Integrative taxonomy reveals overlooked cryptic diversity in the conifer feeding Batrachedra pinicolella (Zeller, 1839) (Lepidoptera, Batrachedridae) Author Berggren, Kai Bravann terrasse 21, NO- 4624 Kristiansand, Norway Author Aarvik, Leif https://orcid.org/0000-0002-0112-8837 Natural History Museum, University of Oslo, P. O. Box 1172 Blindern, NO- 0318 Oslo, Norway Author Huemer, Peter https://orcid.org/0000-0002-0630-545X Tiroler Landesmuseen Betriebgsges. m. b. H., Sammlungs- und Forschungszentrum, Naturwissenschaftliche Sammlungen, Krajnc-Str. 1, A- 6060 Hall in Tirol, Austria p.huemer@tiroler-landesmuseen.at Author Lee, Kyung Min https://orcid.org/0000-0001-9055-1073 Zoology Unit, Finnish Museum of Natural History, University of Helsinki, Finland & Ecology and Genetics Research Unit, University of Oulu, Finland Author Mutanen, Marko https://orcid.org/0000-0003-4464-6308 Ecology and Genetics Research Unit, University of Oulu, Finland text ZooKeys 2022 2022-02-08 1085 165 182 http://dx.doi.org/10.3897/zookeys.1085.76853 journal article http://dx.doi.org/10.3897/zookeys.1085.76853 1313-2970-1085-165 0ED94C608235413B99F5F3D52F2E66D5 D0FD306A27FF562BBDB2A2512B0657AC Batrachedra confusella sp. nov. Figs 3 , 5 , 6 , 9 Material. Holotype 1♂ O , Moss: Rygge, Sildebauen, 59.3268°N ; 10.7101°E ; 9.vii.1980 ; L. Aarvik leg.; NHMO prep. 3943 (NHMO). Paratypes Finland 1♂ A, Sund ; 14.vii.2007 ; M. Mutanen leg.; L. Aarvik prep. 2015.022; BOLD sample ID: MM22065; ZMUO . 1♂ PPe, Oulu; 7.vii.2011 ; M. Mutanen leg.; L. Aarvik prep. 2015.023; BOLD sample ID: MM21051; ZMUO. 1♀ U, Kirkkonummi; 11.vii.2007 ; M. Mutanen leg.; L. Aarvik prep. 2015.025; BOLD sample ID: MM06674; ZMUO. Norway 1♂ AAY, Arendal : Havsoy ; 10.vii.2015 ; K. Berggren leg.; BOLD sample ID: NHMO-DAR-12592; KBE . 1♀ AAY, Arendal : Tromoy , Skottjern ; 22.vii.1983 ; K. Berggren leg.; KBE prep. 13534; KBE ; 1♂ , same locality and date; S. Svendsen leg.; NHMO prep. 2833; NHMO . 1♂ AAY, Arendal : Siring ; 25-30.vi.2002 ; S.A. Bakke leg., NHMO prep. 3936; NHMO . 1♀ AAY, Grimstad : Som ; 8.vii.2017 ; K. Berggren leg., BOLD sample ID: NHMO-DAR-13683; KBE . 1♀ AAY, Grimstad : Som ; 24.vii.2017 ; K. Berggren leg.; BOLD sample ID: NHMO-DAR-14041; KB . 1♂ AAY, Lillesand : Lillesand ; 7.vii.1984 ; K. Berggren leg.; KBE prep. 13515; KBE . 1♂ AAY, Lillesand : Svinoya ; 29.vii.2005 , K. Berggren leg.; KBE prep. 13520; KBE . 1♂ AAY Tvedestrand : Lyngor , Sonnerstrand ; 30.vi-4.vii.2013 ; K. Berggren & B. Johansen leg.; KBE prep. 13524; KBE . 1♂ AK, Asker : Bronnoya ; 11.vii.1981 ; K. Berggren leg.; KBE prep. 13528; KBE . 1♀ AK, Baerum : Ostoya ; 10.vii.1984 ; L. Aarvik leg.; NHMO prep. 2887; NHMO . 1♂ AK, Oslo : Bleikoya ; 14.vii.2009 ; K. Berggren & A. Endrestol leg.; BOLD sample ID: NHMO-DAR-5192; KBE . 1♀ AK, Oslo : Bygdoy , Rodelokka ; 4-19.vii.2016 ; A. Endrestol & K. Berggren leg.; KBE prep. 13536; KBE . 1♂ AK, Oslo : Bygdoy , Rodelokka ; 4-19.vii.2016 ; A. Endrestol & K. Berggren leg.; KBE prep. 13532; KBE . 