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Research Article
Alpha taxonomy of the genus Kessleria Nowicki, 1864, revisited in light of DNA-barcoding (Lepidoptera, Yponomeutidae)
expand article infoPeter Huemer, Marko Mutanen§
‡ Tiroler Landesmuseen-Betriebsgeselllschaft m.b.H., Innsbruck, Austria
§ University of Oulu, Oulu, Finland
Open Access

Abstract

The taxonomy of Kessleria, a highly specialized montane genus of Yponomeutidae with larval host restriction to Saxifragaceae and Celastraceae (Saxifraga spp. – subgenus Kessleria; Saxifraga spp. and Parnassia spp. – subgenus Hofmannia), is revised based on external morphology, genitalia and DNA barcodes. An integrative taxonomic approach supports the existence of 29 species in Europe (the two known species from Asia and North America are not treated herein). A full 658 bp fragment of COI was obtained from 135 specimens representing 24 species, a further seven sequences are >560 bp. Five new species are described: Kessleria cottiensis sp. n. (Prov. Torino, Italy; Dep. Hautes Alpes, France), Kessleria dimorpha sp. n. (Dep. Alpes-de-Haute-Provence, France), Kessleria alpmaritimae sp. n. (Dep. Alpes-Maritimes, France), Kessleria apenninica sp. n. (Prov. Rieti, Prov. L´Aquila, Italy), and Kessleria orobiae sp. n. (Prov. Bergamo, Italy).

Keywords

Lepidoptera, Yponomeutidae, Kessleria, new species, integrative taxonomy, DNA barcode, morphology, cryptic diversity, European mountains

Introduction

The genus Kessleria Nowicki, 1864 is one of the striking examples of long underestimated or neglected diversity in the generally well known fauna of European Lepidoptera. This deficiency of knowledge is reflected in the fact that only 9 out of the 29 European species were described before 1960, and 18 species, or two-thirds of the fauna, after 1990 (Friese 1960, Huemer and Tarmann 1992, 1993). The genus is exceptional in Lepidoptera due to its highly specialized host-plant relationship with the Saxifragaceae, a habit shared e.g. by a section of the Curculionidae genus Dichotrachelus (Merigalli et al. 2013). Whereby the large majority of species is restricted to Saxifraga, two species feed on the herbaceous Celastraceae genus Parnassia, long considered as Saxifragaceae (The Angiosperm Phylogeny Group 2009). Through these host-plant relationships, Kessleria is a genus characteristic of mountain regions in the northern hemisphere, reaching high altitudes of about 3000 m in the European Alps and only exceptionally occurring below 600 m. In such alpine environments adult morphology may be adapted to rough climatic conditions with female brachyptery observed in several families of Lepidoptera (Sattler 1991), including minimum five species of Kessleria. Two major revisions (Friese 1960, Huemer and Tarmann 1992) are primarily based on external and internal morphology of adults, supplemented by ecological data. These authors established a stable and undisputed alpha taxonomy of Kessleria, which has been in use for the last two decades. Recently discovered suspected morphospecies led to the implementation of molecular methods as an additional tool in species delimitation. Sequences of the COI barcode region (Hebert et al. 2003, 2009) confirmed the species status of the vast majority of previously described taxa, and helped in resolving suspected cryptic species-complexes and in delimiting five species new to science.

Material and methods

Extensive descriptions and diagnoses of previously described European species of Kessleria including keys to males and females, colour figures of adults, black-and-white figures of male and female genitalia, last abdominal segments, illustrations of wing venation and figures of larval habits and habitats have been published by Huemer and Tarmann (1992, 1993) and are not repeated here.

Our study was initially based on morphology of the extensive material published in detail by Huemer and Tarmann (1992), and about 100 additional specimens, with DNA barcode sequences as an additional tool for delimitation of cryptic species. Most of the material was set and dried according to standard practice, some were spread, and a few only pinned. Genitalia preparations followed standard techniques for microlepidoptera (Robinson 1976), adapted for Kessleria e.g. by the manual eversion of cornuti (Huemer and Tarmann 1992). Wing venation was not considered for new species descriptions as it proved irrelevant for alpha taxonomy in the genus in the earlier revision by Huemer and Tarmann (1992).

We tried to obtain DNA barcode sequences, a 658 base-pair long segment of the 5’ terminus of the mitochondrial COI gene (cytochrome c oxidase 1), from 150 specimens, three from LMK and ZMUO respectively, and 144 from TLMF. DNA samples (from a single dried leg) were prepared according to the accepted standards. Legs from 150 specimens of Kessleria were processed at the Canadian Centre for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph) using their standard high-throughput protocol described in deWaard et al. (2008). Successfully sequenced voucher specimens are listed in Suppl. material 1 together with species names, sample-IDs, process-IDs, BINs, COI-5P sequence length, and trace counts. Sequences were submitted to GenBank during printing stage; GenBank accession numbers further details including complete voucher data and images can be accessed in the public dataset “Lepidoptera of Europe - Kessleriahttps://doi.org/10.5883/DS-LEAKE in the Barcode of Life Data Systems (BOLD; Ratnasingham and Hebert 2007). Degrees of intra- and interspecific variation in the DNA barcode fragment were calculated under the Kimura 2-parameter (K2P) model of nucleotide substitution using analytical tools in BOLD systems v3.0. (http://www.boldsystems.org). A neighbor-joining tree of DNA barcode data of European taxa was constructed using Mega 5 (Tamura et al. 2011) under the K2P model for nucleotide substitutions. In taxonomic delimitation, we applied principles of integrative taxonomy (Padial et al. 2010) and considered a barcode divergence of roughly 2% supported by at least one morphological character indicating species distinctiveness. We acknowledge that any threshold value of genetic distinctiveness is artificial and should not alone be used as indicating species status (cf. Collins and Cruickshank 2013), for which reason we considered 2% genetic difference associated with at least one morphological character indicating species integrity in the sense of e.g. General Lineage Species Concept (deQueiroz 1998) and Phylogenetic (diagnostic) Species Concept (Cracraft 1989), which both are applicable in delimiting also allopatric populations.

Photographs of the adults were taken with an Olympus SZX 10 binocular microscope and an Olympus E 3 digital camera, and processed using the software Helicon Focus 4.3 and Adobe Photoshop CS4 and Lightroom 2.3. Genitalia photographs were taken with an Olympus E1 Digital Camera from Olympus BH2 microscope.

Measurements were taken with a micrometer eyepiece.

Abbreviations of institutional collections

BMNH Natural History Museum (British Museum, Natural History) London, United Kingdom

LMK Landesmuseum Kärnten, Klagenfurt, Austria

MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain

MNHU Museum für Naturkunde der Humboldt Universität, Berlin, Germany

NHMV Naturhistorisches Museum, Vienna, Austria

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany

SMNK Staatliches Museum für Naturkunde, Karlsruhe, Germany

TLMF Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria

ZMUO Zoological Museum, University of Oulu, Finland

ZMUC Zoological Museum, Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark

ZSM Zoologische Staatssammlung, München, Germany

Results

The checklist of European Kessleria largely follows Huemer and Tarmann (1992). The proposed tentative structure into species groups is based on morphology and as far as available DNA barcode data, but a well-grounded phylogenetic analysis will require further data, particularly from nuclear markers.

