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Article

On Setina irrorella pseudoirrorella Freina and Witt, 1985, and Setina cantabrica Freina and Witt, 1985, in Spain and Portugal: Taxonomic Status, Distribution, Habitat, and Molecular Genetics (Lepidoptera: Erebidae: Arctiinae: Lithosiini)

by
Ramon Macià
1,*,
Josep Ylla
2,
Guillem Masó
3 and
Josep Bau
3
1
Natural Sciences Museum of Barcelona, Natura Laboratory, Arthropod Collection, Passeig Picasso s/n, 08003 Barcelona, Spain
2
C. Principal, Urb. Serrabonica, 08503 Gurb, Spain
3
Biosciences Department, University of Vic—Central University of Catalonia, 08500 Vic, Spain
*
Author to whom correspondence should be addressed.
Taxonomy 2025, 5(2), 23; https://doi.org/10.3390/taxonomy5020023
Submission received: 20 February 2025 / Revised: 14 April 2025 / Accepted: 17 April 2025 / Published: 21 April 2025
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Abstract

:
The taxonomic positions of species in the genus Setina are poorly known and not well established, with ongoing uncertainties in their classification. In this work, the distribution of Setina irrorella pseudoirrorella Freina and Witt, 1985, and Setina cantabrica Freina and Witt, 1985, in Spain and Portugal is updated, expanding their distribution area based on revised bibliographic data and information provided by researchers and public or private collections. The taxonomic status of the binomial flavicans–irrorella and cantabrica–roscida are discussed, and their specific validity is analysed, comparing the differences observed between these taxa, which are studied on the basis of external morphological characteristics, genitalia, and molecular genetics. An updated distribution map is presented. The authors conclude that S. cantabrica and S. i. pseudoirrorella are the only two species of the genus Setina present in the Iberian Peninsula.

1. Introduction

The species of the genus Setina Schrank, 1802, are often poorly known, and their taxonomic position is not always well established. The same occurs with the two taxa present in the Iberian Peninsula, Setina irrorella Linnaeus, 1758, and Setina cantabrica Freina and Witt, 1985, the taxonomic position of which is far from being resolved.
The genus Setina was described by Schrank, 1802, taking Phalaena irrorella Linnaeus, 1758, as the type species. It is represented by a group of species of small average size, exhibiting considerable sexual dimorphism, with larger males. They have ochre or yellow forewings and paler hindwings, and the forewings have three rows of dark spots that can be totally or partially reduced, while in some cases they merge into longer diffuse stripes [1]. Specific external differences are found in the structure of the male antennae and the body colouration and certain aspects of the colouration and intensity of the wing markings, which have a notable degree of variation, and may show, especially the markings on the forewings, a considerable fusion between members of the genus.
It is a difficult group from the taxonomic point of view due to the great individual and interpopulation variability. Furthermore, male genitalia are quite uniform in some species of the genus, with specific characteristics not very prominent. In some of them, the distinctive characteristics are found in the female genitalia. Due to the significant taxonomic complexity, exhaustive research is essential to resolve the existing uncertainties surrounding the genus.
Various species, subspecies, and forms or races have been described from the genus Setina, some relegated to synonymy or of questionable validity.
Several authors [1,2,3,4,5,6] provide us with a diverse but significant overview of this genus. It is true that populations of species of the genus Setina are often poorly known, and their nomenclature is not always well established; the differences between species and subspecies of the genus must be clearly identified, as uncertainties still exist. Certain taxa have also been validated or rejected, some possibly without sufficiently clear scientific justification. As some of these authors explain, Setina populations show dominant morphs depending on altitude [6]. The fact that there are populations from the Mediterranean coast to more than 3000 m in altitude means that the variability due to altitudinal variation is significant.
This work aims to elucidate the specific validity of S. cantabrica and S. irrorella, considering the taxon flavicans a synonym of irrorella, with S. i. pseudoirrorella being the northern Iberian Peninsula subspecies. Based on the reviewed material, we present an updated distribution map of these two taxa in Spain and Portugal.

2. Materials and Methods

The study was based on the compilation of published and unpublished data on both S. i. pseudoirrorella and S. cantabrica in the Iberian Peninsula (Spain and Portugal). Therefore, in addition to carrying out a bibliographic review, the material from the Museum of Natural Sciences of Barcelona (MCNB), from the private collections of the authors and of Eliseo Fernández Vidal, Rafael Magro, Jordi Dantart, and Arcadi Cervelló, was examined. The records with images from GBIF.org (2 January 2025) from Spain and Portugal and those from Virtual Biodiversity were also reviewed, checking in each case the correct identification of the specimens (disregarding those that are doubtful or misidentified). Finally, a good number of unpublished data was provided by colleagues of proven academic rigor.
The data collected allowed for the elaboration of distribution maps, with 10 × 10 km UTM grids, corresponding to S. i. pseudoirrorella (with more than 500 records) and S. cantabrica (with more than 300 records).
Several specimens of S. irrorella and S. roscida from different European localities were also examined to compare them with the Iberian specimens.
For the dissection and preparation of the genitalia, the procedure defined by Robinson [7] was followed with modifications. The images of the genitalia were taken with the Euromex Oxion series microscopes Leica DMLB, Leica MZAPO, NIKON Eclipse a100, and E400 and the digital cameras Leica DFC550, NIKON D3100, and SONY DSLR-A100K with 102.8 AF lens. The images were taken with a Nikon D90 digital camera with a DX105 lens. For image edition, Adobe Photoshop© was used.
From the studied material, total DNA was extracted from the legs of dry specimens. DNA extraction was performed using the NZY Tissue gDNA Isolation kit (nzytech, Lisbon, Portugal) following the manufacturer’s instructions for use or using 10% Chelex 100 (Bio-Rad, Richmond, CA, USA). All laboratory equipment used in this process (scissors, tweezers, and homogenisation instruments) were washed sequentially with 20% bleach and absolute ethanol between consecutive samples. The mitochondrial marker COI-5P was amplified by PCR using primers LCO and HCO [8]. In the case of samples that presented amplification difficulties, intermediate primers modified from SphF4 and SphR3 were used [9], reconstructing the sequence from two resulting fragments. Successful PCR samples were cleaned using the Exo-SAP method; Sanger sequencing was performed by Eurofins Genomics (Ebersberg, Germany) or Macrogen Europe (Amsterdam, The Netherlands).
A total of 30 samples were analysed (Table 1). The data were compiled in the public dataset at BOLD (http://v4.boldsystems.org) under the access code SETI. For the phylogenetic study, a selection of sequences (107 in total) downloaded from the BOLD Public Records Barcode Database on 9 January 2025 and corresponding to specimens of the genus Setina collected in several European countries was also added. The selection criterion for these downloaded sequences was a length between 600 and 700 nucleotides. These sequences (107 in total) were labelled as S. irrorella (n = 57), S. roscida (n = 14), S. aurita (n = 26), S. aurata (n = 3), S. flavicans (n = 2), S. alpestris (n = 1), S. kuhlweini (n = 1), and Setina sp. (n = 3). Two sequences from Nudaria mundana (Hübner, [1809]) were used as outgroup.
Sequence alignment and assembly, phylogenetic analyses, best-fit DNA model determination, and pairwise distance analysis were performed using MEGA 10.1.8 [10].
Estimates of evolutionary divergence between and within groups were calculated as the average number of base substitutions per site across all sequence pairs using the Tamura 3-parameter model [11], with distances computed through a bootstrap analysis of 1000 replicates. Grouping for distance comparisons was based on the main clades identified through Bayesian inference, considering the 30 sequences obtained in this study and the 89 sequences downloaded from BOLD, all of which were assigned to one of the four major clades: the irrorella, aurita, cantabrica, and roscida groups. The generation of phylogenetic trees was carried out under Bayesian inference with BEAST2 v. 2.6.6 [12], using the HKY+G model.

3. Results

3.1. Historical Taxonomic Status of Setina Species in the Iberian Peninsula

Gómez Bustillo (1979) [13] refers to this taxon as Philea irrorella Linnaeus, 1758, found in different Iberian localities. He also considers the possible presence in Granada (Sierra Nevada), though these data are not confirmed. Other authors such as Freina and Witt (1985) [2] and Ylla et al. (2010) [14] state that flavicans is a different species from S. irrorella. They consider that irrorella is not found in the Iberian Peninsula and regard Setina flavicans pseudoirrorella Freina and Witt, 1985, as an endemic subspecies of the north of the Iberian Peninsula, populating the Pyrenees and its northern slopes. Redondo (1990) [15] refers to it as S. irrorella from various localities in the province of Huesca, the Sierra del Moncayo (Zaragoza), and Orihuela del Tremedal (Teruel). According to Leraut. (2006) [5], S. irrorella irrorella Linnaeus, 1758, is the ssp. occurring in northern Europe, Russia, and western Europe (eastern Austria, Savoy, and northern Italy); meanwhile, S. i. flavicans would be the subspecies occurring in the French Alps, central and southern Provence, Pyrenees, Spain, and Portugal. The name pseudoirrorella would be a synonym of the last one. Witt and Ronkay (2011) [1] do not question the validity of flavicans, while Corley (2013) [16] considers that the specimens found in Portugal should be attributed to S. cantabrica. Leraut, G. (2018) [6] states that the genitals of S. flavicans do not differ from the genitals of S. i. irrorella. The yellow colour of the abdomen, the wing pattern, and the reduction of the black markings, are frequent in Provence, in the Pyrenees, and in general in the south of France. Thus, according to his research, flavicans represents the southern subspecies of S. irrorella. It is found in France, Spain, Italy, and certainly throughout the northern Mediterranean basin. The taxon pseudoirrorella, initially described as a subspecies of S. flavicans in the southwest of its distribution (Ariège, Hautes-Pyrénées, etc.), corresponds simply to a form of Setina irrorella flavicans Geyer, 1836, in which the abdomen is devoid of yellow scales. Both forms are allopatric (with a contact zone in some localities of the Pyrenees), and their genitals do not differ. Therefore, he requests for the ssp. flavicans with the black abdomen, the name of S. irrorella ssp. pseudoirrorella Freina and Witt, 1985, to be a new combination.
Regarding S. cantabrica Freina and Witt, 1985, some authors [1,16,17] have followed Fernández Vidal et al. (2003) [4], accepting it as a good species, while others [5,6] have continued to question its specific validity, considering it as Setina roscida cantabrica Freina and Witt, 1985 [2]. According to Leraut (2006) [5], Setina roscida roscida Denis & Schiffermüller, 1775, is the ssp. that flies in Austria and France, east and southwest, while in the Cantabrian Mountains of Spain, it is the ssp. S. r. cantabrica [2]. Leraut (2018) [6] agrees with this view in the study of the genitalia, where he shows in the first place that S. r. roscida differs significantly from the other species of the group. In the male, the subcostal process of the valve is smaller and thicker; in the female, the bursa is thick and rounded. Setina roscida kuhlweini Hübner, 1824, is similar, but the subcostal process of the valve of the male is wider, and the corpus bursa of the female is more elongated. The genitalia of S. r. cantabrica are similar to those of S. r. kuhlweini, the female rather presenting a symmetrical appendix bursae. Based on his diagnosis, he considers S. cantabrica as a subspecies of S. roscida. In his study of Portuguese specimens, Corley (2013) [16] looks at the intensity and arrangement of the markings on the wings and abdomen and finds that the posterior marks agree well with those described for cantabrica, while the anterior marks, especially those on the hind wings, are paler. He also points out that the genitalia of the Portuguese individuals coincide with those of roscida. In his opinion, these two findings would make the assignment of the specimens from Portugal to the subspecies S. r. cantabrica acceptable. However, the final observation that the wingspan of Portuguese individuals is greater than that of roscida specimens (28 mm compared to 20/24 mm) leads him to consider cantabrica as a good species.