1♂ AK, Oslo : Ekebergskraninga ; 25.vi.2008 ; L. Aarvik leg.; NHMO prep. 2835; BOLD sample ID: NHMO-08107; NHMO . 1♀ AK, As : As ; 30.vii.1982 ; L. Aarvik leg.; L. Aarvik prep. 2725; NHMO . 1♀ HES, Asnes : Sonsterud ; 16-25.vii.1998 ; L. Aarvik & A. Bakke leg.; NHMO prep. 2832; NHMO . 1♀ TEY, Kragero : Jomfruland , Oytangen ; 16.vii.2003 ; L. Aarvik leg.; NHMO prep. 3937; NHMO . 1♂ TEY, Porsgrunn : Sandoya ; 10.vii.2003 ; R. Voith leg.; KBE prep. 13521; KBE . 1♂ VAY, Kristiansand : Augland ; 8.vii.1985 ; K. Berggren leg.; KBE prep. 13529; KBE . 1♂ VAY, Kristiansand : Bravann ; 14.vii.2015 ; K. Berggren leg., BOLD sample ID: NHMO-DAR-12591; KBE . 1♂ VAY, Kristiansand : Bravann ; 17.vii.2017 ; K. Berggren leg., BOLD sample ID: NHMO-DAR-14027; KBE . 1♀ VAY, Kristiansand : Bravann ; 10.viii.2012 ; K. Berggren leg.; KBE prep. 13537; KBE . 1♀ VAY, Kristiansand : Bravann ; 26.viii.2016 ; K. Berggren leg.; KBE prep. 13538; KBE . 1♂ VAY, Kristiansand : Bravann ; 26.viii.2016 ; K. Berggren leg.; BOLD sample ID: NHMO-DAR-12591; KBE prep. 13538; KBE . 1♂ VAY, Kristiansand : Flekkeroy , Belteviga ; 28.vii.1999 , K. Berggren leg.; KBE prep. 6141; KBE . 1♀ VAY, Kristiansand : Flekkeroy , Belteviga ; 16-23.vii.2000 , K. Berggren leg.; KBE prep. 6196; KBE . 1♀ VAY, Kristiansand : Nedre Timenes ; 7-14.vii.2001 , K. Berggren leg.; KBE prep. 101696; KBE . 1♂ VAY, Kristiansand : Nedre Timenes ; 28.vii.2015 , K. Berggren leg.; KBE prep. 13533; KBE . 1♀ VAY, Kristiansand : Nedre Timenes ; 31.vii.2017 , K. Berggren leg.; KBE prep. 13535; KBE . 1♂ VAY, Kristiansand : Ostre Randoy ; 9.vi.2006 , K. Berggren leg.; KBE prep. 13530; KBE . 1♂ VAY, Kristiansand : Skalevik ; 12.vii.2019 ; K. Berggren leg., BOLD sample ID: KBE-2019077; KBE . 1♂ VAY, Kristiansand : Stangenes ; 4.vii.1981 ; S. Svendsen leg.; NHMO prep. 3899; NHMO . 1♀ VAY, Kristiansand : Adnevik , Unndalen ; 6.vii.2012 ; K. Berggren leg.; KBE prep. 13512; KBE . 1♀ VAY, Mandal : Hoven ; 24-27.vii.2017 ; K. Berggren & K. Hoven leg.; KBE prep. 13513; KBE . 1♂ VAY, Mandal : Hoven ; 22-24.vii.2017 ; K. Berggren & K. Hoven leg.; KBE prep. 13516; KBE . 1♂ VE, Horten : Knutsrod ; 20-21.vi.2008 ; L. Aarvik leg.; NHMO prep. 3898; NHMO . 1♀ VE, Notteroy : Ostre Bolaerne ; 27.vii.2006 ; R. Voith & K. Berggren leg.; KBE prep. 6104; KBE . 1♂ VE, Notteroy : Ostre Bolaerne ; 9.vii.2006 ; R. Voith & K. Berggren leg.; KBE prep. 10171; KBE . 1♀ VE, Tonsberg : Karlsvikodden ; vii.2005 ; R. Voith leg.; KBE prep. 10170; KBE . 1♂ O , Fredrikstad : Blasopp ; 4-19.vii.2013 ; O.J. Lonnve leg.; KBE prep. 13531; KBE . 1♂ O , Halden : Orod Grustak ; 5.vi-1.vii.2009 , F. Odegaard leg.; BOLD sample ID: NHMO-DAR-4107 (failed); KBE prep. 9438; KBE . 1♂ O , Hvaler : Asmaloy , Huser ; 3.vii.1994 ; L. Aarvik leg.; NHMO prep. 3965; NHMO . 1♂ O , Hvaler : Vesteroy , Slettevika ; 25.vi.2018 ; J.R. Gustad leg.; NHMO prep. 3980; NHMO . 1♂ O , Moss : Rygge , Sildebauen ; 13.vii.1980 ; L. Aarvik leg.; NHMO prep. 3890; NHMO . 