Checklist of European Kessleria

Kessleria Nowicki, 1864

Subgenus Kessleria Nowicki, 1864

Kessleria alpicella-group

K. alpicella (Stainton, 1851)

= K. alpicella (Herrich-Schäffer, 1855), Homonym

K. mixta Huemer & Tarmann, 1992

Kessleria alternans-group

K. alternans (Staudinger, 1871)

K. cottiensis sp. n.

K. dimorpha sp. n.

K. wehrlii Huemer & Tarmann, 1992

K. alpmaritimae sp. n.

Kessleria petrobiella-group

K. nivescens Burmann, 1980

K. petrobiella (Zeller, 1868)

Kessleria albanica-group

K. macedonica Huemer & Tarmann, 1992

K. albanica Friese, 1960

K. burmanni Huemer & Tarmann, 1992

K. insubrica Huemer & Tarmann, 1993

K. hauderi Huemer & Tarmann, 1992

Kessleria apenninica-group

K. apenninica sp. n.

K. diabolica Huemer & Tarmann, 1992

K. brevicornuta Huemer & Tarmann, 1992

K. pyrenaea Friese, 1960

K. brachypterella Huemer & Tarmann, 1992

Kessleria zimmermanni-group

K. zimmermanni Nowicki, 1864

= K. tatrica Friese, 1960

K. albomaculata Huemer & Tarmann, 1992

K. caflischiella (Frey, 1880)

Kessleria albescens-group

K. klimeschi Huemer & Tarmann, 1992

K. helvetica Huemer & Tarmann, 1992

K. inexpectata Huemer & Tarmann, 1992

K. orobiae sp. n.

K. albescens (Rebel, 1899)

Subgenus Hofmannia Heinemann & Wocke, 1877

K. saxifragae (Stainton, 1868)

K. fasciapennella (Stainton, 1849)

= K. longipenella Friese, 1960

Molecular analysis

Sequencing resulted in a full barcode fragment of 658 bp for 135 specimens, covering 24 species. A further seven sequences that were longer than 560 bp were included in the analysis. A single short sequence of 307 bp was not considered, and sequencing failed for seven voucher specimens. Mean intraspecific divergence is 0.61%. It ranges from 0–4.27%, exceeding 2% only in three species, which, however, may include further cryptic diversity (e.g. K. alpicella, K. albanica and K. inexpectata) and should be tested accordingly with more material (Table 1, Fig. 1). On the contrary, interspecific divergence in the genus is much higher with a mean divergence of 10.38% and maximum of 16.22%. Interspecific divergence to the nearest neighbour ranges from 1.86–9.29%, with the only exception being K. inexpectata and K. helvetica, which overlap in DNA barcode (Table 1, Fig. 1).

Figure 1.

Neighbour-joining tree (Kimura 2 parameter, built with MEGA 5; cf. Tamura et al. 2011). Note: the scale bar only applies to internal branches between species. The width of the triangles represents the sample size, and the depth the relative genetic variation within the cluster (2× scale bar). Source: DNA Barcode data from BOLD (Barcode of Life Database, cf. Ratnasingham and Hebert 2007).

From sequence analysis of 20 Kessleria species based on at least three sequences, 17 species are delimited by a minimum of one to a maximum of 10 diagnostic characters whereas K. inexpectata, K. cottiensis and K. alpmaritimae have no diagnostic character (Table 1).

Intraspecific distance and interspecific divergence to the nearest neighbour in the genus Kessleria. Source: DNA Barcode data from BOLD (Barcode of Life Database, cf. Ratnasingham and Hebert 2007).

Species # sequ Mean intra Max intra Nearest neighbour Nearest species Nearest species Diagnostic characters
Kessleria albanica 5 2.05 2.98 PHLAB1059-10 Kessleria burmanni 9.29 8
Kessleria albescens 3 0 0 PHLAD145-11 Kessleria orobiae 2.66 3
Kessleria albomaculata 1 N/A N/A PHLAD138-11 Kessleria petrobiella 6.76 -
Kessleria alpicella 12 1.52 4.27 PHLAD119-11 Kessleria wehrlii 6.9 6
Kessleria alpmaritimae 6 0 0 PHLAD119-11 Kessleria wehrlii 1.87 0
Kessleria alternans 10 0.12 0.31 PHLAD122-11 Kessleria cottiensis 2.65 3
Kessleria apenninica 4 1.06 1.71 PHLAI438-13 Kessleria pyrenaea 5.47 3
Kessleria burmanni 6 0 0 PHLAD140-11 Kessleria hauderi 7.61 6
Kessleria caflischiella 8 0.04 0.15 PHLAD118-11 Kessleria alpmaritimae 6.39 6
Kessleria cottiensis 5 0 0 PHLAB957-10 Kessleria dimorpha 1.86 0
Kessleria dimorpha 4 0.08 0.15 PHLAD122-11 Kessleria cottiensis 1.86 1
Kessleria fasciapennella 8 0.04 0.15 PHLAI063-12 Kessleria saxifragae 7.21 8
Kessleria hauderi 2 0 0 PHLAB1059-10 Kessleria burmanni 7.61 -
Kessleria helvetica 1 N/A N/A PHLAB1065-10 Kessleria inexpectata 0.31 -
Kessleria inexpectata 7 1.4 2.18 LASTS544-14 Kessleria helvetica 0.31 0
Kessleria insubrica 4 0.08 0.15 PHLAB1059-10 Kessleria burmanni 8.95 9
Kessleria klimeschi 5 0.06 0.15 PHLAB1065-10 Kessleria inexpectata 8.83 10
Kessleria nivescens 14 1.09 2.5 PHLAD138-11 Kessleria petrobiella 3.29 4
Kessleria orobiae 5 0.31 0.46 PHLAB1067-10 Kessleria albescens 2.66 1
Kessleria petrobiella 4 0 0 PHLAD132-11 Kessleria nivescens 3.29 1
Kessleria pyrenaea 1 N/A N/A PHLAB861-10 Kessleria apenninica 5.47 -
Kessleria saxifragae 20 0.44 1.29 LEFIB126-10 Kessleria fasciapennella 7.21 9
Kessleria wehrlii 4 0 0 PHLAD118-11 Kessleria alpmaritimae 1.87 1
Kessleria zimmermanni 5 0 0 PHLAD138-11 Kessleria petrobiella 5.73 6

Taxonomy

New species of Kessleria

Kessleria alternans-group

The K. alternans-group is characterized by strong sexual dichroism and to a lesser extent dimorphism, with females being smaller and lighter, but not strongly brachypterous (Figs 211). The genitalia are characterized by the strong reticulate sculpture of the apical part of the phallus (Figs 1221) and the ductus bursae, which is extended into the corpus bursae (Figs 2226). Larval host-plants, as far as known, belong to the small-leaved Saxifraga spp., particularly the S. oppositifolia-complex, and to broad-leaved congeners such as S. paniculata. Five species belong to this group: K. alternans, K. wehrlii and the new taxa K. cottiensis, K. dimorpha and K. alpmaritimae.

Figures 2–7.

Kessleria adults in dorsal view. 2 K. alternans, ♂, Switzerland, Graubünden, SE Sils-Maria, 1820–1870 m, 13.7.1989, leg. Huemer, Karsholt & Tarmann (TLMF) 3 K. alternans, ♀, same data (TLMF) 4 K. cottiensis sp. n., ♂, paratype, Italy, Prov. Torino, Alpi Cozie, V. delle Finestre, 1700 m, 27.7.1990, leg. Huemer & Tarmann (TLMF) 5 K. cottiensis sp. n., ♀, paratype, same data (TLMF) 6 K. dimorpha sp. n., ♂, paratype, France, Dep. Hautes-Alpes, Col Agnel, 2770 m, 4.8.2010, leg. Huemer (TLMF) 7 K. dimorpha sp. n., ♀, paratype, same data (TLMF).

Figures 8–11.

Kessleria adults in dorsal view. 8 K. wehrlii, ♂, paratype, France, Dep. Alpes Maritimes, Mont Gelas Massiv, Mont Colomb W, 2450 m, 24.7.1990, leg. Huemer & Tarmann (DNA barcode ID TLMF Lep 01857) (TLMF) 9 K. wehrlii, ♀, paratype, same data (TLMF) 10 K. alpmaritimae sp. n., ♂, paratype, France, Dep. Alpes Maritimes, Marguareis W-Hang, Navela, 2100–2200 m, 18.–19.7.1991, leg. Huemer & Tarmann (TLMF) 11 K. alpmaritimae sp. n., ♀, paratype, same data (DNA barcode ID TLMF Lep 01851) (TLMF).

Figures 12–17.