3.2. Setina irrorella pseudoirrorella Freina and Witt, 1985. Stat. rev.

Setina flavicans pseudoirrorella Freina and Witt, 1985.
Original combination: Setina flavicans pseudoirrorella Freina and Witt, 1985. LT. Spain. Pyrenees Orientales, based on various captured specimens, both males and females, with the Holotype being: Pyrenees Orientales, Port de Toses (Girona) l800 m, 20,8.198l, leg. W. PAVLAS.

3.2.1. Material Examined for Setina irrorella pseudoirrorella

The following material was examined:
ARABA: 2 ♂, Urbasa, Mendiak, 800 m, 30TWN64, 6.VIII.1876, Col. Rius, MCNB.
ASTURIAS: 1 ♂, Los Llanos, Mocin, 889 m, 30TTN69, 22.VI.2019, BV.; 1 ♂, Aleiva, 592 m, 30TUP60, 14.VII.2019, BV.; 1 ♂, Tuiza de Arriba, 1589 m, 30TTN67, 15.VII.2016, BV.; 1 ♂, Aller, 1399 m, 30TUN07, 1.VIII.2016, BV.; 1 ♂, Somiedo, 1636 m, 29TQH27, 27.IX.2014, BV.; 1 ♂, Braña de Mumián, Somiedo, 1396 m, 29TQH27, 2.IX.2013, BV.; 1 ♂, Fuensanta, 580 m, 30TTP90, 15.VII.2012, BV.; 1 ♂, Puerto Ventana, 1760 m, 30TTN57, 21.VI.2012, BV.; 1 ♂, Gamoniteiro, 1745 m, 30TTN68, 12.VIII.2011, BV.; 1 ♂, Cangas de Onis, 1080 m, 30TUN39, 15.V.2011, BV.; 1 ♂, Lena, 1332 m, 30TTN68, 7.IX.2009, BV.; 1 ♂, Dolia, 839 m, 29TQH29, 24.V.2007, BV.; 1 ♂, Collado de Pandébano, 1200 m, 30TUN58, 18.VIII.2012, Guerrero leg.; 1 ♂, Valle del Lago, 1220 m, 30TQH27, 17.VIII.2012, Guerrero leg.
BARCELONA: 2 ♂, Coll de la Bena, Berguedà, 1440 m, 31TCG9879, 1.VII.2000, J. Dantart leg.; 2 ♂, Cal Tasconet, Coll de la Baena, Berguedà, 1420 m, 31TCG9879, 1.VII.2000, J. Dantart leg.; 2 ♂, Castellar de n’Hug, Berguedà, 1397 m, 31TDG1981, 27.VIII.2011, J. Pibernat leg.; 1 ♂, Gisclareny, Berguedà, 1300 m, 31TDG9978, 1.VII.2000, Dantart leg.; 6 ♂, 1 ♀, Castell de Montesquiu, Osona, 681 m, 31TDG3563, 15.IX.2007, 12.X.2007, R. Macià leg.; 1 ♂, Font Pomereta, Osona, 1180 m, 31TDG4530, 8.VII.2004, J. Dantart leg.; 1 ♂, Rasos de Peguera, 1813 m, 31TCG96, 31.VIII.1921, M. Rondós leg.; 1 ♂, Coll de Pal, 1963 m, 30TDG08, 29.VIII.2016, BV.; 1♂, Guardiola de Berguedà, 1192 m, 31TDG08, 22.VIII.2015, BV.; 1 ♂, Castellar de n’Hug, Berguedà, 1395 m, 31TDG18, 4.VIII.2005, J. Martí leg.; 1 ♂, Gòsol, Berguedà, 1423 m, 31TCG87, 19.IX.2014, J. Martí leg.; 2 ♂, Serra de Cabrera, Puigsacalm, 1100 m, 31TDG46, 24.VI.1989, Col. Rius, MCNB; 4 ♂, Urb. La Roca, Taradell, 520 m, 31TDG43, 28.VIII.1988, R. Macià leg.; 1 ♂, Castellar de n’Hug, 1390 m, 31TDG18, 8.VIII.1980, Col. Rius, MCNB; 1 ♂, Camí de Sobrepuny, La Nou, 1548 m, 31TDG06, 12.IX.2021, J. Planes leg.; 1 ♂, Serrabonica, Vespella, Gurb, 31TDG34, J. Ylla leg.; 1 ♂, Castellà de n’Hug, 1400 m, 31TCG28, 6.VIII.2004, A. Cervelló leg.; 1 ♂, Boca sud del Cadí, 1175 m, 31TCG87, 2.VII.2011, A. Cervelló leg.; 1 ♂, Sant Sadurní d’Osormort, 500 m, 31TDG54, 18.IX.1978, J.Ylla leg.; 3 ♂, Parc del Castell de Montesquiu, Osona, 707 m, 31TDG3463, 15.IX.2007, J. Ylla leg.
BURGOS: 1 ♂, Páramo de Masa, Masa, 950 m, 30TVN41, 24.VII.1999, A. Blázquez leg.; 1 ♂, San Pantaleón del Páramo, 900 m. 30TVN30, 1.VIII.1998, A. Blázquez leg.; 1 ♂, Montoiro, 1033 m, 30TVN31, 6.VII.2011, BV.; 1 ♂, Fuentelsoto, Cobarrubias, 1020 m. 30TVM65, 4.VII.2019, R. Macià leg.; 3 ♂, 1 ♀, Loma del Rei, Quintanilla Sobresierra, 1003 m, 30TVN41, 24.VI.2009, 13.VI.2011, 13.VII.2012, R. Macià leg.; 2 ♂, El Cerro, Gredilla la Polera, 980 m, 30TVN40, 14.VI.2017, R. Macià leg.;2 ♂, El Robledo, Paramos de Lora, 992 m, 30TVN33, 7.VII.2015, R. Macià leg.; 1 ♂, Sedano, 804 m, 30TNV32, 27.VII.2002, R. Magro leg.; 1 ♂, Paramo Mozuelos de Sedano, 922 m, 30TVN32, 1.VI.2009, R. Magro leg.; 1 ♂, Mata, 900 m, 30TNV40, 21.VII.2012, R. Magro leg.; 1 ♂, Burgos, 861 m, 30TVM48, 14.VII.1993, R. Magro leg.; 3 ♂, Paramo de Masa, 1045 m, 30TVN41, 16.IX.1986, 19.VIII.1990, R. Magro leg.; 4 ♂, Quintanilla Sobresierra-Mata, Páramos de Masa, 900–1000 m, 30TVN40, 8.VII.1998, 2.VII,2000, J.Ylla leg.
CANTABRIA: 10 ♂, Fuente De, El Cable, 1834 m, 30TUN57, 23.VII.1991, R. Macià leg.; 1 ♂, Pujayo, 395 m, 30TVN17, 27.VI.2018, VB; 1♂, Horcadina de Covarrobres, 1908 m, 30TUN58, 8.VI.2011, BV; 1 ♂, Fuente De, 1095 m, 30TUN56, 11.VII.1990, R. Magro leg.; 3 ♂, Fuente De, El Cable, 1834 m, 23.VII.1991, J. Ylla leg.
GUIPUZCOA: 1 ♂, Zegama, 1300 m, 30TWN55, 28.VII.2007, BV.
GIRONA: 2 ♂, Pla de la Baga, Cerdanya, 1415 m, 31TDG0890, 12.IX.2010, E. Guzman leg.; 2 ♂, Torrent dels Bous, Gréixer, Cerdanya, 1250 m, 31TDF0395, 11.IX.2010, J. Dantart leg.; 2 ♂, Pla de Capllong, Cerdanya, 1700 m, 31TCH9801, 11.IX.2010, J. Dantart leg.; 1 ♂, Camprodón, Ripollès, 988 m, 31TDG4784, VII.1975, J. Dantart leg.; 2 ♂, Campelles, El Ripollès, 1303 m, 31TDG28, 19.VIII.1985, A. Moliné leg., J. Ylla col.; 1 ♂, Fontalba-Puigmal, El Ripollès, 2000–2900 m, 31TDG38, 29.VII.1990, J. Ylla leg.; 2 ♂, Ribes Altes, El Ripollès, 1850 m, 31TDG18, 21.VIII.1988, J. Ylla leg,; 1 ♂, Queralbs, El Ripollès, 1180 m, 31TDG38, 28.VII.1987, J. Ylla leg.; 1 ♂, Coll de Jou, El Ripollès, 1250 m, 31TDG38,12.VII.990, J. Ylla leg.; 2 ♂, Setcases-Espinavell, El Ripollès, 1700 m, 31TDG49, 24.VIII.1989; J. Ylla leg.; 2 ♂, Guils de Cerdanya, La Cerdanya, 1455 m, 31TDH00, 10.VIII.2003, 13.VIII.2005, J. Ylla leg.; 2 ♂, Serra de l’Estremera, 1768 m, 31TDG28, 29.VII.1995, 9.VII.2003, J. Ylla leg.; 2 ♂, Palau. La Cerdanya, 1400 m, 31TDG19, 16.VIII.2008, J. Ylla leg,; 1 ♂, Cereja, Llívia, La Cerdanya, 1441 m, 31TDH10, 15.VIII.2008, J. Ylla leg.; 1 ♂, Les Planes de Nevà, El Ripollès, 1650 m, 31TDG28, 1/VII/1986, J. Ylla leg.; 1 ♂, La Molina, El Ripollès, 2000 m, 31TDG18, 5.VII.!986, J. Ylla leg.; 3 ♂, Vall Ter, El Ripollès, 31TDG4287/3996, 2000–2220 m, 9.VIII.2023, J. Ylla leg.; 1 ♂ Serra dels Clots, Vall d’Eina, Alta Cerdanya, 2000 