / Brandenberg / Tiefenbachklamm / 11°51'52"E, 47°29'4"N / 645 m, 16.6.2013 / leg. Huemer” “DNA Barcode / TLMF Lep 10354”. (Coll. TLMF).

Tinea rusticella was figured twice by Hübner in the eighth volume of his Sammlung europäischer Schmetterlinge, first it was validly described on page 61, pl. 3, fig. 17 (1796) and later a different species was figured on pl. 49, fig. 339 (1813). Hübner (1825) considered them conspecific, and he referred to both figures when he erected the monotypic genus Monopis.

Zeller (1852) was probably the first to question whether Hübner’s two figures of Tinea rusticella represented the same species. He referred to Hübner’s fig. 339 (1813) when dealing with the species, which became known as Monopis rusticella [= Monopis laevigella ([Denis & Schiffermüller], 1775)], and rejected that Hübner’s fig. 17 (1796) could be of a specimen of that species, suggesting that it could be Tinea misella Zeller, 1839 [= Haplotinea insectella (Fabricius, 1794)]. Tinea rusticella Hübner, 1813 is both a misidentification and a homonym of Tinea rusticella Hübner, 1796 and thus permanently invalid.

Haworth (1828: 548) named the species twice. First with reference to Hübner’s pl. 3, fig. 17 as Recurvaria rustica, which is an unjustified emendation and thus an objective synonym of Tinea rusticella (Hübner, 1796) [= Haplotinea insectella (Fabricius)], and later in the same work Haworth (op. cit.: 339), again with reference to Hübner’s pl. 3, fig. 17, proposed the name Tinea saturella in synonymy with Tinea rusticella. Because Tinea saturella was described in synonymy with Tinea rusticella it was always considered a synonym of that species (viz. Monopis rusticella), but because Haworth referred only to Hübner’s fig. 17 (and not to fig. 339) it is an objective junior synonym of Tinea rusticella Hübner, 1796, and thereby a subjective junior synonym of Haplotinea insectella (Fabricius). However, as the name Tinea saturella has never been made available under the provision of Art. 11.6. of the Code (ICZN 1999) and adopted as the name of a taxon before 1961, we consider it as unavailable.

Although Monopis Hübner 1825 was described as a monotypic genus, it is based on a partly misidentified species. We consider Zeller (1852) as First Reviser of Tinea rusticella Hübner, restricting the name to the species now (and also by Zeller 1852) known as Monopis laevigella ([Denis & Schiffermüller], 1775).

Discussion

Compared with many other groups of Lepidoptera, the species diversity of Tineidae is generally poorly investigated. Hundreds of species deposited in museum collections remain undescribed (Robinson 2009). It is likely that many more species remain entirely undiscovered globally. The European fauna is comparatively well understood, and the fauna of the entire continent has recently been taxonomically reviewed in two monographs (Gaedike 2015, 2019). New species discoveries are uncommon, particularly for central and northern parts of Europe. An example of a recent species discovery is that of Monopis neglecta Šumpich & Liška, 2011, a species that morphologically is nearly indistinguishable from M. weaverella (Scott, 1858) (see Gaedike 2019). While no genetic data were provided for M. neglecta in the original description, the DNA barcode sequences provided in the present study confirm its status as a separate species from M. weaverella. It is encouraging that although the species of Tineidae are often difficult to tell apart from each other morphologically, no cases of barcode sharing in the European fauna are known. Evidently, therefore, DNA barcoding provides an efficient way to investigate their diversity in less thoroughly explored areas as well.

Based on the available distributional data, Monopis jussii has a much more limited range than M. laevigella. It is possible, if not likely, that it is a member of boreo-montane faunal elements, being distributed in the boreal region on the one hand and in the Alps below the timberline on the other hand. It is likely absent from the lowlands of Central Europe. It would not be surprising if the species turns out to be present in other European mountain systems and the eastern Palearctic. Based on the large number of examined museum specimens from the ZMUO and MZH collections, the species is widely present in northern Finland south to ca. 64° N but becomes much scarcer towards the more southern localities. The southernmost verified records from Finland are from the province of Tavastia australis (ca. 61° N).

Based on our own and other experiences (Robinson 2009, Gaedike 2019), Monopis laevigella is not strict regarding the source of its food, but it seems to prefer cavity-breeding birds, possibly because their nests are usually dry. Several extensive rearing experiments of nest bottoms of various birds, mostly the Tawny Owl (Strix aluco Linnaeus, 1758) and the Ural Owl (S. uralensis Pallas, 1771), from southern Finland have yielded large numbers of M. laevigella, which is usually present in every nest in large numbers. In an experiment by MM in 2017 with 13 nest bottoms of Strix spp., probably thousands of M. laevigella emerged. Among several dozen pinned specimens sampled from each nest, none represents M. jussii. Other species that are regularly or often present in the nests of Strix spp. in Finland are Niditinea striolella (Matsumura, 1931) (usually emerges in great numbers too), Tinea svenssoni Opheim, 1965 (present in almost all nests), Tinea steueri Petersen, 1966 (not present in every nest) and Monopis fenestratella (Heyden, 1863) (present in most nests but is cryptic in behaviour). While it is possible that M. jussii has stricter habitat requirements and that it has a strong preference for the Boreal Owl, we find this possibility unlikely. The Boreal owl, the Ural owl, as well as the Great tit are all cavity breeders, rendering the nest conditions between these species very similar. In rearing conditions, tineids are not selective for the origin of food and readily feed on mammal hairs too. It is more likely that Monopis jussii has been reared mostly from the nests of the Boreal owl just because it is a more common owl species within the moth’s main distribution in Finland than either of the two Strix species present in Finland. Further rearing experiments, optimally systematically from different species of birds, would bring additional valuable information on the habitat requirements of M. jussii and several other species of Tineidae.

Monopis laevigella has a Holarctic distribution (Landry and Pohl 2018, Gaedike 2019). Many specimens of this species have been barcoded from the Nearctic region, both from Canada and the U.S.A. They fall in two clusters, both of which are highly distinct from the clade consisting of M. jussii and the Palearctic M. laevigella (data only partially public in BOLD). In the Neighbor-Joining trees neither of these clusters is placed as sister to the Palearctic M. laevigella + M. jussii clade, suggesting that they represent distinct taxa and even that their closest relative is not M. laevigella. However, due to the limited phylogenetic information content of the DNA barcode region, verification of both scenarios requires more rigorous and thorough taxonomic and phylogenetic scrutiny.

Acknowledgements

We are grateful to Leif Aarvik, Reinhard Gaedike, Peter Buchner, Bob Heckford, and Juhani Itämies for providing information and help with many kinds of matters during the preparation of this study. DNA barcoding was conducted at the Centre for Biodiversity Genomics, to whose staff we are grateful for their continuous support. Thomas Pape, Natural History Museum of Denmark, kindly advised on questions about nomenclature. Sequencing was financially supported by the Academy of Finland, Kone foundation and Finnish Cultural foundation through grants to the Finnish Barcode of Life project and furthermore supported by the Promotion of Educational Policies, University and Research Department of the Autonomous Province of Bolzano - South Tyrol with funds to the projects “Genetische Artabgrenzung ausgewählter arktoalpiner und boreomontaner Tiere Südtirols” and “Erstellung einer DNA-Barcode-Bibliothek der Schmetterlinge des zentralen Alpenraumes (Süd-, Nord- und Osttirol)”. Andrew Liston kindly checked the English language. Finally, we are indebted to Bengt Å. Bengtsson, an anonymous reviewer, and Erik J. van Nieukerken for many useful comments on the earlier version of this paper.

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