Kessleria male genitalia. 12 K. alternans, Switzerland, Graubünden, SE Sils-Maria, 1820–1870 m, 13.7.1989, leg. Huemer, Karsholt & Tarmann, gen. slide YPO 22 (TLMF) 13 idem, distal part of phallus enlarged 14 K. cottiensis sp. n., paratype, Italy, Prov. Torino, Alpi Cozie, V. delle Finestre, 1700 m, 27.7.1990, leg. Huemer & Tarmann, gen. slide YPO 66 (TLMF) 15 idem, distal part of phallus enlarged 16 K. dimorpha sp. n., paratype, France, Dep. Hautes-Alpes, Col Agnel, 2770 m, 4.8.2010, leg. Huemer gen. slide YPO 149 (TLMF) 17 idem, holotype, gen. slide YPO 158, distal part of phallus enlarged.

Figures 18–21.

Kessleria male genitalia. 18 K. wehrlii, paratype, France, Dep. Alpes Maritimes, Mont Gelas Massiv, Mont Colomb W, 2450 m, 24.7.1990, leg. Huemer & Tarmann (DNA barcode ID TLMF Lep 01857), gen. slide YPO 64 (TLMF) 19 idem, distal part of phallus enlarged 20 K. alpmaritimae sp. n., paratype, France, Dep. Alpes Maritimes, Marguareis W-Hang, Navela, 2100–2200 m, 18.-19.7.1991, leg. Huemer & Tarmann, gen. slide YPO 55 (TLMF) 21 idem, distal part of phallus enlarged.

Figures 22–23.

Kessleria female genitalia. 22 K. alternans, Switzerland, Graubünden, SE Sils-Maria, 1820–1870 m, 13.7.1989, leg. Huemer, Karsholt & Tarmann, gen. slide YPO 6 (TLMF) 23 K. cottiensis sp. n., paratype, Italy, Prov. Torino, Alpi Cozie, V. delle Finestre, 1700 m, 27.7.1990, leg. Huemer & Tarmann, gen. slide YPO 67 (TLMF).

Figures 24–26.

Kessleria female genitalia. 24 K. dimorpha sp. n., paratype, France, Dep. Hautes-Alpes, Col Agnel, 2770 m, 4.8.2010, leg. Huemer, gen. slide YPO 159 (TLMF) 25 K. wehrlii, paratype, France, Dep. Alpes Maritimes, Mont Gelas Massiv, Mont Colomb W, 2450 m, 24.7.1990, leg. Huemer & Tarmann, gen. slide YPO 69 (TLMF) 26 K. alpmaritimae sp. n., paratype, France, Dep. Alpes Maritimes, Marguareis W-Hang, Navela, 2100–2200 m, 18.-19.7.1991, leg. Huemer & Tarmann (TLMF).

Figures 27–31.

Kessleria female genitalia, details of VIII abdominal segment enlarged. 27 K. alternans, Switzerland, Graubünden, SE Sils-Maria, 1820–1870 m, 13.7.1989, leg. Huemer, Karsholt & Tarmann, gen. slide YPO 6 (TLMF) 28 K. cottiensis sp. n., paratype, Italy, Prov. Torino, Alpi Cozie, V. delle Finestre, 1700 m, 27.7.1990, leg. Huemer & Tarmann, gen. slide YPO 67 (TLMF) 29 K. dimorpha sp. n., paratype, France, Dep. Hautes-Alpes, Col Agnel, 2770 m, 4.8.2010, leg. Huemer, gen. slide YPO 159 (TLMF) 30 K. wehrlii, paratype, France, Dep. Alpes Maritimes, Mont Gelas Massiv, Mont Colomb W, 2450 m, 24.7.1990, leg. Huemer & Tarmann, gen. slide YPO 69 (TLMF) 31 K. alpmaritimae sp. n., paratype, France, Dep. Alpes Maritimes, Marguareis W-Hang, Navela, 2100–2200 m, 18.–19.7.1991, leg. Huemer & Tarmann (TLMF).

Figure 32.

Distribution pattern of the Kessleria alternans-group from examined material.

Figures 33–34.

Kessleria adults in dorsal view. 33 K. apenninica sp. n., ♂, holotype, Italy, L´Aquila, NP Gran Sasso, ex Miniera di Lignite, 1750 m, 14.-15.7.2010, leg. Huemer (DNA barcode ID TLMF Lep 01663) (TLMF) 34 K. pyrenaea, ♂, Spain, Aragon, Parzan (Bielsa) env. Pico de la Rubinera, 2700–3005 m, 10.-11.7.2010, leg. Cesanek (DNA barcode ID TLMF Lep 08933) (TLMF) (coll. Tokár).

Figures 35–38.

Kessleria male genitalia. 35 K. apenninica sp. n., ♂, holotype, Italy, L´Aquila, NP Gran Sasso, ex Miniera di Lignite, 1750 m, 14.-15.7.2010, leg. Huemer gen. slide YPO 147 (DNA barcode ID TLMF Lep 01663) (TLMF) 36 idem, distal part of phallus enlarged 37 K. pyrenaea, ♂, Spain, Aragon, Parzan (Bielsa) env. Pico de la Rubinera, 2700–3005 m, 10.–11.7.2010, leg. Cesanek, gen. slide 15/1391 P.Huemer (DNA barcode ID TLMF Lep 08933) (coll. Tokár) 38 idem, distal part of phallus enlarged.

Kessleria cottiensis sp. n.

Type material

Holotype. ♂, „ITALIA, Prov. Torino Alpi Cozie, 2150 m Colle delle Finestre 27.7.1990 leg. Huemer & Tarmann“ „YPO 58 ♂ P. Huemer“ (TLMF).

Paratypes. Italy: 13 ♂, 7 ♀, same data, genitalia slides YPO 59 ♂ P. Huemer, YPO 77 ♂ P. Huemer, DNA barcode IDs TLMF Lep 03106, TLMF Lep 03107, TLMF Lep 03108 (TLMF); 3 ♂, 6 ♀, same data, but V. delle Finestre, 1700 m, genitalia slides YPO 66 ♂ P. Huemer, YPO 67 ♀ P. Huemer (TLMF). France: 1 ♂, Dep. Hautes-Alpes, Nevache, 1950m, 31.7.2001, leg. Nel, genitalia slide 12937 J. Nel, DNA barcode ID TLMF Lep 03144 (TLMF); 1 ♂, Dep. Hautes-Alpes, Vallee de la Claree, 2000 m, 3.7.2002, leg. Nel, genitalia slide 14644 J. Nel, DNA barcode ID TLMF Lep 03142 (TLMF).

Diagnosis

K. cottiensis resembles other taxa of the K. alternans-group in wing markings and colour (Figs 211), but the male differs in the on average smallest forewing length of 7.59 mm (n=14) vs. 8.75 mm (n=13) in K. wehrlii, 8.55 mm (n=26) in K. alternans, 8.05 mm (n=6) in K. alpmaritimae and 8.25 mm (n=6) in K. dimorpha. Compared to K. wehrlii and K. alpmaritimae, the whitish suffusion of the forewing is largely reduced. The ochre-brown markings, which are well present in K. alternans, are almost completely absent. The female of K. cottiensis is also distinctly smaller than K. alternans, with a forewing length of 6.13 mm (n=8) on average vs. 7.58 mm (n=11), whereas females of K. cottiensis and K. alpmaritimae are indistinguishable. K. wehrlii is insufficiently known from only a single worn female specimen. Compared to the genetically nearest neighbour K. dimorpha, which is similar in forewing length (6.0 mm, n=4), the hindwings are slightly less reduced and the ground colour of the forewing is much lighter. Diagnostic characters in genitalia are generally weak. The male genitalia differ from K. alpmaritimae by the medially strongly widened valva, from K. alternans by the more slender cornuti, from K. wehrlii by three instead of four cornuti, and from the nearest neighbour K. dimorpha by the distinctly longer phallus (1.65–1.70 mm vs. 1.32–1.36 mm) (Figs 1221). The female genitalia show no diagnostic characters to related species of the K. alternans-group (Figs 2231).