m, S1TDG2799, VII.1996, J. Pibernat leg.; 2 ♂, Comaoriola, Cóms de Das, Cerdanya, 1980 m, 31TDG0887, 12.IX.2010, E. Guzman leg.; 4 ♂, 1 ♀, Girul, Cerdanya, 1540 m, 31TCH9900, 2.VII.2005, J. Dantart leg.; 3 ♂, Gombrèn, Ripollès, 912 m, 31TDG2577, 27.VIII.2011, M. Rondós leg.; 1 ♂, La Molina, Cerdanya, 1420 m, 31TDG1488, VIII.1990, J. Dantart, leg.; 6 ♂, Mosoll, Cerdanya, 1100 m, 31TDG0692, 2.IX.2013, 21.VIII.2014, J. Dantart leg.; 1 ♂, Olot, Garrotxa, 447 m, 31TDG5770, 25.VIII.2010, E. Guzman leg.; 1 ♂, Les Planelles del Ramonet, Cerdanya, 1238 m, 31TDG1190, 19.VI.2015, J. Dantart leg.; 1 ♂, Bosc del Quer, Eina, Alta Cerdanya, 1700 m, 31TDH2501, VII.1996, J. Dantart leg.; 2 ♂, Rupit, Osona, 845 m, 31TDG5552, 5.VIII.2000, J. Dantart leg.; 1 ♂, St. Hilari Sacalm, La Selva, 805 m, 31TDG5396, 10.VIII.1979, J. Dantart leg.; 1 ♂, Setcases, Ripllès, 1265 m, 31TDG4291, 1.VIII.1911, A. Codina leg.; 2 ♂, Surroca de Baix, Ripollès, 1170 m, 31TDG4080, 10.IX.2010, Ll. Abós, leg.; 1 ♂, Susqueda, La Selva, 320 m, 31TDG6148, 2.VIII.1910, A. Codina leg.; 3 ♂, Vallter, Ripollès, 2160 m, 31TDG3997, 26.VIII.1989, J. Dantart leg.; 2 ♂, Vidrà, Osona, 996 m, 31TDG4363, 9.VIII.1980, J. Dantart leg.; 1 ♂, Rierol de la Vila Vella, Vidrà, Osona, 920 m, 31TDG4463, 11.VIII.2916, A. Cervelló leg.; 1 ♂, Bruguera, 1176 m, 31TDG38, 17.VIII.2018, BV.; 1 ♂, Queralbs, 1878 m, 31TDG29, 30.VII.2017, BV.; 1 ♂, Mosoll, 1107 m, 31TDG09, 21.VIII.2014, BV.; 1 ♂, Campelles, 883 m, 31TDG37, 13.VIII.2013, BV.; 2 ♂, 1 ♀, Montgrony, Ripollés, 1350 m, 31TDG27, 4.VIII1990, J. Martí leg.; 1 ♀, Gréixer, Baixa Cerdanya, 1340 m, 31TDG09, 12.VIII.2002, J. Martí leg.; 1♂, Cal Tinent, Greixer, Moixeró, 1000 m, 31TDG09, 25.VIII.1985, Col. Rius, MCNB; 1 ♂, Puigsacalm, 1170 m, 31TDG46, 15.VI.1990, Col. Rius, MCNB; 1 ♂, Serrat d’Eurti, Vilallobent, 1410 m, 31TDG19, 21.VIII.1992, Col. Rius, MCNB; 1 ♂, Torrent set Fonts, Tosa d’Alp, La Molina, 2080 m, 31TDG08, 24.VII.1993, Col. Rius, MCNB; 1 ♂, Viladrau, 890 m, 31TDG43, 10.VIII.1919, Novellas, MCNB; 1 ♂, Pla Traver, Ciuret, 1200 m, 31TDG46, 6.VIII.1926, Codina, MCNB; 1 ♂, Coll de Santigosa, 1057 m, 31TDG47, 10.VIII.1978, Pérez de Gregorio, MCNB; 1 ♂, Alp, 1250 m, 31TDG09, 26.VIII.1982, Vallhonrrat, MCNB; 1 ♂, Puigsacalm, 1170 m, 31TDG46, 4.VII.1980, Pérez de Gregorio, MCNB; 1 ♂, Queralbs, 1075 m, 31TDG29, 7.VIII.1981, Pérez de Gregorio, MCNB; 2 ♂, Coll de Toses, 1800, 31TDG18, 7.VII.1986, Pérez de Gregorio, MCNB; 3 ♂, Vall Ter, 2000 m, 31TDG39, 2.VII.1986, Pérez de Gregorio, MCNB; 1 ♂, Molló, 1176 m, 31TDG58, 25.VIII.1987, Artigas, MCNB; 1 ♂, Camprodón, 963 m, 31TDG48, 2.VIII.1927, M. Ibarra, MCNB; 1 ♂, Queralbs, 1236 m, 31TDG38, 16.IX.2011, A. Cervelló leg.; 1 ♂, Central Hidroelectrica de Deió, Queralbs, 1300 m, 31TDG39, 21.VII.2021, A. Cervelló leg.; 2 ♂, Navá, 1265 m, 31TDG28, 4.VIII.1990, R. Macià leg.; 1 ♂, Vilamanya, 1048 m, 31TDG38, 7.VIII.2015, R. Macià leg.; 3 ♂, Fontalba, 2000 m, 31TDG39, 29.VII.1990, R. Macià leg.; 2 ♂, Espinavell, 1500 m, 31TDG49, 15.VIII.2002, R. Macià leg.; 1 ♂, La Molina, 1900 m, 31TDG18, 18.VII.1992, R. Macià leg.; 4 ♂, Refugi de Ribes Altes, 1522 m, 31TDG38, 20.VIII.1988, R. Macià leg.; 2 ♂, Serra d’Estremera, Queralbs, 1890 m, 31TDG28, 26.VIII.2000, R. Macià leg.; 4 ♂, 1 ♀, Font Home Mort, Queralbs, 1500 m, 31TDG28, 9.VII.1999, 22.VII.2006, R. Macià leg.; 2 ♂, Vidrà, 980 m, 31TDG46, 11.VIII.2016, R. Macià leg.
HUESCA: 1♂, Canal de Izas, Canfranc, 1491 m, 30TYN0537, 26.VI.1999, T. Latasa leg.; 1 ♂, Canfranc, 1200 m, 30TYN03, 27.VIII.2019, BV.; 1 ♂, Montany, 2417 m, 31TCH01, 23.VII.2011, BV.; 1 ♂, Chisagües, 1431 m, 31TBH62, 14.VIII.2010, BV.; 1 ♂, Valle de Ordesa, 1033 m, 30TYN32, 2.VIII.2018, Weiss, MCNB; 1 ♂, Port de Benasque, 1138 m, 30TBH92, 24.VII.1921, Novellas, MCNB; 1 ♂, Hecho, 834 m, 30TXN83, 6.VIII.1999, Escolà, MCNB; 2 ♂, Fuente de la Salud, Canfranc, 1370 m, 30TYN03, 9.VII.2021, R. Macià leg.; 1 ♂, Estación de Candanchú, 1640 m, 30TYN03, 10.VII.12021, R. Macià leg.; 1 ♂, Ibon de Acherito, 1800 m, 30TXN84, 10.VIII.2001, V. Redondo leg.; 1 ♂, Forau de Aiguallut, 2000 m, 30TCH02, 8.VIII.1976, V. Redondo leg.; 1 ♂, Collado de Sahún, 1900 m, 31TBH91, 5.VIII.1979, V. Redondo leg.; 1 ♂, La Serra, Respomuso, 1800 m, 30TYN14, 26.VII.1986, V. Redondo leg.; 1 ♂, Ibon de Piedrafita, 1600 m, 30TYN13, 5.VIII.1985, V. Redondo leg.; 1 ♂, Collado las Blancas, Borau, 2100 m, 30TXN92, 13.VIII.1977, V. Redondo leg.; 1 ♂, Balneario de Panticosa, 1700 m, 30TYN23, 2.IX.1984, V. Redondo leg.; 1 ♂, Camino de la Mina, Cerler, 1600 m, 31TBH91, 27.VII.1988, V. Redondo leg.; 1 ♂, Astún, 1800 m, 30TYN04, 20.VII.2022, V. Redondo leg.; 4 ♂, 1 ♀, Lago Urdiceto, Bielsa, 2000 m, 31TBH72, 12.VII.1995, R. Macià leg.; 1 ♂, Bielsa, 1053 m, 31TBH72, 16.VII.1999, R. Macià leg.; 2 ♂, Coll del Portalet, 1700 m, 30TYN14, 11.VII.2015, R. Macià leg.; 1 ♂, Coll del Portalet, 1600–1800 m, 30TYN14, 11.VII.2015, J. Ylla leg.
LA RIOJA: 4 ♂, Pineda, Hoyuelas, Lumbreras, 1300 m, 30TWM3660, 13.VII.2001, T. Latasa leg.; 1 ♂, La Blanca, Villoslada de Cameros, 1100 m, 30TWM2556, 9.VIII.2000, Garzón leg.; 1 ♂, Hoyos de Iregua, Volloslada de Cameros, 1800 m, 30TWM2152, 7.VII.2000, T. Latasa leg.; 2 ♂, Camping Villoslada de Cameros, 1100 m, 30TWM2658, 15.VI.2001, T. Latasa leg.; 1 ♂, Barranco de los Cayos, Cornago, 750 m, 30TWM7559, 29.VII.2016, T. Latasa leg.; 1 ♂, Viniegra de Arriba, 1456 m, 30TWM15, 19.VI.2022, T. Latasa leg.; 4 ♂, P.N. Sierra de Cebollera, 1480 m, 30TWM25, 27.VII.2007, R. Macià leg.