1♂ O , Sarpsborg : Tune , Rakil ; 4.vii.2002 ; T.J. Olsen leg., NHMO prep. 3935; NHMO . Sweden 1♂ Oeland , Borgholm: Byrums Sandvik; 23.vii.1985 ; K. Berggren leg.; KBE prep. 13527; KBE . 1♂ Oeland , Borgholm: Byrums Sandvik; 23.vii.1985 ; K. Berggren leg.; KBE prep. 6142; KBE. Denmark 1♂ EJ, Anholt ; 1.viii.1975 ; E. S. Nielsen leg., Lundquist prep. 2050; ZMUC . 1♀ F, Aebelo ; 25.vi.-9.vii.1943 ; Worm-Hansen leg.; Rasmussen prep. 3936; ZMUC . 1♂ NWZ, Karuphoj ; 18.vii.2003 ; H. Hendriksen leg.; Hendriksen prep. 4194; ZMUC . 1♂ LFM, Boto ; 29.vi.1968 ; E. Traugott-Olsen leg.; Traugott-Olsen prep. 1242; ZMUC . 1♂ , 1♀ LFM, Valse Skov ; 10.vii.1982 ; J. Lundqvist leg.; Lundqvist prep. 1343, 1344; ZMUC . 1♀ LFM, Ulfshale ; 19.vii.1987 ; O. Karsholt leg.; Karsholt prep. 5381; ZMUC . 1♀ B, Slotslyngen ; 31.vii.1967 ; H.K. Jensen leg.; Jensen prep. 692; ZMUC . 1♀ B, Boderne ; 8.viii.1968 ; H.K. Jensen leg., Jensen prep. 810; ZMUC . Austria 1♀ , Oberoesterreich , NP Kalkalpen , Spering-Lackerbodenstrasse , ca 700 m ; 16.vii.2004 ; J. Wimmer leg ; genitalia in glycerine; TLMF. 1♀ , ditto, but 653 m , 14.vi.2009 ; J. Wimmer leg ; gen. slide GEL 1287 P. Huemer; TLMF. 1♂ Nordtirol , Umhausen N , unterh. Farst , 1370 m , 27.vi.2017 , P. Huemer leg ; GEL 1276 P. Huemer; TLMF. 1♂ Nordtirol , Umhausen N , unterh. Farst , 1370 m , 11.vi.2017 , P. Huemer leg ; DNA Barcode ID TLMF Lep 26819; TLMF. 1♂ Nordtirol , Zirl , Eigenhofen , linke Innau , 600 m , 7.vi.2012 , P. Huemer leg. ; DNA Barcode ID TLMF Lep 09322; TLMF. 1♂ Nordtirol , Brandenberg , Tiefenbachklamm , 645 m , 16.vi.2013 , P. Huemer leg. ; DNA Barcode ID TLMF Lep 10349; TLMF. 1♂ Osttirol , Lengberg , Drau-Auen , Schattseite , 630 m , 25.vi.2008 , H. Deutsch leg. ; DNA Barcode ID TLMF Lep 24185; TLMF. 1♂ Vorarlberg , Froedischtal , Schoenebuchweg , Klausen , 755 m , 14.vi.2017 , A. Mayr leg ; DNA Barcode ID TLMF Lep 25068; coll. A. Mayr. Germany 1♀ , Bavaria , Inning /A, 550 m , mid vii.1970 Zuernbauer F. leg.; genitalia in glycerin PH ; TLMF. Italy 1♂ Suedtirol , Oberrasen , Biotop Rasner Moeser S , 1100 m , 5.vii.2015 , P. Huemer leg.; DNA Barcode ID TLMF Lep 17982; TLMF . 1♂ Suedtirol , Laas , Tschenglser Au , Biotop Rasner Moeser S , 900 m , 20.vi.2015 , P. Huemer leg.; DNA Barcode ID TLMF Lep 18779; TLMF . Figures 5-8. Male genitalia of Batrachedra 5 B. confusella sp. nov., genitalia slide NHMO 3899 6 B. confusella sp. nov., distal end of phallus, genitalia slide NHMO 3899 7 B. pinicolella , genitalia slide NHMO 3891 8 B. pinicolella , distal end of phallus, genitalia slide NHMO 3892. Armenia 1♂ , Tavush Province , Diljan , 1395 m ; 13.vii.2011 ; O. Karsholt leg.; BOLD sample ID: ZMUC00029754; ZMUC . Morphological diagnosis. Batrachedra confusella sp. nov. (Fig. 3 ) and B. pinicolella (Fig. 4 ) cannot be separated externally with certainty though it is remarkable that none of the 56 specimens of B. confusella with images in BOLD has a spot at about the one-third the length of the forewing fold, a character commonly