Description

Male (Fig. 4). Head covered with erected whitish hair-like scales; antennae dark grey-brown, indistinctly lighter ringed; thorax and tegulae dark grey-brown. Forewing length 7.0–8.8 mm (Ø 7.59 mm; n=14); ground colour dark grey-brown, mottled with whitish scales, particularly in distal half, forming indistinct patches in fold and on costa at about 4/5; irregular black dots on veins and few brown scales in medial part of wing; oblique blackish fascia at about 1/3 to 1/2 indistinct; termen mixed whitish-grey, dark grey-brown in apical part; fringes light grey, darker in apical area. Hindwing dark grey, fringes with dark grey base, distal part light grey.

Female (Fig. 5). Head covered with erected whitish hair-like scales; antennae grey-brown, distinctly ringed whitish; thorax and tegulae whitish. Forewing length 5.8–6.3 mm (Ø 6.13 mm; n=8); ground colour whitish, mottled with black scales, particularly along veins and in tornal part, patches of brown scales in medial part of wing from base to end of cell; oblique blackish fascia at about 1/3 to 1/2 indistinct, separated into larger dash-like patch and reduced dot; termen whitish with some dark grey-brown mottling in apical part; fringes greyish-white, darker in apical area.

Male genitalia (Figs 14–15). Socii long and slender, with apical spine; anterior margin of tegumen with medial process; gnathos broadly tongue-shaped, smooth; valva moderately slender, length 0.66–0.70 mm, medially distinctly widened, maximum width of about 0.24–0.26 mm, densely covered with long hairs in medial part and short setae on ventromedial margin, ventromedial part weakly convex, distal part slender with ventrally convex and dorsally projected apex, costa strongly sclerotized with some distal dentation; sacculus oval, weakly confined, densely covered with strong setae; saccus sizeable in length, about 0.38–0.42 mm, stout, about same width throughout, apex rounded; phallus ca. 1.65–1.70 mm long and slender, straight, apically with distinct reticulate sculpture, uneverted vesica with ca. 0.76–0.80 mm long sclerotized part, three prominent needle-shaped, one single, the other basally connected, cornuti of about 0.46–0.51 mm in length.

Female genitalia (Figs 23, 28). Genitalia ca. 4.9 mm in length; papilla analis large, densely covered with long setae; apophysis posterior rod like, ca. 0.70 mm, about length of apophysis anterior; apophysis anterior rod like; posterior part bifurcated with straight dorsal and inwardly curved ventral branch; lamella postvaginalis with large sclerotized mediolateral patches, covered with microtrichia, medial area less sclerotized, posteriolateral part with hump, covered with some long setae; ostium bursae membranous; antrum weakly sclerotized, funnel-shaped; ductus bursae very long, about 2.8 mm, from entrance of ductus seminalis to transition into corpus bursae covered with finely granulous sculpture, particularly in posterior and anterior part, ductus bursae extended into posterior part of corpus bursae, entrance to corpus bursae weakly widened; corpus bursae well delimited, about 1.2 mm in length, ovoid, with small plate-like signum.

Molecular data

The average intraspecific divergence of the barcode region is 0.0% (n=5). The minimum distance to the nearest neighbour K. dimorpha is 1.86%, whereas the minimum divergence to K. alternans, K. alpmaritimae and K. wehrlii ranges from 2.65% and 2.98% to 3.63%, respectively.

Etymology

The species name refers to the type locality in the Cottian Alps (Alpi Cozie, Alpes cottiennes).

Distribution

(Fig. 32). Only known from a small area in the southwestern Alps (Cottian Alps) of Italy and France. An alleged K. alternans from the Graian Alps (Huemer and Tarmann 1992) likely refers to K. cottiensis, but the specimen in question could not be re-examined.

Ecology

Host-plant and early stages unknown. The adults were collected in late July. The flight period can most likely be further prolonged, depending on snow coverage and elevation. A specimen collected earlier during the summer, on June 9th, by Jäckh in Valle delle Finestre (Huemer and Tarmann 1992) probably belongs to K. cottiensis. The adults were collected during the day, flying freely in the morning hours and flushed out from their resting places with a bee-smoker. The species occurs in alpine grassland interspersed with calcareous rocks. Vertical distribution: from about 1700 m to 2150 m.

Remarks

K. cottiensis described here was already suspected to be distinctive from K. alternans by Huemer and Tarmann (1992), who illustrated adults (Figs 6–7) and cornuti of male genitalia (Fig. 105).

Kessleria dimorpha sp. n.

Type material

Holotype. ♂, „Frankreich Dep. Hautes-Alpes Col Agnel, 2770 m 6°59'02"E, 44°41'10"N 4.8.2010, leg. Huemer TLMF 2011-010“ „BC TLMF Lep 01756“ „YPO 158 ♂ P. Huemer” (TLMF).

Paratypes. France: 7 ♂, 5 ♀, same data, genitalia slides YPO 149 ♂ P. Huemer, YPO 159 ♀ P. Huemer, DNA barcode IDs TLMF Lep 01757, TLMF Lep 01758, TLMF Lep 01759 (TLMF); 4 ♂, 1 ♀, same data, leg. Wieser (LMK).

Diagnosis

K. dimorpha resembles other taxa of the K. alternans-group in wing markings and colour (Figs 211), but the male differs from the genetically nearest neighbour K. cottiensis by the on average distinctly larger forewing length of 8.25 mm (n=6) vs. 7.59 mm (n=14). Larger species are K. wehrlii with forewing length 8.75 mm (n=13) and K. alternans with 8.55 mm (n=26), whereas K. alpmaritimae with 8.05 mm (n=6) is of similar size. Furthermore, K. wehrlii and K. alpmaritimae have a much more prominent whitish suffusion on the forewing, whereas the ochre-brown markings of K. dimorpha rather resemble K. alternans. The female of K. dimorpha reflects a tendency to reinforced brachyptery and is distinctly smaller than K. alternans with a forewing length of only 6.0 mm (n=4) on average vs. 7.58 mm (n=11), whereas females of K. cottiensis and K. alpmaritimae are strongly suffused with whitish scales. K. wehrlii is insufficiently known from only a single worn female specimen. The male genitalia differ from all other taxa of the K. alternans-group by the distinctly shorter phallus with <1.40 mm vs. a minimum of 1.50 mm in other species (Figs 1221). The female genitalia show no diagnostic characters to related species of the K. alternans-group (Figs 2231).

Description

Male (Fig. 6). Head covered with ochre-brown hair-like scales; antennae almost unicolorous dark grey-brown; thorax and tegulae mixed dark grey-brown and ochre-brown. Forewing length 8.0–8.4 mm (Ø 8.25 mm; n=6); ground colour dark grey, intensively mottled with light grey, ochre-brown and whitish scales, white medial patch in fold; black dots particularly on costal and subcostal veins; black patch near base and at end of cell, oblique blackish fascia at about 1/3 to 1/2 reduced to large patch in fold; termen mixed dark and light grey; fringes basally dark grey, distal part whitish-grey, darker in apical area. Hindwing dark grey, fringes with dark grey base, distal part whitish-grey.

Female (Fig. 7). Head covered with erected whitish hair-like scales; antennae grey-brown, indistinctly lighter ringed; thorax and tegulae whitish. Forewing length 6.0 mm (Ø 6.0 mm; n=4); ground colour whitish, mottled with dark grey and black, particularly along fold and in tornal part, few black dots along costal and subcostal vein, small patches of ochre-brown scales in medial part of wing particularly in fold and at end of cell; oblique blackish fascia at about 1/3 to 1/2 indistinct, separated into larger dash-like patch and reduced dot; termen mixed whitish and dark grey; fringes whitish-grey, with dark grey basal part near apex. Hindwing grey, fringes whitish-grey with darker basal part.