LLEIDA: 1♂, Saut deth Pish, Arrós, Val d’Aran, 1554 m, 31TCH2238, 18.VII.2008, A. Blazquez leg.; 1 ♂ Camping la Verneda, El Pont d’Arrós, 950 m, 31TCH13, 3.VII.1994, R. Macià leg.; 3 ♂, Adons, Alta Ribagorça, 1350 m, 31TCH2189, 3.VIII.2019, A. Cervelló leg.; 1 ♂, Bassa d’Arrés, Val d’Aran, 1520 m, 31TCH1337, 6.VII.1984, J. Dantart leg.; 4 ♂, Bagergue, Val d’Aran, 1419 m, 31TCH2931, 5.VII.1983, J. Dantart leg.; 2 ♂, Coll del Boix, Solsonès, 1250 m, 31TCG6668, 11.VII.2000, J. Dantart leg.; 2 ♂, Coll del pas del Coro, Pallars Sobirà, 1950 m, 31TCH4021, 28.VI.2003, J. Dantart leg.; 1 ♂, Estaon, Pallars Sobirà, 1237 m, 31TCH5316, 21.VII.2007, J. Dantart leg,; 1 ♂, Cortal del Mateu, Lles, Cerdanya, 1265 m, 31TDG9292, 23.IX.2017, A. Cervelló leg.; 1 ♂, Tuc de Maubèrme, Val d’Aran, 2880 m, 31TCH2940, 16.VII.1998, J. Dantart, leg.; 1 ♂, Plans de Mont, Val d’Aran, 1680 m, 31TCH2437, 18.VII.2014, A. Cervelló leg.; 2♂, 1 ♀, Pletiu dera Montanheta, Val d’Aran, 1860 m, 31TCH2982, 26.VII.2008, A. Cervelló leg.; 1 ♂, Còth de Montoliu, Val d’Aran, 2480 m, 31TCH3038, 23.VII.1988, J. Dantart leg.; 5 ♂, Plans de Nera, Valarties, Val d’Aran, 1410 m, 31TCH2524, 1.VIII.2019, E. Guzman leg.; 2 ♂, Salardú, Val d’Aran, 1268 m, 31TCH2830, 18.VII.1981, J. Dantart leg.; 8 ♂, 2 ♀, Tredós, Val d’Aran, 1798 m, 31TCH3025, 1.VIII.2019, J. Dantart leg.; 6 ♂, 1 ♀, Vanhs de Tredós, Val d’Aran, 1800 m, 31TCH3024, 26.VII.2008, 3.VIII.2009, 4.VII.2010, R. Macià leg.; 8 ♂, Vanhs de Tredós, Val d’Aran, 1800 m, 31TCH3024, 26.VII.2008, A. Xaus leg.; 1 ♂, Baqueira-Beret, 1515 m, 31TCH32, 7.7.2018, BV.; 1 ♂, Abella de la Conca, 1214 m, 31TCG46, 21.VIII.2018, BV.; 1 ♂, Boixols, 1270 m, 31TCG57, 21.VIII.2018, BV.; 1 ♂ Tabascan, 1905 m, 31TCH52, 2.VIII.2017, BV.; 1 ♂, Arrós, 1427 m, 31TCH13, 14.VII.2017, BV.; 1 ♂, Espot, 1318 m, 31TCH41, 6.VIII.2014, BV.; 1 ♂, Bagergue, 2071 m, 31TCH23, 16.VIII.2011, BV.; 1 ♂, Vall de Bohí, 2529 m, 31TCH21, 22.VII.2022, BV.; 1 ♂, La Guingueta d’Àneu, 949 m, 31TCH41, 8.VII.2010, BV.; 1 ♀, Alinyà, Alt Urgell, 957 m, 31TCG67, 11.VII.2000, J. Martí leg.; 1 ♂, St. Joan de Toran, Val d’Aran, 1035 m, 31TCH24, 4.VII.1994, J. Martí leg.; 2 ♂, Port de la Bonaigua, Pallars Sobirà, 2050 m, 31TCH32, 30.VII.1994, J. Martí leg.; 1 ♂, Tirvia, Pallatrs Subirà, 990 m, 31TCH50, 28.VII.1995, J. Martí leg.; 1 ♂, Riu Escrita, Espot, 980 m, 31TCH41, 3.VIII.1985, Col Rius, MCNB; 1 ♂, Estanys de la Pera, 2330 m, 31TCH80, 30.VII.1988, Col. Rius, MCNB; 1 ♂, Estany petit de la Pera, 2300 m, 31TCH80, 9.VIII.1985, Col. Rius, MCNB; 1 ♂, Coll de Jou, 1468 m, 31TCG76, 24.VIII.1982, F. Vallhonrrat, MCNB; 3 ♂, Port de la Bonaigua, 2072 m, 31TCH32, 14.VIII.2005, 16.VII.2007, 15.VII.2010, Pérez de Gregorio, MCNB; 1 ♂, Llac Gerber, 2340 m, 31TCH32, 17.VII.1986, Col. Artigas, MCNB; 1 ♂, Cap de Baqueira, 2500 m, 31TCH32, 3.VIII.1986, Col. Artigas, MCNB; 1 ♂, Salardú, 1260 m, 31TCH23, 18.VII.1957, M. Ibarra, MCNB; 1 ♂, Font de Tarrés, Gòsol, 1502 m, 31TCG87, 19.IX.2014, J. Planes leg.; 1 ♂, La Bargadera, 1950 m, 31TCH22, 4.VIII.2013, J. Planes leg.; 1 ♂, Plans de Sotllo, Alins, 1250 m, 31TCH62, 2.IX.2007, J. Planes leg.; 1♂, Es Bordes, 865 m, 31TCH13, 20.VIII.2012, A. Cervelló leg.; 2 ♂, Loseron, 1850 m, 31TCH22, 23.VII.2006, A. Cervelló leg.; 5 ♂, Vall d’Horno, 1550 m, 31TCH12, 13.VIII.2012, 15.VII.2017, A. Cervelló leg.; 2 ♂, Tirvia, 31TCH50, 11.VIII.1986, A. Cervelló leg.; 1 ♂, Alós d’Isil, 1270 m, 31TCH52, 16.VII.2011, A. Cervelló leg.; 1 ♂, Sorribes, 1370 m, 31TCG97, 2.VII.2011, A. Cervelló leg.; 10 ♂, Sanillers, Travesseres, 1140 m, 31TCG99, 23.IX.2017, 31.VIII.2019, 7.VIII.2020, 23.VII.2022, A. Cervelló leg.; 2 ♂, Josa del Cadí, Tuixent, 1431 m, 31TCG87, 21.VIII.2014, R. Macià leg.; 2 ♂, Pla de Beret, 31TCH33, 17.VII.1999, R. Macià leg.; 1 ♂, Tuc de Cortajàs, 2300 m, 31TCH23, 20.VII.2000, R. Macià leg.; 4 ♂, 1 ♀, Bassa d’Arrès, Vilamós, 1425 m, 31TCH13, 20.VI.1998, R. Macià leg.; 1 ♂, Camí de Valarties, 1350 m, 27.VI.2012, R. Macià leg.; 3 ♂, 1 ♀, Port de la Bonaigua, 2100 m, 31TCH32, 13.VII.1997, R. Macià leg.; 3 ♂, Pla de l’Artiga de Lin, Val d’Aran 6.VIII.1988, J.Ylla and J. Junyent, leg.; 1 ♂, Arabell, Alt Urgell, 992 m, 31TCG69, 15.VIII.2004, J. Ylla leg.
LEÓN: 2 ♂, Villanueva de Arriba, 31.VII. 1998, 1200 m, 30TUN53, R. Macià leg.; 1 ♂, Valporquero de Torio, 1347 m., 30TTN95, 30.VII.2021, A. Blazquez leg.; 1 ♂, Lugán, 900 m, 30TUN03, 30.VII.1998, A. Blazquez leg.; 1 ♂, Collado Jermoso, 2064 m, 30TUN48, 14.VIII.2017, BV.; 1 ♂, Cordiñanes, 2065 m, 30TUN48, 22.VIII.2013, BV.; 1 ♂, Puerto de Agüera, Cubilla de Arbás, 1340 m, 30TTN75, 11.VIII.1991, Col. Rius, MCNB; 1 ♂, Portillo de la Reina, 1239 m, 30TUN 46, 9.VII.2011, A. Blázquez leg.
LUGO: 1 ♂, Fogoso du Courel, 863 m, 29TPH52, 26.VII.2009, E. Fernández leg.; 3 ♂, Alto do Couto, Sierra do Courel, 1340 m, 29TPH51, 20–21.VII.1991, 4.VII.1993, E. Fernández leg.
NAVARRA: 1 ♂, Ollide, Sierra de Urbasa, 1072 m, 30TWN7846, 17.VII.2012, T. Latasa Leg.
OURENSE: 1 ♂, A Veiga, 1026 m, 29TPG68, 19.VI.2011, BV.
PALENCIA: 1 ♂, Cardaño de Abajo, 2224 m, 30TUN55, 10.VIII.2020, BV.; 1 ♂, Vidrieros, 2073 m, 30TUN65, 26.VII.2020, BV.; 1 ♂, Villabasta de Valdavia, 923 m, 30TUN61, 31.VII.2018, BV.; 1 ♂, Cardaño de Arriba, 2357 m, 30TUN56, 10.VIII.2012, BV.; 1 ♂, Velilla del Rio Carrión, 1200 m, 30TUN44, 4.V.2009, V. Redondo leg.; 1 ♂, Villanueva de Arriba, 1200 m, 30TUN53, 31.VII.1998, A. Blázquez leg.
SANTANDER: 1 ♂, Pujayo, 395 m, 30TVN17, 27.VI.2018, BV.; 1 ♂, Fuente Dé, 1877 m, 30TUN58, 25.VII.2013, BV.; 1 ♂, Horcadina de Covarrobres, 1908 m, 30TUN58, 8.VI.2011, BV.
SÓRIA: 1 ♂, Puerto de Santa Inés, 1770 m, 30TWM15, 18.VII.1976, V. Redondo leg.
TERUEL: 1 ♂, Puerto de Orihuela, Orihuela del Tremedal, 1700 m, 30TXK18, 13.VIII.1975, V. Redondo leg.
ZARAGOZA: 2 ♂, Moncayo, 1800 m, 30TWM92, 6.VIII.1976, V. Redondo leg.