present in B. pinicolella . All 47 specimens of Norwegian dissected or barcoded specimens of B. confusella sp. nov. are without the spot on the fold. In eight of 20 dissected or barcoded Norwegian specimens of B. pinicolella the spot is present. Thus, the presence of the spot in the fold strongly indicates that the actual specimen belongs to B. pinicolella . With a wingspan of 11.0-12.0 mm, B. pinicolella is slightly larger than B. confusella sp. nov. on average. In the male genitalia we found a diagnostic character in the shape of the valva and the uncus. On average B. pinicolella (Fig. 7 ) has a longer and more slender valva than B. confusella sp. nov. (Fig. 5 ). The uncus is slightly different in the two species in shape and width. In B. confusella sp. nov. the lateral sides are smoother and with finer and longer setae than in B. pinicolella . In B. pinicolella the uncus is laterally distinctly rugose with coarse setae. The tip of the phallus in B. confusella sp. nov. (Fig. 6 ) is armed with a distinct cornutus. In B. pinicolella (Fig. 8 ) the cornutus is narrower and often will appear merely as a fold inside the phallus. The gnathos tends to be broader distally in B. pinicolella than in B. confusella sp. nov. In the female genitalia, the two species differ in the length of the signum. In B. pinicolella (Fig. 10 ) the signum is in the range 290-380 μm . In B. confusella sp. nov. (Fig. 9 ) the length of the signum ranges from 490 to 600 μm . In B. pinicolella the sclerotized portion of the ductus bursae is narrower and straighter than in B. confusella sp. nov. Fig. 22 of the female genitalia given by Koster and Sinev (2003) represents B. pinicolella in the present interpretation of the name. Figures 9-10. Female genitalia of Batrachedra 9 B. confusella sp. nov., genitalia slide NHMO 3937 10 B. pinicolella , genitalia slide NHMO 3961. Molecular diagnosis. Sequences of B. pinicolella and B. confusella sp. nov. form two well-defined clusters which received separate Barcode Index Numbers (BIN-codes): AAF0078 and AAF0077 respectively. The DNA barcode region of B. confusella sp. nov. shows 0.95% intraspecific divergence and a minimum divergence of 6.82% to its closest relative B. pinicolella (Fig. 11 ). The latter species shows no intraspecific variability in our material (the slight apparent variation in the neighbor-joining tree results only from variation in sequence lengths). Therefore, DNA barcodes allow safe identification of the two species. The 407-bp long fragment of the nuclear MDH gene differs by 5.8% from that of B. pinicolella . The 506-bp long fragment of EF-1a differs by 3.1% from that of B. pinicolella . Therefore, both examined nuclear genes support the status of the two species and permit their identification by these markers as well. All examined five specimens were infected by Wolbachia , while none of the specimens of B. pinicolella were infected, suggesting a difference between the two species also in this regard. Figure 