Male genitalia (Figs 16–17). Socii long and slender, with apical spine; anterior margin of tegumen with medial process; gnathos broadly tongue-shaped, smooth; valva moderately slender, length 0.71–0.72 mm, medially weakly widened, maximum width of about 0.26–0.28 mm, densely covered with long hairs in medial part and short setae on ventromedial margin, ventromedial part weakly convex, distal part moderately slender with ventrally convex and dorsally projected apex, costa strongly sclerotized with weak distal dentation; sacculus oval, weakly confined, densely covered with strong setae; saccus short, about 0.32 mm, stout, about same width throughout, apex rounded; phallus ca. 1.32–1.36 mm long and slender, straight, apically with distinct reticulate sculpture, uneverted vesica with ca. 0.58–0.60 mm long sclerotized part, three to four prominent needle-shaped cornuti of about 0.38–0.40 mm in length.

Female genitalia (Figs 24, 29). Genitalia ca. 4.9 mm in length; papilla analis large, densely covered with long setae; apophysis posterior rod like, ca. 0.72 mm, about length of apophysis anterior; apophysis anterior rod like; posterior part bifurcated with straight dorsal and inwardly curved ventral branch; lamella postvaginalis with large sclerotized mediolateral patches, covered with microtrichia, medial area less sclerotize, posterolateral part with hump, covered with some long setae; ostium bursae membranous; antrum weakly sclerotized, funnel-shaped; ductus bursae long, ca. 2.3 mm, posterior part from entrance of ductus seminalis anterior and anterior part covered with finely granulous sculpture, medial part with weak and hardly discernible sculpture, ductus bursae extended into posterior part of corpus bursae, entrance to corpus bursae weakly widened; corpus bursae well delimited, about 1.4 mm in length, ovoid, with small plate-like signum.

Molecular data

The average intraspecific divergence of the barcode region is low with 0.08%, ranging from a minimum of 0% to a maximum of 0.15% (n=4). The minimum distance to the nearest neighbour K. cottiensis is 1.86%, whereas the minimum divergence to K. alternans, K. alpmaritimae and K. wehrlii ranges from 3.15% and 3.64% to 4.3%, respectively.

Etymology

The species name refers to the remarkable sexual dimorphism.

Distribution

(Fig. 32). Only known from the type locality, the French side of Col Agnel (Cottian Alps), close to the Italian border.

Bionomics

Host-plant and early stages unknown. Based on the type locality, the host-plant is most likely Saxifraga cf. oppositifolia. The adults have been collected in early August during the early morning hours from about 7–10a.m. at low temperatures between 2–5 °C. Males were flying actively during this period in search for females. Both sexes were later found in copula, often sitting on cushions of their suspected host-plant. A single female was found at light, attracted from its nearby habitat and crawling upwards to the light tower, but unable to fly actively. From personal observations of PH, it is likely that the slightly reinforced brachyptery of K. dimorpha is combined with flightlessness. The species occurs in rocky habitat on siliceous soil. Vertical distribution: about 2800 m.

Remark

Fringes of the examined females seem partially lost and thus may lead to a biased impression of the extent of wing reduction.

Kessleria alpmaritimae sp. n.

Type material

Holotype. ♂, „FRANKREICH Dep. Alpes Maritimes Marguareis W-Hang Navela 2100–2200 m 21.–23.7.1990“„leg. Huemer, Tarmann“„YPO 79 ♂ P. Huemer“ (TLMF).

Paratypes. France: 9 ♂, 5 ♀, same data, genitalia slide YPO 55 ♂ P. Huemer (TLMF); 7 ♂, 2 ♀, same data, but 18.-19.7.1991, DNA barcode IDs TLMF Lep 01850, TLMF Lep 01851, TLMF Lep 03100, TLMF Lep 03101, TLMF Lep 03102, TLMF Lep 03103 (TLMF); 3 ♂, 5 ♀, same data, but 23.7.1990 (TLMF); 1 ♂, same data, but Punta Marguareis, 2450–2650 m, 23.7.1990 (TLMF).

Diagnosis

K. alpmaritimae resembles other taxa of K. alternans-group in wing markings and colour (Figs 211), but the male with average forewing length of only 8.05 mm (n=6) is distinctly smaller than K. wehrlii with 8.75 mm (n=13) and K. alternans with 8.55 mm (n=26) and larger than K. cottiensis with only 7.59 mm (n=14). K. dimorpha with an average forewing length of 8.25 mm (n=6) is similar in size, but clearly differs by the largely reduced whitish suffusion of the forewing, a character stage which also applies to K. cottiensis and K. alternans, whereas K. wehrlii is intensely mottled whitish. The female with forewing length of only 6.06 mm (n=3) is distinctly smaller than that of K. alternans with 7.58 mm (n=11), but hardly separable from other species in size. Compared to the genetically nearest neighbour K. dimorpha, the hindwings are less reduced and the ground colour of the forewing is much lighter. The female of K. wehrlii is insufficiently described due to limited material, and the females of K. cottiensis and K. alpmaritimae are indistinguishable. The male genitalia differ from K. cottiensis by the medially weakly widened valva, from K. alternans by the more slender cornuti, from the nearest neighbour K. wehrlii by three instead of four cornuti, and from K. dimorpha by the distinctly longer phallus (1.52–1.58 mm vs. 1.32–1.36 mm) (Figs 1221).

The female genitalia show no diagnostic characters to related species of the K. alternans-group (Figs 2231).

Description

Male (Fig. 10). Head covered with erected whitish hair-like scales; antennae dark grey-brown, indistinctly lighter ringed; thorax and tegulae dark grey-brown. Forewing length 7.0–8.5 mm (Ø 8.05 mm; n=6); ground colour blackish to dark grey-brown, intensely mottled with whitish scales, particularly from basal fifth to 4/5; veins with distinct black dots, particularly along costa, subcosta, cubital and anal veins, medial and radial veins with indistinct brown lines; oblique blackish fascia at about 1/3 to 1/2 indistinct; termen dark grey-brown; fringes light grey, with slightly darker base and indistinct fringe linge. Hindwing dark grey, fringes light greyish-white with slightly darker base.

Female (Fig. 11). Head covered with erected whitish hair-like scales; antennae grey-brown, indistinctly ringed whitish; thorax and tegulae whitish. Forewing length 5.9–6.3 mm (Ø 6.06 mm; n=3); ground colour whitish, mottled with few black scales, particularly along veins, patches of black scales near base and at distal end of cell; oblique blackish fascia at about 1/3 to 1/2 distinct, separated into two dashes; termen whitish with some blackish dots in apical part; fringes white. Hindwing grey, fringes whitish-grey.

Male genitalia (Figs 20–21). Socii long and slender, with apical spine; anterior margin of tegumen with medial process; gnathos broadly tongue-shaped, smooth; valva moderately slender, length 0.72–0.76 mm, medially weakly widened, maximum width of about 0.26–0.27 mm, densely covered with long hairs in medial part and short setae on ventromedial margin, ventromedial part weakly convex, distal part slender with ventrally convex and dorsally projected apex, costa strongly sclerotized with weak distal dentation; sacculus oval, weakly confined, densely covered with strong setae; saccus short, about 0.32–0.33 mm, stout, about same width throughout, apex rounded; phallus ca. 1.52–1.58 mm long and slender, straight, apically with distinct reticulate sculpture, uneverted vesica with ca. 0.62–0.66 mm long sclerotized part, three prominent needle-shaped, one single, the other basally connected, cornuti of about 0.42–0.44 mm in length.

Female genitalia (Figs 26, 31). Genitalia ca. 4.9 mm in length; papilla analis large, densely covered with long setae; apophysis posterior rod like, ca. 0.76 mm, about length of apophysis anterior; apophysis anterior rod like; posterior part bifurcated with straight dorsal and inwardly curved ventral branch; lamella postvaginalis with large sclerotized mediolateral patches, covered with microtrichia, medial area less sclerotized, posterolateral part with hump, covered with some long setae; ostium bursae membranous; antrum weakly sclerotized, funnel-shaped; ductus bursae very long, ca. 2.4 mm, from entrance of ductus seminalis to transition into corpus bursae covered with finely granulous sculpture, particularly in posterior and anterior part, ductus bursae extended into posterior part of corpus bursae, entrance to corpus bursae weakly widened; corpus bursae well delimited, about 1.2 mm in length, ovoid, with largely reduced plate-like signum.