3.2.2. Taxonomic Notes for Setina irrorella pseudoirrorella

The taxon S. i. irrorella, of Palearctic distribution, with a very widespread and quite common distribution, is present in much of France, Germany, Poland, the Balkans, Romania and southern Russia, east of Kamchatka, and Mongolia. It is also frequent in the northern part of the Alpes from Haute Provence to Haute Savoie, where it can reach high altitudes, even of more than 3000 metres. S. i. irrorella is the typonominal ssp. (Figure 1).
Regarding the taxon flavicans, the authors agree with what Leraut (2018) [6] states. The yellow colouration of the abdomen, the ground colour of the wings, and the reduction of the black drawings are frequent. This taxon, flavicans, is none other than the southern subspecies of S. irrorella and is present in France and throughout the Mediterranean basin (Figure 2).
According to Leraut (2018) [6], the form without yellow scales corresponds to the ssp. S. i. flavicans, but he considers that both forms are allopatric with a contact zone in some localities of the Pyrenees, and their genitals do not differ (Figure 2). Therefore, as they also have a black abdomen, the name S. i. pseudoirrorella applies as the Iberian subspecies of irrorella. (Figure 3).
The genitalia of S. flavicans show minimal or insignificant differences compared to those of S. irrorella in both sexes, well within the typical of the variability of the species and insufficient to consider them as different species. The genital structure of S. i. pseudoirrorella also does not show significant differences with the typonominal ssp. (Figure 4 and Figure 5). Taking these considerations into account, the taxon S. flavicans is a synonym of S. irrorella.
In general, in Spain, the vast majority of the specimens observed have a black abdomen, but in the eastern Pyrenees and Pre-Pyrenees, the two forms coexist: abdomen with yellow scales and black abdomen (Figure 3). Therefore, we could say that both subspecies are present, but being sympatric and sharing area and flight time, we consider that both forms belong to the ssp. pseudoirrorella, with no evidence of genetic difference between them.
Indeed, molecular analysis did not show genetic discrimination by the COI-5P marker between S. irrorella and S. flavicans. The phylogenetic trees (see Section 3.4) show that the specimens analysed are positioned within the same clade without any grouping between the specimens tagged as S. irrorella and S. flavicans. On the other hand, the irrorella–flavicans group forms a distinct clade, with a very high Bayesian support (PP = 1), including 63 sequences with low genetic distance within the group (T92-dist = 0.82%, see Section 3.4).

3.2.3. Biology of Setina irrorella pseudoirrorella

As for phenology, S. i. pseudoirrorella is a univoltine species, with adults flying in a single, long generation from May to September at altitudes between 400 and 2900 m. It is not ruled out that a partial second generation could occur at lower and warmer altitudes. It should be noted that these phenological data have been obtained by reviewing the material over a very wide range of years (1908–2023). Consequently, they are extremely significant, and it will be interesting to see whether the climate change affecting us causes future phenological changes. It has been noted that some colonies located at low altitudes have disappeared, probably due to the alteration of their habitat.
Ylla et al. (2010) [14] comment that their larvae feed on lichens and have been observed ingesting leaves of Saxifraga L.; however, most of the data on their biology remain unknown. Some of its stages are represented in Figure 6.

3.2.4. Distribution of Setina irrorella pseudoirrorella

S. i. pseudoirrorella is present throughout the Pyrenees, including its northern slopes, where it can be frequent. It is scarcer and in isolated colonies in the Pre-Pyrenees. In Catalonia, it reaches the Osona and Montseny regions but with colonies that are currently testimonial. It is also present in the Basque–Navarrese provinces, León, Burgos, and Palencia, and in the Cantabrian Mountains reaching Galicia, where it is less frequent and sympatric with S. cantabrica, maintaining isolated colonies. Further south, a probably small population is known in La Rioja and in the Iberian System.
Although a more southern distribution cannot be ruled out, we have not found any evidence of its presence in the south of Spain, as Gómez Bustillo (1979) [13] indicates in the province of Granada.
More than 500 records from all over Spain have been compiled and reviewed, which have allowed us to draw up a new, more up-to-date distribution map (Figure 7).

3.3. Setina cantabrica Freina and Witt, 1985. Stat. rev.

The original combination for Setina roscida cantabrica Freina and Witt, 1985, is as follows: Setina cantabrica Freina and Witt, 1985, Entomofauna 6 (16): 211 LT. Spain; Cantabrian Mts. Picos de Europa, Riaño, 1000 m. described based on 17 male specimens captured by Marten el 1.VII.1952.

3.3.1. Material Examined for Setina cantabrica

The follow material was examined:
ASTURIAS: 3 ♂, Camin de Mesa, Saliencia, Somiedo, 1400 m, 29TPH37, 18.VII.1996, E. Fernández leg.; 2 ♂, Lagos de Saliencia, Somiedo, 1800 m, 29TPH37, 27.VII.2003, R. Macià leg.; 2 ♂, Puerto de Leitariegos 1600 m, 29TQH16, 23.VII.1994, 13.VII.2000, E. Fernández leg.; 1 ♂, La Farropona, Somiedo, 1900, 29TQH37, 17.VII.2000, E. Fernández leg.; 1 ♂, Auteiro, Valle del Lago, 1340 m, 29TQH27, 19.VII.1999, E. Fernández leg.; 3 ♂, Sotres, 1250 m, 30TUN58, 23.VII.1991, 27.VII.1993, R. Macià leg.; 2♂, Lago de la Cueva, Sotres, 1800 m, 29TQH37, 23.VII.2003, R. Macià leg.; 1 ♂, Aixete, Tuiza, 1380 m, 30TTN66, 11.VII.2013, R. Macià leg.
LEON: 2 ♂, Villanueva de Arriba, 1200 m, 30TUN53, 31.VII.1998, R. Macià leg.; 1 ♂, Peña Ubiña, Torrebarrio, 1850 m, 30TTN56, 24.VII.2003, A. Blazquez leg.; 2 ♂, Roguera, Mina Ventana pequeña, 1450 m, 29TQH47, 22.VII.2003, A. Blazquez leg.; 1 ♂, Minas de Ventana, 1460 m, 29TQH47, 13.VII.2004, A. Blazquez leg.; 1 ♂, Portilla de la Reina, 1239 m, 30TUN46, 9.VII.2011, A. Blazquez leg.; 52 ♂, 3 ♀, Roguera, Mina Ventana, 1450 m, 30TTN5570, 17.VII.2002, 22.VII.2003, 13.VII.2004, 18.VII.2005, 7.VII.2013, 18.VII.2015, 2.VII.2020, 3.VII.2022, 5.VII.2023, R. Macià leg.; 25 ♂, 1 ♀, Caldas de Luna, 1284 m, 30TTN5767, 17.VII.2002, 22.VII.2003, 23.VII.2005, 8.VII.2012, 6.VII.2022, R. Macià leg.; 16 ♂, 1 ♀, Peña Ubiña, Torrebarrio, 1850 m, 30TTN5866, 14.VII.2004, 22.VII.2005, 9.VII.2011, R. Macià leg.; 31 ♂, Puerto de Piedrafita, 1540 m, 30TTN8866, 17.VII.2002, 22.VII.2003, 13.VII.2004, 18.VII.2005, 7.VII.2013, 18.VII.2015, 8.VII.2023, R. Macià leg.; 16 ♂, Puerto de la Mesa, 1850 m, 29TQH1163, 21.VII.1999, 16.VII.2000, 19.VII.2003, E. Fernández leg.; 1 ♂, Túnel de Alceo, Villamanin, 1495 m, 30TTN65, 28.VIII.2010, E. Fernández Leg.; 1 ♀, Villafeliz de Babia, 1180 m, 30TTN55, 10.VIII.2007, E. Fernández leg.; 3 ♂, Casa Mieres, Puerto de la Cubilla, 1400 m, 30TTN66, 11.VIII.2007, E. Fernández leg.; 2 ♂, Cubilla de Arbás, 1336 m, 30TTN75, 30.VII.2011, E. Fernández leg.; 1 ♂, Puerto de Pajares, Busdongo, 30TTN76, VII.1985, F. Murciego leg.; 1 ♂, Puerto de las Señales, 1740 m, 30TUN17, 15.VII.2000, E. Fernández leg.; 12 ♂, Puerto la Cubilla, 1683 m, 30TTN66, 26.VII.2002, 22.VII.2003, 23.VII.2005, 8.VII.2012, R. Macià leg.
LUGO: 2 ♂, Donis, Sierra de los Ancares, 1350 m, 29TPH74, 22.VII.1995, E. Fernández leg.; 1 ♂, Puente Vales, Vilarello, 1300 m, 29TPH63, 22.VII.1995, E. Fernández leg.
OURENSE: 16 ♂, 2 ♀, Finte da Cova, Foyo Grande, 1700 m, 29TPG88, 2.VII.1994, 29.VI.1996, 29.VII.1998, 17.VI.2000, 31.VII.2000, E. Fernández leg.; 3 ♂, 1 ♀ Carballeda de Valdeorras-Benuza, 1780 m, 29TPG88, 29-VII-1998, E. Fernández leg.
SANTANDER: 2 ♂ Fuente De, 1895 m, 30TUN56, 16.VII.1989, R. Magro leg.; 1 ♂, Espinama, 1890 m, 30TUN56, 8.VIII.1988, R. Magro leg.
PORTUGAL: 1 ♂, Serra das Meadas, Lamego, 1000 m, VI.1968, T. Monteiro leg.; 3 ♂, Souzanil, VI.1960, T. Monteiro leg.; 2 ♂, Tras-os-Montes, Zimão, VIII.1978, 7.VII.1981, Monteiro; 1 ♂, Tras-os-Montes, Pedras Salgadas, 7.1938, (M.F.V. Corley).