11. Neighbor-joining visualization of genetic divergences of DNA barcode fragment of COI gene within and between European species of Batrachedra . The scale indicates 1% genetic divergence as calculated under Kimura 2 parameter model for nucleotide substitution. We generated a genomic dataset from nine individuals of B. pinicolella using ddRAD sequencing. We obtained 1.24 million reads per individual on average after quality filtering steps. After clustering 88% sequence similarity, we recovered 19,292 clusters per sample were retained with an average of 82.26 per sample for cluster depth (Table 1 ). A total length of ddRAD data is 926,224 bp, of which 9,242 are single nucleotide polymorphisms (SNP). Phylogenetic analysis using SNP data produced robust support for the relationship between the individuals (Fig. 12 ). The two revealed lineages corresponding to B. confusella sp. nov. and B. pinicolella , which have 100% bootstrap support. STRUCTURE also identified two genetic clusters (Fig. 11 ). Table 1. A summary of the ddRAD data.

Species	SampleID	Reads passed filter	Clusters at 88%	Coverage	Retained loci	Consensus loci
B. confusella	MM22066	1520904	29400	40.06	11608	4488
B. confusella	MM22068	1624893	31776	45.71	12030	4346
B. confusella	MM23464	2976135	62919	43.18	19568	4557
B. confusella	TLMF Lep 10349	1353160	16809	60.85	5292	2563
B. confusella	TLMF Lep 18779	624001	11396	47.99	3177	917
B. pinicolella	MM06705	808698	4251	163.44	572	149
B. pinicolella	MM21052	971904	6440	131.25	1305	200
B. pinicolella	MM21054	931540	7060	118.22	1344	370
B. pinicolella	TLMF Lep 17972	370227	3579	89.61	611	155
Total		1242385	19292	82.26	6167	1972

Figure 12. Maximum-likelihood tree inferred from the ddRAD SNP data. Bootstrap support values are indicated above the branches and only the values> 50% are shown. The barplot shows the assignments of individuals into two genetic clusters, the red clusters referring to Batrachedra confusella , the blue clusters to B. pinicolella . Each bar represents one individual and colours represent the proportion of the individuals that belong to each of the genetic cluster. Description. Male (Fig. 3 ). Wingspan 10-11 mm. Labial palp 2.3 times diameter of eye, cream, porrect, slightly curved, second segment longer than third, outer side with brownish suffusion, third segment on outer side with two bands. Head cream, with appressed scaling. Antenna glabrous, with appressed scales, pale ochreous brown, weakly ringed, rings becoming more distinct distally. Thorax ochreous yellow. Forewing ochreous yellow, with scattered dark brown scales, denser distally and on costa, discal spot small; cilia brownish grey; hindwing grey, cilia grey. Legs cream, inner sides with brownish suffusion, which on tarsi forms bands. Abdomen cream, with lateral grey suffusion. Female . Externally similar to male. Male genitalia (Figs 5 , 6 ) Uncus gradually narrowed towards tip, slightly concave at two-thirds length, medially with lateral setae, lateral margins