Molecular data

The average intraspecific divergence of the barcode region is 0.0% (n=6). The minimum distance to the nearest neighbour K. wehrlii is 1.87%, whereas the minimum divergence to K. cottiensis, K. dimorpha and K. alternans ranges from 2.98% and 3.64% to 3.75%, respectively.

Etymology

The species name is a made-up word which refers to the area of the type locality, the Alpes Maritimes.

Distribution

(Fig. 32). Only known from the type locality, the Marguareis Massif, in the French Alpes Maritimes.

Ecology

Host-plant and early stages unknown. The adults have been collected in the last third of July during the day, flying freely in the morning hours and flushed out from their resting places with a bee-smoker. The species occurs in rocky habitat on calcareous soil. Vertical distribution: from about 2100 m to 2650 m.

Remarks

K. alpmaritimae described here was already suspected to be distinctive from K. alternans by Huemer and Tarmann (1992), who illustrated adults (Figs 8–9).

Kessleria apenninica-group

The K. apenninica-group s.str. only includes the new species K. apenninica which is characterized e.g. by slender forewings. From characters of the male genitalia, such as the short cornuti, closer relatives are likely to be the Iberian K. diabolica, K. brevicornuta, K. brachypterella and K. pyrenaea, which all differ in adult morphology (see Figs 3338, and Huemer and Tarmann 1992). These species together with K. apenninica may form a larger species-group, but at present material is scarce and supporting molecular data are lacking. The nearest neighbour of the new species with a full DNA barcode is tentatively attached to K. pyrenaea and considered for the differential diagnosis.

Kessleria apenninica sp. n.

Type material

Holotype. ♂, „Italia Prov. Rieti Monte Terminillo 13°00,6'E, 42°29,0'N 1730–1780 m, 16.7.2010 leg. Huemer TLMF 2010-020“ „YPO 147 ♂ P. Huemer“ „TLMF Lep 01662“ (TLMF).

Paratypes. Italy: 1 ♂, same data, DNA barcode ID TLMF Lep 01661 (TLMF); 2 ♂, Prov. L´Aquila, NP Gran Sasso, ex Miniera di Lignite, 13°42,8'E, 42°25,6'N, 1750 m, 14.-15.7.2010, leg. Huemer, genitalia slide YPO 148 ♂ P. Huemer, DNA barcode IDs TLMF Lep 01663, TLMF Lep 01664 (TLMF).

Diagnosis

K. apenninica is characterized by unusually slender forewings and a pure white colour with black pattern. Species from the K. apenninica-group are externally unmistakably distinguishable from one another both by wing pattern and colour (Figs 33–34, and Huemer and Tarmann 1992), but the genitalia of males are similar (Figs 35–38). However, in K. apenninica the saccus is distinctly shorter than in all other species with 0.23 vs. minimum 0.29 mm.

Description

Male (Fig. 33). Head covered with whitish hair-like scales; antennae almost unicolorous dark grey with light grey apex; thorax and tegulae mixed dark grey and whitish. Forewing length 5.8–6.9 mm (Ø 6.4 mm; n=4); forewing slender; ground colour white, mottled with black; black dots on veins and in terminal area; black patch near base and oblique blackish fascia at about 1/3 to 1/2; fringes white with indistinct dark grey fringe line. Hindwing dark grey, fringes with dark grey base, distal part white.

Female. Unknown.

Male genitalia (Figs 35–36). Socii long and slender, with apical spine; anterior margin of tegumen with medial process; gnathos broadly tongue-shaped, smooth; valva slender, length 0.61 mm, max width 0.19 mm; densely covered with long hairs in medial part and short setae on ventromedial margin, ventromedial part weakly convex, distal part with ventrally convex and dorsally projected apex, costa strongly sclerotized with indistinct distal dentation; sacculus oval, weakly confined, densely covered with strong setae; saccus short, about 0.24 mm, slender, about same width throughout, apex rounded; phallus 0.95 mm long, slender, uneverted vesica with ca. 0.35 mm long sclerotized apical part, including 3 short cornuti of about 0.22 mm length [hardly discernible in situ].

Female genitalia. Unknown.

Molecular data

K. apenninica splits into two geographically separated haplogroups, which in our examination – based on limited material – did not reveal any morphological differences. The average intraspecific divergence of the barcode region is considerable with 1.05%, ranging from a minimum of 0% to a maximum of 1.69% (n=4). The minimum distance to the nearest neighbour K. pyrenaea is 5.47%.

Etymology

The species name refers to the Apennines where all type specimens have been collected.

Distribution

Only known from the Apennines in Central Italy.

Ecology

Host-plant and early stages unknown, but the species probably feeds on an unidentified broad-leaved Saxifraga species growing on steep rocks. The adults have been collected in the last third of July from light. The species occurs in rocky habitat on calcareous soil. Vertical distribution: from about 2100 m to 2200 m.

Kessleria albescens-group

The K. albescens-group is characterized by small and predominantly whitish-coloured species without obvious sexual dichroism or dimorphism (Figs 3948). The male genitalia are recognizable by the strongly spinous sacculus, the long and stout saccus, and particularly the phallus with two long cornuti with bases of similar length (Figs 4958). The female genitalia are characterized by the curved entrance of the ductus bursae and the finely granulated sculpture of the entire ductus bursae (Figs 5966). Larval host-plants, as far as is known, belong to broad-leaved Saxifraga spp., e.g. S. paniculata and S. incrustata. Five species are known: K. albescens, K. inexpectata, K. helvetica, K. klimeschi and the new species K. orobiae.

Figures 39–44.

Kessleria adults in dorsal view. 39 K. klimeschi, ♂, holotype, Italy, Prov. Udine, Montasio, Malga Pecol, 1600 m, 24.6.1989 e.l., leg. Huemer & Tarmann (TLMF) 40 K. klimeschi, ♀, paratype, same data, but 3.7.1989 e.l. (TLMF) 41 K. helvetica, ♂, holotype, Switzerland, Wallis, Zermatt, 1850 m, 10.8.1980, leg. Whitebread (DNA barcode ID TLMF Lep 01868) (TLMF) 42 K. helvetica, ♀, Switzerland, Wallis, Zermatt, Triftschlucht, 1820 m, 10.6.2014 e.l., leg. Schmid (DNA barcode ID TLMF Lep 14996) (TLMF) 43 K. inexpectata, ♂, paratype, France, Dep. Alpes Maritimes, Marguareis W-Hang, Navela, 2100–2200 m, 21.–23.7.1990, leg. Huemer & Tarmann (TLMF) 44 K. inexpectata, ♀, paratype, same data (TLMF).

Figures 45–48.

Kessleria adults in dorsal view. 45 K. orobiae sp. n., ♂, paratype, Zambla Alta – Plassa, 1160 m, 24.6.2013 leg. Mayr (coll. Mayr) 46 K. orobiae sp. n., ♀, paratype, same data (coll. Mayr) 47 K. albescens, ♂, Italy, Monte Baldo, Bocca di Navene, 14.7.1987, leg. Huemer & Tarmann (DNA barcode ID TLMF Lep 03131) (TLMF) 48 K. albescens ♀, same data, but 10.9.1987 e.l. (DNA barcode ID TLMF Lep 01866) (TLMF).

Kessleria orobiae sp. n.

Type material

Holotype. ♂, „ITALIA sept. Prov. Bergamo, Alpi Orobie Zambla Alta – Plassa 9°47'48"E, 45°54'12"N 1160 m, 24.6.2013 leg. Huemer“ „DNA Barcode TLMF Lep 09972“ „YPO 160 ♂ P. Huemer“ (TLMF).

Paratypes. Italy: 6 ♂, 6 ♀, same data, DNA barcode IDs TLMF Lep 09971, TLMF Lep 09973 (TLMF); 1 ♂, 1 ♀, same data, but e.l. 11.7.2013 (TLMF); 7 ♂, 2 ♀, same data, but leg. Mayr (coll. Mayr, Feldkirch); 1 ♂, Prov. Bergamo, Alpi Orobie, Val d´Arera, 2000 m, 14.–15.8.1992, leg. Huemer (TLMF); 1 ♀, Prov. Bergamo, Alpi Orobie, W. Ca. San Marco, 2100 m, e.l. 31.7.1992, leg. Huemer & Tarmann, DNA barcode ID TLMF Lep 03175 (TLMF).