3.3.2. Taxonomic Notes for Setina cantabrica

The authors are of the opinion that S. cantabrica should maintain the status of good species based on the following arguments:
Despite the molecular studies indicating that the genetic divergence of the COI marker between S. roscida and S. cantabrica is minimal, which justifies by itself specific separation (T92-dist = 0.92%, see Table 2), the sequences of both species are split in two different branches, with high Bayesian support (PP = 1, see Section 3.4) and with very low divergence within the groups (T92-dist = 0.10% for the cantabrica group and 0.37% for the roscida group (see Section 3.4, Table 2)). Only two specimens downloaded from public data and tagged as S. roscida are included in the cantabrica group, and in this case, it can be said that they correspond to specimens captured in the distribution area of S. cantabrica. On the other hand, all the specimens included in the roscida group correspond to specimens captured outside the Iberian Peninsula (see Section 3.4).
At a morphological level, there are differences both externally and in genitalia that make us consider it a good species and different from S. roscida. In this case, despite sharing a similar wing pattern with roscida, the wingspan of the male imagos of cantabrica is greater. The wingspan is 28.66 ± 1.37 (=49) mm for S. cantabrica and 23.54 ± 1.46 (=22) mm for S. roscida (Figure 8 and Figure 9). It also presents evident differences in the antennae of the males.
As for the antennae (Figure 10), these are ciliated or fasciculated in both species, characterised by the presence of a large number of cilia located on the ventral side of the antenna along its entire length, although it is in the segments of the central part, where this hairiness is most evident. The differences between the two species are found in a greater density of cilia in S. cantabrica and in the shape of the segments in lateral view, being more trapezoidal in S. cantabrica and more rounded in S. roscida. In the ventral view, the presence of two types of cilia is evident in both species, some shorter and more abundant (at a rate of about 10–15 per segment) and others longer at a rate of 2 per segment and twice as long, arranged in their central part.
Regarding geographical separation, S. r. roscida is native to western Europe (southwest France, south Germany, Switzerland, Austria, the Balkans, north Italy, south Romania, and Bulgaria), while S. r. kuhlweini colonises southern Sweden, northeast Germany, and the Baltic countries of Kazakhstan. In the case of S. cantabrica, it is only found in the Cantabrian Mountains and in northern Portugal. Therefore, geographical isolation occurs between both species, which facilitates their evolutionary divergence.
Despite presenting similar male genitalia, compared to S. roscida (Figure 10B), S. cantabrica (Figure 10A) has a reduced, rounded ampullary process situated quite towards the centre of the valve; a generally long, slender, and pointed saccular process extending beyond the terminal part of the valve; and a long, thin, and pointed vinculum, triangular in both cases, but smaller in the case of S. cantabrica.
Furthermore, the phallus of S. roscida generally has 1–2 large cornuti at its terminal end, while S. roscida has 3–5 large cornuti in the same area, with the size of the aedeagus being larger in the latter case. The differences in the female genitalia are more representative, with S. cantabrica presenting a wider ostium than S. roscida, which makes the geometry of the antrum also different. The appendix bursae of S. cantabrica (Figure 11A) is more or less symmetrical, wider in the median part, rather drop-shaped with rounded proximolateral margins, while in S. roscida, it is triangularly asymmetrical and rounded, is wider proximally, and has a much wider union with the corpus bursae. (Figure 11B).

3.3.3. Biology of Setina cantabrica

S. cantabrica is a monovoltine species; adults emerge in a single generation in June, July, and early August in colonies that can sometimes be abundant. It prefers open biotopes, mainly with calcareous substrate, located between 800 and 2100 m in altitude, with adults flying during the day, with special intensity from early morning to midday, coming to artificial light less frequently. Corley (2013) [16] mentions that most of the affected Portuguese areas have acidic rocks, and the altitude range is smaller.
There are no reliable data on the host plants of the caterpillars nor on most other aspects of biology. Some of their stages are represented in Figure 12.

3.3.4. Distribution of Setina cantabrica

This is an Iberian endemic species, occupying a wide area of the Cantabrian Mountains, from Riaño (León) to the Sierra de los Ancares in the province of Lugo and even further south in the Trevinca massif in Ourense [4], reaching as far as northern Portugal [16]. In some localities, it is sympatric with S. i. pseudoirrorella.
The updated distribution of S. cantabrica is presented on a 10 × 10 km2 UTM grid map (Figure 13). Most of the records reviewed, a total of about 300, correspond to the area of the Cantabrian Mountains in the provinces of León, Cantabria, and Asturias, reaching as far as Galicia and northern Portugal.
The Portuguese records extend the distribution area of S. cantabrica to northern Portugal. The specimens from southern Portugal in the Monteiro collection, from Lagoa on the southern coast of the Algarve, as Corley (2013) comments [16], are very likely of doubtful origin. They are shown on the map in yellow, indicating that they have not been verified.

3.4. DNA Analysis

The Bayesian inference analysis of the DNA barcoding marker COI-5P for the specimens included in this study (listed in Table 1) rvealed two well-supported clades: one comprising the roscida–cantabrica sequences and the other including the irrorella–flavicans specimens. This result is illustrated in the combined analysis with public records (Figure 14), which confirms the highly supported separation between S. roscida and S. cantabrica (PP = 1), while no clear distinction is evident within the irrorella–flavicans group. As shown in Table 2, the inter-specific distances are around 2%, except for the divergence between S. roscida and S. cantabrica, which is substantially lower than the rest.
The remaining group detected through Bayesian inference corresponds to S. aurita, a species not included in our dataset but often difficult to distinguish morphologically from S. irrorella.
Table 2. Estimates of evolutionary divergence on sequence pairs between groups for the marker COI-5P. The number of base substitutions per site from the mean of all sequence pairs between species is shown in percentages with standard error estimates under each value. Analyses were performed using the Tamura 3-parameter model [11], and rate variation between sites was modelled with a gamma distribution (shape parameter = 1). The analysis involved 121 nucleotide sequences with a total of 658 positions. The intraspecific (within group) number of base substitutions obtained under the same conditions is shown by the diagonal line (bold). Light green shaded cells highlight T92 distances <1.5%, and orange shaded cells indicate T92 distances >5%. The comparison is made based on sequences grouped in the same clade or group.
Table 2. Estimates of evolutionary divergence on sequence pairs between groups for the marker COI-5P. The number of base substitutions per site from the mean of all sequence pairs between species is shown in percentages with standard error estimates under each value. Analyses were performed using the Tamura 3-parameter model [11], and rate variation between sites was modelled with a gamma distribution (shape parameter = 1). The analysis involved 121 nucleotide sequences with a total of 658 positions. The intraspecific (within group) number of base substitutions obtained under the same conditions is shown by the diagonal line (bold). Light green shaded cells highlight T92 distances <1.5%, and orange shaded cells indicate T92 distances >5%. The comparison is made based on sequences grouped in the same clade or group.
COI-5P T92 DistanceIrrorella-flavicans GroupAurita GroupCantabrica GroupRoscida Group
Irrorella-flavicans Group (n = 60)0.82%
±0.16 SE
Aurita Group(n = 33)2.16%
±0.46 SE		1.30%
±0.27 SE
Cantabrica Group(n = 11)2.60%
±0.60 SE		2.29%
±0.52 SE		0.10%
±0.08 SE
Roscida Group(n = 15)2.78%
±0.61 SE		2.37%
±0.51 SE		0.56%
±0.23 SE		0.37%
±0.14 SE
Nudaria mundana12.44%
±1.66 SE		11.86%
±1.59 SE		12.74%
±1.71 SE		12.74%
±1.68 SE

4. Discussion

Due to its complexity, the taxonomic status of the Setina genus has been a subject of considerable debate. Our findings indicate that both morphological and genetic characteristics of S. irrorella and S. flavicans do not justify their specific separation. In terms of genitalia, S. flavicans show minimal differences compared to those of S. irrorella in both sexes, which fall within the typical variability of the species. This lack of significant differentiation in genital structure suggests that S. flavicans should be considered a synonym of S. irrorella. Further supporting this, molecular studies using the COI-5P marker show no genetic separation between S. irrorella and S. flavicans. The specimens analysed are positioned within the same clade, indicating no distinct genetic grouping. The extensive morphological studies combined with the geographic distribution of S. irrorella indicates that the taxon flavicans represents the southern subspecies S. i. pseudoirrorella, which have a contact zone in the Pyrenees with the European form S. i. irrorella.
Regarding S. cantabrica, our results support its status as a distinct species from S. roscida. Although the genetic divergence of the COI marker between S. roscida and S. cantabrica is reduced, the sequences of both species form separate branches with high Bayesian support. This genetic distinction is corroborated by morphological differences, particularly in the wingspan and antennae structure of the males. Among other differences, S. cantabrica males have a greater wingspan and a higher density of cilia on the antennae compared to S. roscida, with significant differences in the shape of the antenna segments. Additionally, S. cantabrica and S. roscida exhibit geographical separation, with S. cantabrica only found in the Iberian Peninsula and S. roscida outside this region.
Considering all this evidence, the authors consider it justified to maintain S. cantabrica as a valid species while resolving the irrorella–flavicans binomial into the subspecies S. irrorella pseudoirrorella. Further studies, particularly involving molecular data, will still be essential to resolve remaining uncertainties and refine the classification of these taxa.
It should be noted that the original description by Freina and Witt (1985) [2] is listed as “Holotypus ♂: Kantabrisches Geb., Picos Europa, Riano, 1000 m, 1.7.1952, MÄRTEN lg., Franz DANIEL, München (in Museum WITT, München) 16 ♂♂ idem. (mit Gen.Präp.WITT Nr.2621, 2622, 2623 und TARMANN A 45) (all in Museum WITT, München, ♀ unbekannt”. The female as unknown, and the same is stated in Freina and Witt (1987) [3]. We are surprised that in Witt and Ronkay (2011) [1], the female genitalia is represented as “295. Setina cantabrica, Spain Asturia MWM2622 Paratype”. It is assumed that this is an error; the original identification “MWM2622 Paratype” corresponded to a male, the female was unknown, and it was represented for the first time in Fernández Vidal et al. (2003) [4].