smooth, with fine hairs; gnathos gradually narrowed, becoming parallel-sided before distal end, distal end spinose; valva nearly parallel-sided, dorsal margin curved distally, costa angled or slightly hooked at distal end; anellus lobes rounded; phallus long and slender, sub-distally with bulge, cornutus near distal end distinct, tapered proximally. Female genitalia (Fig. 9 ) Papillae anales broad with short setae; apophyses posteriores and anteriores of similar length, apophyses anteriores with basal fork; antrum funnel-shaped with medial ridge; ductus bursae with medial portion sclerotized and curved, anterior portion with two or three coils; corpus bursa with long (490-600 μm ) and oval signum; signum with numerous short, transverse ridges. Etymology. The species' name, Batrachedra confusella , indicates the confusion with its sister species, B. pinicolella . Biology. Due to the confusion of the two species, the biology is insufficiently known. Batrachedra confusella sp. nov. is the well-known species affecting Pinus , and it has been found in several localities of pure pine forests. However, a female beaten from an artificial afforestation of Larix at a lowland locality in Switzerland without Pinus in the nearby surroundings (Bryner in litt.) also belongs to B. confusella sp. nov., indicating that Larix may be an additional host-plant. Batrachedra pinicolella , in contrast, seems to be restricted to forests of Norway spruce ( Picea abies ), which is considered to be the host-plant. This hypothesis is proved by a dissected Finnish female specimen bred from Picea . Bolov and Sinev (1990) recorded outbreaks of B. pinicolella on Picea orientalis , P. abies , P. pungens , Abies sp. ( "European" ), A. nordmanniana , and less frequently on Pinus sp. ( "eastern" ), P. silvestris , and P. pityusa from the Kabardino-Balkaria republic in northern Caucasus. Possibly both species were involved. Verified specimens of Batrachedra pinicolella . Finland 1♂ PPe, Oulu; 7.vii.2011; M. Mutanen leg.; L. Aarvik prep. 2015.024; BOLD sample ID: MM21054; ZMUO. 1♀ PPe, Oulu; 7.vii.2011; M. Mutanen leg.; L. Aarvik prep. 2016.001; BOLD sample ID: MM21052; ZMUO. 1♀ A, Eckeroe ; 12.vii.2007; M. Mutanen leg.; L. Aarvik prep. 2016.002; BOLD sample ID: MM06705; ZMUO. 1♀ St, Rauma; la. Picea abies ; 5.iv.2019; J. Itaemies leg.; L. Aarvik prep. 2021.01; ZMUO. Norway 1♀ AAI, Bygland: Heddevika; 29.vii.2008; K. Berggren leg.; BOLD sample ID: NHMO-DAR-5194; KBE. 1♂ AAY, Birkenes: Nordasen ; vii-viii.2016; S. Svendsen leg.; KBE prep. 13526; KBE. 1♂ AAY, Birkenes: Bjorvand; 30.vii.2002; K. Berggren & S. Svendsen leg.; KBE prep. 13510; KBE. 1♀ AAY, Lillesand: Kjostvedt ; 9.vii.2007; K. Berggren leg.; KBE prep. 13514; KBE. 1♂ AK, Baerum : Isi; 11.vii.2003; P. Seglen leg; KBE prep. 13525; KBE. 1♂ AK, Baerum : Sandvika; 20.vii.1928; E. Barca leg.; NHMO prep. 2834; NHMO. 1♂ AK, As : As ; 1.viii.1982; L. Aarvik leg.; NHMO prep. 3900; NHMO. 1♂ AK, As : As ; 11.vii.1983; L. Aarvik leg.; NHMO