Diagnosis

K. orobiae largely resembles other taxa of the K. albescens-group in wing markings and colour, and cannot be unambiguously separated (Figs 3948). Similarly, female genitalia exhibit no significant diagnostic characters for discrimination at species level (Figs 5966, and Huemer and Tarmann 1992). The most reliable diagnostic characters in the species-group are found in the male genitalia (Figs 4958). K. orobiae differs from K. klimeschi, K. albescens and K. helvetica by a much shorter saccus (0.32 mm vs. minimum 0.38–0.58 mm) which is only about half the length of the valva compared to the minimum 0.75 times the length of the valva in the other species. In K. inexpectata the saccus is slightly longer and furthermore without the apical widening of K. orobiae, K. albescens and K. helvetica. The two needle-shaped cornuti are similar in all species, with the exception of K. klimeschi with only one needle-shaped and one sub-ovate cornutus, and K. helvetica with cornuti of about 0.50 mm in length. The female genitalia largely resemble other species of the K. albescens-group with only quantitative differences, such as a longer ductus bursae than in K. albescens (1.6 mm vs. 1.3–1.4 mm), and the overall length of genitalia which exceeds K. klimeschi (4.2 mm vs. 3.5 mm).

Figures 49–54.

Kessleria male genitalia. 49 K. klimeschi, holotype, Italy, Prov. Udine, Montasio, Malga Pecol, 1600 m, 24.6.1989 e.l., leg. Huemer & Tarmann, genitalia slide YPO 17 (TLMF) 50 idem, cornuti enlarged 51 K. helvetica, holotype, Switzerland, Wallis, Zermatt, 1850 m, 10.8.1980, leg. Whitebread, gen. slide 350 Whitebread (DNA barcode ID TLMF Lep 01868) (TLMF) 52 idem, distal part of phallus enlarged 53 K. inexpectata, paratype, France, Dep. Alpes Maritimes, Marguareis W-Hang, Navela, 2100–2200 m, 21.–23.7.1990, leg. Huemer & Tarmann, gen. slide YPO 63 (TLMF) 54 idem, distal part of phallus enlarged.

Figures 55–58.

Kessleria male genitalia. 55 K. orobiae sp. n., holotype, Zambla Alta – Plassa, 1160 m, 24.6.2013 leg. Huemer, gen. slie YPO 160 (TLMF) 56 idem, distal part of phallus enlarged 57 K. albescens, ♂, Italy, Monte Baldo, Bocca di Navene, 14.7.1987, leg. Huemer & Tarmann, gen. slide YPO 19 (DNA barcode ID TLMF Lep 03131) (TLMF) 58 idem, distal part of phallus enlarged.

Figures 59–62.

Kessleria female genitalia. 59 K. klimeschi, paratype, Italy, Prov. Udine, Montasio, Malga Pecol, 1600 m, 24.6.1989 e.l., leg. Huemer & Tarmann, gen. slide YPO 76 (TLMF) 60 K. inexpectata, paratype, France, Dep. Alpes Maritimes, Marguareis W-Hang, Navela, 2100–2200 m, 21.–23.7.1990, leg. Huemer & Tarmann, gen. slide YPO 74 (TLMF) 61 K. orobiae sp. n., paratype, Zambla Alta – Plassa, 1160 m, 24.6.2013, leg. Huemer, gen. slide YPO 161 (TLMF) 62 K. albescens, Italy, Monte Baldo, Bocca di Navene, 14.7.1987, leg. Huemer & Tarmann (TLMF).

Figures 63–66.

Kessleria female genitalia, details of VIII abdominal segment enlarged. 63 K. klimeschi, paratype, Italy, Prov. Udine, Montasio, Malga Pecol, 1600 m, 24.6.1989 e.l., leg. Huemer & Tarmann, gen. slide YPO 76 (TLMF) 64 K. inexpectata, paratype, France, Dep. Alpes Maritimes, Marguareis W-Hang, Navela, 2100–2200 m, 21.-23.7.1990, leg. Huemer & Tarmann, gen. slide YPO 74 (TLMF) 65 K. orobiae sp. n., paratype, Zambla Alta – Plassa, 1160 m, 24.6.2013, leg. Huemer, gen. slide YPO 161 (TLMF) 66 K. albescens, Italy, Monte Baldo, Bocca di Navene, 14.7.1987, leg. Huemer & Tarmann (TLMF).

Description

Male (Fig. 45). Head covered with white hair-like scales; antennae ringed dark grey and whitish; thorax and tegulae mixed dark grey and white, distally predominantly white, particularly tegulae. Forewing length 6.7–7.3 mm (Ø 7.03 mm; n=6); ground colour whitish-grey, intensively mottled with blackish-grey spots all over wing, few ochre-brown scales in dorsal part; blackish-grey patches at base of costa and at end of cell, oblique blackish-grey fascia at about 1/3 to 1/2 narrow and indistinct, medially separated; termen whitish-grey; fringes white, basally with distinct blackish-grey cilia line, apex with small dark grey tip. Hindwing dark grey, fringes with dark grey base, distal part white.

Female (Fig. 46). As male. Head covered with white hair-like scales; antennae ringed dark grey and whitish; thorax and tegulae mixed dark grey and white, distally predominantly white, particularly tegulae. Forewing length 5.9–6.6 mm (Ø 8.18 mm; n=5); ground colour whitish-grey, intensively mottled with blackish-grey spots all over wing, few ochre-brown scales in dorsal part; blackish-grey patches at base of costa and at end of cell, oblique blackish-grey fascia at about 1/3 to 1/2 narrow and indistinct, medially separated; termen whitish-grey; fringes white, basally with distinct blackish-grey cilia line, apex with small dark grey tip. Hindwing dark grey, fringes with dark grey base, distal part white.

Male genitalia (Figs 55–56). Socii long and slender, with apical spine; anterior margin of tegumen with medial process; gnathos broadly tongue-shaped, smooth; valva slender, length 0.60 mm, maximum width 0.18 mm; densely covered with long hairs in medial part and short setae on ventromedial margin, apical part ventrally convex, costa strongly sclerotized without dentation; sacculus oval, weakly confined, densely covered with strong setae; saccus moderate in length, about 0.32 mm, stout, distally widened with rounded apex; phallus 1.34 mm long, slender, uneverted vesica with ca. 0.70 mm long sclerotized apical part, including two prominent needle-shaped cornuti of about 0.40 mm length.

Female genitalia (Figs 61, 65). Genitalia ca. 4.2 mm in length; papilla analis large, densely covered with long setae; apophysis posterior rod like, ca. 0.70 mm, about length of apophysis anterior; apophysis anterior rod like; posterior part bifurcated with straight dorsal and inwardly curved ventral branch; ventral branch descending into patch like sclerite; lamella postvaginalis with sclerotized lateral patches, covered with microtrichia, medially membranous, posterolaterally with hump, covered with some long setae; ostium bursae membranous; antrum weakly developed, ring-like; ductus bursae long, about 1.6 mm, weakly curved in posterior part, from entrance of ductus seminalis to almost transition to corpus bursae densely covered with finely granulous sculpture; sculpture in posterior part nearly bacillary, distally increasingly granulous, entrance to corpus bursae widened, without sculpture; corpus bursae about 1.7 mm in length, saccate, posterior part folded, without signum.

Molecular data

The average intraspecific divergence of the barcode region is low with 0.31%, ranging from a minimum of 0.15% to a maximum of 0.46% (n=5). The minimum distance to the nearest neighbour K. albescens is 2.66%, whereas the minimum divergence to K. inexpectata, K. helvetica and K. klimeschi ranges from 3.14% and 3.46% to 9.53%, respectively.

Etymology

The species name refers to the Orobian Alps (Alpi Orobie) in northern Italy, where the type locality is situated.