Author Contributions

Investigation, R.M., J.Y., G.M. and J.B. All authors have read and agreed to the published version of the manuscript.

Funding

This study received no external funding.

Data Availability Statement

Molecular data has been compiled in the public dataset and is available at BOLD (http://v4.boldsystems.org) under the access code SETI.

Acknowledgments

This work would not have been possible without the collaboration and assistance of the following people, who have provided us with data from their research and collections for the preparation of the distribution maps: Eliseo Fernández Vidal, Víctor Redondo, Rafael Magro, Martí Rondós, Josep Muñoz, Arcadi Cervelló, Pere Passola, Eduard Guzman, Albert Xaus, Jordi Dantart, Francesc Vallhonrat, Diego Fernández, Josep Planes, Antonio Correas, Tomás Latasa, Éric Drouet, and Ángel Blázquez. In particular, we would like to thank Javier Gastón for his invaluable help and selfless collaboration in the preparation and discussion of the genitalia; to Berta Caballero-López and Irene Lobato-Vila, curators of the arthropod collection at the Barcelona Museum of Natural Sciences (MCNB), who provided us with all kinds of facilities to review the deposited material; and to Emma Asensio, librarian at the Barcelona Museum of Natural Sciences (MCNB), for her help in consulting the Museum’s bibliographic collections. Josep Martí provided us with the complete data corresponding to the images stored in Virtual Biodiversity. We also appreciate the comments of the anonymous reviewers, which have helped us improve the work.

Conflicts of Interest

The authors declare no conflicts of interest.