prep. 3892; NHMO. 1♂ OS, Nordre Land: Tranligrenda; 5.vii.2010; K. Berggren leg.; KBE prep. 13519; KBE. 1♂ VAY, Flekkefjord: Helle; 18.vii.2013; K. Berggren leg.; BOLD sample ID: NHMO-DAR-4109; KBE prep. 13504; KBE. 1♂ VAY, Kristiansand: Gimle Gard ; 11.vii.2003; K. Berggren leg.; BOLD sample ID: NHMO-DAR-5193 (failed); KBE prep. 13523; KBE, 1♀ VAY, Kristiansand: Kuholmen; 17.vii.1970; K. Berggren leg.; KBE prep. 13511; KBE. 1♂ VAY, Kristiansand: Nedre Timenes; 31.vii.2011; K. Berggren leg.; BOLD sample ID: NHMO-DAR-4108; KBE prep. 13503; KBE. 1♂ VAY, Kristiansand: Stokken; 8.vii.1978; K. Berggren leg.; KBE prep. 13522; KBE. 1♂ VAY, Kristiansand: Sogne , Torvesanden; 27.vi.2014; K. Berggren leg.; KBE prep. 13517; KBE. 1♂ VE, Larvik: Frydenlund; 12.vii.2013; K. Berggren leg.; KBE prep. 13518; KBE. 1♀ O , Fredrikstad: Onsoy , Rauer; 21.vii.1920; E. Barca leg.; NHMO prep. 3961; NHMO. 1♂ O , Moss: Rygge, Sildebauen; 10.vii.1981; L. Aarvik leg.; NHMO prep. 3891; NHMO. Switzerland 1♀, BE, La Neuveville, Ligeresse, 810 m, 4.vii.2002, Larix decidua; leg. R. Bryner; coll. R. Bryner. Austria 1♂ Vorarlberg, Gaschurn-Partenen, Schuttfluren Lifinar, 1150 m, 31.vii.2018; P. Huemer leg.; BOLD sample ID: TLMF Lep 26889; TLMF. 1♂ Vorarlberg, Gaschurn-Partenen, Schuttfluren Lifinar, 1150 m, 2.vii.2018; P. Huemer leg.; GEL 1275 P. Huemer; TLMF. 1♀ Vorarlberg, Gaschurn-Partenen, u. Ganifer-Schrofen, 1460-1500 m, 19.vii.2019; P. Huemer leg.; GEL 1282 P. Huemer; TLMF. 1♀ Oberoesterreich , Nationalpark Kalkalpen, Lackerbodenstrasse , 653 m, 14.vi.2009; J. Wimmer leg.; GEL 1287 P. Huemer; TLMF. Germany 1♀ Baden-Wuerttemberg , Schwarzwald, Buchenberg, 21.vii.1954; H.G. Amsel leg; genitalia in gylcerine; TLMF. Italy 1♂ Suedtirol , Oberrasen, Biotop Rasner Moeser S, 1100 m, 5.vii.2015, P. Huemer leg.; DNA Barcode ID TLMF Lep 17972; TLMF. Distribution. The species pair B. confusella sp. nov. and B. pinicolella is widely distributed in Europe but seems to be absent from the Mediterranean (http://www.faunaeur.org; accessed on 25.iv.2021). However, as former records have been summarized among the latter taxon, a detailed study of distribution of both species is required for future studies. From our sequenced and/or genitalized material, B. confusella sp. nov. together with its major hostplant, Pinus sylvestris , is distributed in the temperate zones between the Alps in the South and Fennoscandia in the north. Batrachedra pinicolella in concordance with its major host-plant, Picea abies , shows a boreo-montane distribution pattern with isolated records from the Alps and mountainous areas of Central Europe as well as northern and north-western Europe. A barcoded specimen of B. confusella sp. nov. from Armenia (coll. ZMUC) confirms the presence of this species in Caucasus. According to images of genitalia on the mothdissection.co.uk website, both species are present in the United Kingdom (https://mothdissection.co.uk/species.php?Tx=Batrachedra_pinicolella; accessed on 30.iv.2021)