References

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Figure 1. S. i. irrorella. (A,B) ♂♀ Col de Glandon, Savoie (Francia), 1807 m, 8.VII.2004, R. Macià leg.; (C) ♂ Val Tourmanché, Aosta (Italia), 2250 m, 1.VIII.1999, A. Bonora leg.; (D) ♀ Schnalstal, South Tirol (Italia), 1800 m, 31.VII.1978, O. Achenbach leg.
Figure 1. S. i. irrorella. (A,B) ♂♀ Col de Glandon, Savoie (Francia), 1807 m, 8.VII.2004, R. Macià leg.; (C) ♂ Val Tourmanché, Aosta (Italia), 2250 m, 1.VIII.1999, A. Bonora leg.; (D) ♀ Schnalstal, South Tirol (Italia), 1800 m, 31.VII.1978, O. Achenbach leg.
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Figure 2. S. i. flavicans. (A,B) ♂♀ Marcoux, Alpes, Alta Provenza (Francia) 850 m. 15.VII.2006, R. Macià leg.; (C) ♂ Col de Lombarde, Alpes marítimos (Francia), 2800 m, 13.VIII.1996, R. Macià leg.; (D) ♀ Col de Lombarde, Alpes marítimos (Francia), 2800 m, 19.VII.2007, R. Macià leg.
Figure 2. S. i. flavicans. (A,B) ♂♀ Marcoux, Alpes, Alta Provenza (Francia) 850 m. 15.VII.2006, R. Macià leg.; (C) ♂ Col de Lombarde, Alpes marítimos (Francia), 2800 m, 13.VIII.1996, R. Macià leg.; (D) ♀ Col de Lombarde, Alpes marítimos (Francia), 2800 m, 19.VII.2007, R. Macià leg.
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Figure 3. S. i. pseudoirrorella. Different Iberic locations (yellow and black abdomen). (A) ♂ Caldas de Luna (León), 1430 m, 23.VII.2005, R. Macià leg.; (B) ♀ Caldas de Luna (León), 1430 m, 23.VII.2005, R. Macià leg.; (C) ♂ El Cerro, Gredilla la Polera (Burgos), 980 m, 21.VII.2017, R. Macià leg.; (D) ♀ El Cerro, Gredilla la Polera (Burgos), 980 m, 21.VII.2017, R. Macià leg.; (E) ♂ Loma del Rei, Quintanilla Sobresierra (Burgos), 1003 m, 13.VII.2012, R. Macià leg.; (F) ♂ Lago Urdiceto, Bielsa (Huesca), 2000 m, 17.VII.1995, R. Macià leg.; (G) ♀ Lago Urdiceto, Bielsa (Huesca), 2000 m, 17.VII.1995, R. Macià leg.; (H) ♂ Banhs de Tredós, Val d’Aran (Lleida), 1750 m, 12.VII.2007, R. Macià leg.; (I) ♀ Banhs de Tredós, Val d’Aran (Lleida), 1750 m, 12.VII.2007, R. Macià leg.; (J) ♂ Bosc Nera, Vanhs de Tredós, Val d’Aran (Lleida) 1800 m, 26.VII.2008, R. Macià leg.; (K) ♂. Vallter, Ripollès (Girona) 2100 m. 9.VIII.2023, R. Macià leg.; (L) ♀ Vallter, Ripollès (Girona) 2100 m. 9.VIII.2023, R. Macià leg.; (M) ♂ Vallter, Ripollès (Girona) 2100 m. 9.VIII.2023, R. Macià leg.; (N) ♂ Font Home Mort, Queralbs (Girona), 1564 m, 29.VII.2006, R. Macià leg.; (O) ♂ P.N. Castell de Montesquiu, Montesquiu (Barcelona) 637 m. 21.IX.2007, R. Macià leg.; (P) ♀ P.N. Castell de Montesquiu, Montesquiu (Barcelona) 637 m. 21.IX.2007, R. Macià leg.
Figure 3. S. i. pseudoirrorella. Different Iberic locations (yellow and black abdomen). (A) ♂ Caldas de Luna (León), 1430 m, 23.VII.2005, R. Macià leg.; (B) ♀ Caldas de Luna (León), 1430 m, 23.VII.2005, R. Macià leg.; (C) ♂ El Cerro, Gredilla la Polera (Burgos), 980 m, 21.VII.2017, R. Macià leg.; (D) ♀ El Cerro, Gredilla la Polera (Burgos), 980 m, 21.VII.2017, R. Macià leg.; (E) ♂ Loma del Rei, Quintanilla Sobresierra (Burgos), 1003 m, 13.VII.2012, R. Macià leg.; (F) ♂ Lago Urdiceto, Bielsa (Huesca), 2000 m, 17.VII.1995, R. Macià leg.; (G) ♀ Lago Urdiceto, Bielsa (Huesca), 2000 m, 17.VII.1995, R. Macià leg.; (H) ♂ Banhs de Tredós, Val d’Aran (Lleida), 1750 m, 12.VII.2007, R. Macià leg.; (I) ♀ Banhs de Tredós, Val d’Aran (Lleida), 1750 m, 12.VII.2007, R. Macià leg.; (J) ♂ Bosc Nera, Vanhs de Tredós, Val d’Aran (Lleida) 1800 m, 26.VII.2008, R. Macià leg.; (K) ♂. Vallter, Ripollès (Girona) 2100 m. 9.VIII.2023, R. Macià leg.; (L) ♀ Vallter, Ripollès (Girona) 2100 m. 9.VIII.2023, R. Macià leg.; (M) ♂ Vallter, Ripollès (Girona) 2100 m. 9.VIII.2023, R. Macià leg.; (N) ♂ Font Home Mort, Queralbs (Girona), 1564 m, 29.VII.2006, R. Macià leg.; (O) ♂ P.N. Castell de Montesquiu, Montesquiu (Barcelona) 637 m. 21.IX.2007, R. Macià leg.; (P) ♀ P.N. Castell de Montesquiu, Montesquiu (Barcelona) 637 m. 21.IX.2007, R. Macià leg.
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Figure 4. (A) S. i. irrorella; male and female genitalia. Sölden, Tirol (Austria) 2800 m 21.VII.1997, 9460JG, 9463JG. (B) S. i. flavicans; male and female genitalia. Col de Lombarde, Alpes marítimos (Francia) 2300 m 18.VII.2007,9458JG, 9462JG.
Figure 4. (A) S. i. irrorella; male and female genitalia. Sölden, Tirol (Austria) 2800 m 21.VII.1997, 9460JG, 9463JG. (B) S. i. flavicans; male and female genitalia. Col de Lombarde, Alpes marítimos (Francia) 2300 m 18.VII.2007,9458JG, 9462JG.
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Figure 5. S. i. pseudoirrorella. (A) Male genitalia. Bosc Nera, Vanhs de Tredós, Val d’Aran (Lleida) 1740 m. 26.VII.2015, 2862RM; (B) male genitalia. Mina Ventana, Roguera, Torrebarrio (León) 1450 m. 27.VII.2002, JY2641; (C) female genitalia, Lago Urdiceto, Bielsa (Huesca) 2000 m. 17.VII.95, JY2740; (D) male genitalia. Vallter, Ripollès (Girona) 2200 m. 10.VIII.2024, 2859RM.
Figure 5. S. i. pseudoirrorella. (A) Male genitalia. Bosc Nera, Vanhs de Tredós, Val d’Aran (Lleida) 1740 m. 26.VII.2015, 2862RM; (B) male genitalia. Mina Ventana, Roguera, Torrebarrio (León) 1450 m. 27.VII.2002, JY2641; (C) female genitalia, Lago Urdiceto, Bielsa (Huesca) 2000 m. 17.VII.95, JY2740; (D) male genitalia. Vallter, Ripollès (Girona) 2200 m. 10.VIII.2024, 2859RM.
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Figure 6. Caterpillar, chrysalis, and adult of S. i. pseudoirrorella.
Figure 6. Caterpillar, chrysalis, and adult of S. i. pseudoirrorella.
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Figure 7. S. i. pseudoirrorella. A distribution map according to data reviewed from the Iberian Peninsula.
Figure 7. S. i. pseudoirrorella. A distribution map according to data reviewed from the Iberian Peninsula.
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Figure 8. S. cantabrica: (A,B) ♂♀ Mina Ventana, Roguera, Torrebarrio (León), 1450 m, 15.VII.2010, R. Macià leg.; (C,D) ♂♂ Mina Ventana, Roguera, Torrebarrio (León), 1450 m, 13.VII.2004, R. Macià leg.; (E,F) ♂♀ Peña Ubiña, Torrebarrio (León), 1850 m, 24.VII.2003, R. Macià Leg.; (G,H) ♂♀ Puerto de Piedrafita, Cármenes, (León), 1683 m, 14.VII.2004. R. Macià leg.
Figure 8. S. cantabrica: (A,B) ♂♀ Mina Ventana, Roguera, Torrebarrio (León), 1450 m, 15.VII.2010, R. Macià leg.; (C,D) ♂♂ Mina Ventana, Roguera, Torrebarrio (León), 1450 m, 13.VII.2004, R. Macià leg.; (E,F) ♂♀ Peña Ubiña, Torrebarrio (León), 1850 m, 24.VII.2003, R. Macià Leg.; (G,H) ♂♀ Puerto de Piedrafita, Cármenes, (León), 1683 m, 14.VII.2004. R. Macià leg.
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Figure 9. S. roscida: (A,B) ♂♀ Tanas, Trentino-Alto Adigio, Tirol del Sud (Italia) 850 m. 28.VI.2004; (C,D) ♂♀ Solü, Vezprém región (Hungría), 10.V.2008, J. Darebnik leg.; (E) ♂ Möchelmossen (Suecia), 23.V.1994, K. Nupponen leg.; (F) ♂ Csakbereny (Hungría), 17.V.2008, Vorlind leg.; (G) ♀ Altenbach (Austria), 30.V.1997, I. Richter leg.; (H) ♂ Csakbereny, Bucka-hegy (Hungría), 8.V.2003, I. Richter leg.
Figure 9. S. roscida: (A,B) ♂♀ Tanas, Trentino-Alto Adigio, Tirol del Sud (Italia) 850 m. 28.VI.2004; (C,D) ♂♀ Solü, Vezprém región (Hungría), 10.V.2008, J. Darebnik leg.; (E) ♂ Möchelmossen (Suecia), 23.V.1994, K. Nupponen leg.; (F) ♂ Csakbereny (Hungría), 17.V.2008, Vorlind leg.; (G) ♀ Altenbach (Austria), 30.V.1997, I. Richter leg.; (H) ♂ Csakbereny, Bucka-hegy (Hungría), 8.V.2003, I. Richter leg.
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Figure 10. Structures of the male antennae: (A,B) S. roscida, ventral and lateral view. (C,D) S. cantabrica, ventral and lateral view.
Figure 10. Structures of the male antennae: (A,B) S. roscida, ventral and lateral view. (C,D) S. cantabrica, ventral and lateral view.
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Figure 11. (A) S. cantabrica: male genitalia. Puerto de Piedrafita (León) 1450 m. 10.VIII.2007, 3138 JG; female genitalia. Mina Ventana, Roguera, Torrebarrio (León) 1450 m. 27.VII.2002, JY2739. (B) Setina roscida: male genitalia. Tanas, south Tirol (Italy) 800 m. 28.VI.1994, 9461JG; female genitalia. Gran Sasso, Apenines (Italia) 2500 m. 12.VII.2009, 9456JG.
Figure 11. (A) S. cantabrica: male genitalia. Puerto de Piedrafita (León) 1450 m. 10.VIII.2007, 3138 JG; female genitalia. Mina Ventana, Roguera, Torrebarrio (León) 1450 m. 27.VII.2002, JY2739. (B) Setina roscida: male genitalia. Tanas, south Tirol (Italy) 800 m. 28.VI.1994, 9461JG; female genitalia. Gran Sasso, Apenines (Italia) 2500 m. 12.VII.2009, 9456JG.
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Figure 12. Caterpillar, chrysalis, and adult of S. cantabrica.
Figure 12. Caterpillar, chrysalis, and adult of S. cantabrica.
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Figure 13. S. cantabrica. The distribution map according to data reviewed from the Iberian Peninsula.
Figure 13. S. cantabrica. The distribution map according to data reviewed from the Iberian Peninsula.
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Figure 14. The phylogenetic tree representation of the extended dataset that includes the sequences obtained in this study together with those published in BOLD under the genus Setina. Posterior probability values are shown next to the nodes. The main clades or groups have been highlighted in colours (blue: irrorella–flavicans group; green: aurita group; yellow: cantabrica group; red: roscida group). The specimens downloaded from BOLD are shown in black font, and those sequenced in the framework of this work in colour font. For clarity, the branching of the outgroup (Nudaria mundana) has been shortened.
Figure 14. The phylogenetic tree representation of the extended dataset that includes the sequences obtained in this study together with those published in BOLD under the genus Setina. Posterior probability values are shown next to the nodes. The main clades or groups have been highlighted in colours (blue: irrorella–flavicans group; green: aurita group; yellow: cantabrica group; red: roscida group). The specimens downloaded from BOLD are shown in black font, and those sequenced in the framework of this work in colour font. For clarity, the branching of the outgroup (Nudaria mundana) has been shortened.
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Table 1. A list of specimens investigated by genetic analysis of the COI-5P marker.
Table 1. A list of specimens investigated by genetic analysis of the COI-5P marker.
Sample IDTagged SpeciesCollectorCollection DateLocationGenbank Access Code.
UVIC_JBM2302S. cantabricaR. Macià13 July 2024Spain, León,PV173886
UVIC_JBM2305S. cantabricaR. Macià5 July 2022Spain, LeónPV173887
UVIC_JBM2306S. cantabricaR. Macià5 July 2022Spain, LeónPV173889
UVIC_JBM2317S. cantabricaR. Macià5 July 2022Spain, LeónPV173885
UVIC_MBT412S. cantabricaR. Macià3 July 2024Spain, LeónPV173890
UVIC_MBT413S. cantabricaR. Macià3 July 2024Spain, LeónPV173891
UVIC_MBT414S. cantabricaR. Macià3 July 2024Spain, LeónPV173892
UVIC_MBT416S. cantabricaR. Macià3 July 2024Spain, LeónPV173888
UVIC_JBM2301S. flavicansR. Macià12 July 2007Spain, LleidaPV173907
UVIC_JBM2309S. flavicansR. Macià8 August 2021Spain, LeónPV173899
UVIC_JBM2310S. flavicansR. Macià14 July 2020Spain, LleidaPV173900
UVIC_JBM2316S. flavicansR. Macià14 July 2020Spain, LleidaPV173901
UVIC_MBT402S. flavicansR. Macià10 August 2024Spain, GironaPV173902
UVIC_MBT403S. flavicansR. Macià25 June 2024Spain, BurgosPV173903
UVIC_MBT404S. flavicansR. Macià10 August 2024Spain, GironaPV173904
UVIC_MBT405S. flavicansR. Macià3 July 2024Spain, LeónPV173905
UVIC_MBT406S. flavicansR. Macià3 July 2024Spain, LeónPV173893
UVIC_MBT407S. flavicansR. Macià16 July 2024Spain, LleidaPV173894
UVIC_MBT408S. flavicansR. Macià14 July 2020Spain, LleidaPV173895
UVIC_MBT409S. flavicansR. Macià16 July 2024Spain, LleidaPV173896
UVIC_MBT411S. flavicansR. Macià25 June 2024Spain, BurgosPV173897
UVIC_RMV12S. flavicansHerzet12 July 2004FrancePV173908
UVIC_RMV13S. flavicansR. Macià18 July 2012Spain, LleidaPV173909
UVIC_RMV14S. flavicansR. Macià15 September 2007Spain, BarcelonaPV173910
UVIC_RMV15S. flavicansR. Macià13 July 2012Spain, BurgosPV173911
UVIC_RMV16S. flavicansR. Macià8 August 2021Spain, LeónPV173912
UVIC_RMV17S. flavicansR. Macià6 July 2017Spain, LeónPV173906
UVIC_RMV18S. irrorellaR. Macià21 July 1997AustriaPV173898
UVIC_JBM2314S. roscidaR. Macià17 July 2019ItalyPV173913
UVIC_MBT417S. roscidaCsakbereny8 May 2003HungaryPV173914
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MDPI and ACS Style

Macià, R.; Ylla, J.; Masó, G.; Bau, J. On Setina irrorella pseudoirrorella Freina and Witt, 1985, and Setina cantabrica Freina and Witt, 1985, in Spain and Portugal: Taxonomic Status, Distribution, Habitat, and Molecular Genetics (Lepidoptera: Erebidae: Arctiinae: Lithosiini). Taxonomy 2025, 5, 23. https://doi.org/10.3390/taxonomy5020023

AMA Style

Macià R, Ylla J, Masó G, Bau J. On Setina irrorella pseudoirrorella Freina and Witt, 1985, and Setina cantabrica Freina and Witt, 1985, in Spain and Portugal: Taxonomic Status, Distribution, Habitat, and Molecular Genetics (Lepidoptera: Erebidae: Arctiinae: Lithosiini). Taxonomy. 2025; 5(2):23. https://doi.org/10.3390/taxonomy5020023

Chicago/Turabian Style

Macià, Ramon, Josep Ylla, Guillem Masó, and Josep Bau. 2025. "On Setina irrorella pseudoirrorella Freina and Witt, 1985, and Setina cantabrica Freina and Witt, 1985, in Spain and Portugal: Taxonomic Status, Distribution, Habitat, and Molecular Genetics (Lepidoptera: Erebidae: Arctiinae: Lithosiini)" Taxonomy 5, no. 2: 23. https://doi.org/10.3390/taxonomy5020023

APA Style

Macià, R., Ylla, J., Masó, G., & Bau, J. (2025). On Setina irrorella pseudoirrorella Freina and Witt, 1985, and Setina cantabrica Freina and Witt, 1985, in Spain and Portugal: Taxonomic Status, Distribution, Habitat, and Molecular Genetics (Lepidoptera: Erebidae: Arctiinae: Lithosiini). Taxonomy, 5(2), 23. https://doi.org/10.3390/taxonomy5020023

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