Distribution

(Fig. 67). Only known from Zambla Alta – Plassa and few nearby localities in the Orobian Alps (Prov. Bergamo, Italy).

Figure 67.

Distribution pattern of the Kessleria albescens-group from examined material.

Ecology

The larval habits are insufficiently known, but based on our observations, the larva lives in the shoots and as a leaf-miner in basal leaves of Saxifraga paniculata and Saxifraga sp. Mined leaves are partially spun together and covered with a fine silken web. The adults have been collected from the Saxifraga-cushions or nearby rock during the day. In the first few hours of the night they have been observed with a head-lamp flying actively around the larval habitat or sitting near the host-plant. The adult is on the wing from late June to mid-August, depending on altitude and snow coverage. Bred specimens date from mid to late July. K. orobiae occurs in rocky habitat both on calcareous and silicous soil. Vertical distribution: from about 1100 m to 2100 m.

Established European species of Kessleria

A brief overview of established species lists including original description, type locality, type material, references of published images of adults and images of genitalia, and hitherto unpublished molecular data. For extensive generic and species descriptions and diagnoses, see Huemer and Tarmann (1992).

Genus Kessleria Nowicki, 1864

Kessleria Nowicki 1864: 12. Type species: Kessleria zimmermanni Nowicki, 1864 by monotypy and original designation.

Subgenus Kessleria Nowicki, 1864
Kessleria alpicella (Stainton, 1851)

Tinea (Oecophora!) alpicella [Fischer von Röslerstamm, Mann in litt.] Stainton 1851: 17. Type locality: Europe [?Austria]. Type material: Lectotype (designated by Huemer and Tarmann 1992: 16) (BMNH) [examined].

Swammerdamia alpicella Herrich-Schäffer 1855: 272. Type locality: Austria, Niederösterreich, Schneeberg. Type material: Syntypes [not traced]. Homonym and synonym.

Redescription and diagnosis see Huemer and Tarmann 1992: 16–18, figs 1–2 (adult), figs 85–93 (male genitalia), figs 220–221 (female genitalia).

Molecular data. K. alpicella splits into three geographically separate haplogroups, indicating potential cryptic diversity. The average intraspecific divergence of the barcode region is high with 1.52%, ranging from a minimum of 0% to a maximum of 4.27% (n=12). The minimum distance to the nearest neighbour K. wehrlii is 6.9%.

Kessleria mixta Huemer & Tarmann, 1992

Kessleria mixta Huemer and Tarmann 1992: 18. Type locality: Albania, Korab. Type material: Holotype (designated by Huemer and Tarmann 1992: 19) (NHMV) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 18–19, Fig. 3 (adult), figs 94–96 (male genitalia).

Molecular data. Unavailable.

Remarks. Female unknown.

Kessleria alternans (Staudinger, 1871)

Kessleria alternans Staudinger 1871: 291. Type locality: Switzerland, Graubünden, Sils-Maria. Type material: Lectotype (designated by Friese 1960: 75) (MNHU) [examined].

Redescription and diagnosis see Huemer and Tarmann 1992: 19–22, figs 4–6 (adult), figs 97–102 (male genitalia), figs 223–224 [misidentified figures depicting newly described species in this paper are not listed].

Molecular data. The intraspecific divergence of the barcode region is low, ranging from a minimum of 0% to a maximum of 0.32% (mean 0.12%) (n=10). The minimum distance to the nearest neighbour K. cottiensis is 2.65%.

Remarks. Huemer and Tarmann (1992) already recognized and described a remarkable amount of individual and geographical variation. At that time, this variation was considered as intraspecific, and the authors hesitated to describe further species.

Kessleria wehrlii Huemer & Tarmann, 1992

Kessleria wehrlii Huemer and Tarmann 1992: 22. Type locality: France, Dep. Alpes-Maritimes, Mont Colomb. Type material: Holotype (designated by Huemer and Tarmann 1992: 23) (TLMF) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 22–23, figs 11–12 (adult), figs 109–111 (male genitalia), fig. 211 (8th abdominal segment), fig. 222 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is 0.0% (n=4). The minimum distance to the nearest neighbour K. alpmaritimae is 1.87%.

Kessleria nivescens Burmann, 1980

Kessleria nivescens Burmann 1980: 105. Type locality: Italy, Prov. Verona, Monte Baldo. Type material: Holotype (designated by Burmann 1980: 107) (SMNK) [examined].

Redescription and diagnosis see Huemer and Tarmann 1992: 23–26, figs 13–17 (adult), figs 115–120 (male genitalia), fig. 219 (female genitalia).

Molecular data. K. nivescens splits into three geographically separate haplogroups. The intraspecific divergence of the barcode region is high, ranging from a minimum of 0% to a maximum of 2.5% (mean 1.09%) (n=14). The minimum distance to the nearest neighbour K. petrobiella is 3.29% (mean 4.52%, max. 4.92%).

Remarks. Huemer and Tarmann (1992) already recognized and described a considerable amount of individual and geographical variation, particularly in phenotypic appearance, but the authors hesitated to describe further species due to the lack of diagnostic genitalia characters. Molecular data suggest possible cryptic diversity, but further investigations are required.

Kessleria petrobiella (Zeller, 1868)

Scythropia petrobiella Zeller 1868: 607. Type locality: Slovenia, [Log Pod Mangrtom]. Type material: Lectotype (designated by Huemer and Tarmann 1992: 37) (BMNH) [examined].

Redescription and diagnosis see Huemer and Tarmann 1992: 37–39, figs 32–33 (adult), figs 156–161 (male genitalia), fig. 229 (female genitalia).

Molecular data. The intraspecific divergence of the barcode region is 0% (n=4). The minimum distance to the nearest neighbour K. nivescens is 3.29%.

Kessleria macedonica Huemer & Tarmann, 1992

Kessleria macedonica Huemer and Tarmann 1992: 26. Type locality: Kosovo/Macedonia, Shar Planina, Crni vrh. Type material: Holotype (designated by Huemer and Tarmann 1992: 26) (ZSM) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 26–27, fig. 18 (adult), figs 121–123 (male genitalia).

Molecular data. Unavailable.

Remarks. Female unknown.

Kessleria albanica Friese, 1960

Kessleria albanica Friese 1960: 68. Type locality: Albania, Nikai. Type material: Holotype (designated by Friese 1960: 68) (ZSM) [examined].

Redescription and diagnosis see Huemer and Tarmann 1992: 27–28, fig. 19 (adult), figs 124–125 (male genitalia).

Molecular data. K. albanica splits into three geographically separate haplogroups, indicating potential cryptic diversity. The average intraspecific divergence of the barcode region is high with 2.08%, ranging from a minimum of 0% to a maximum of 3.12% (n=5). The minimum distance to the nearest neighbour K. burmanni is 9.29%.

Remarks. Female unknown.

Kessleria burmanni Huemer & Tarmann, 1992

Kessleria burmanni Huemer and Tarmann 1992: 28. Type locality: Austria, Nordtirol, Nordkette. Type material: Holotype (designated by Huemer and Tarmann 1992: 30) (TLMF) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 28–30, figs 20–22 (adult), figs 126–131 (male genitalia), fig. 225 (female genitalia), figs 63–65 (SEM egg structures).

Molecular data. The average intraspecific divergence of the barcode region is 0.0% (n=6). The minimum distance to the nearest neighbour K. hauderi is 7.61%.

Kessleria insubrica Huemer & Tarmann, 1993

Kessleria insubrica Huemer and Tarmann 1993: 41. Type locality: Italy, Prov. Bergamo, Val d´Arera. Type material: Holotype (designated by Huemer and Tarmann 1993: 45) (TLMF) [examined].

Description and diagnosis see Huemer and Tarmann 1993: 41–46, fig. 1 (adult), figs 5–6 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is low with 0.08%, ranging from a minimum of 0% to a maximum of 0.15% (n=4). The minimum distance to the nearest neighbour K. burmanni is 8.95%.

Remarks. Male unknown.

Kessleria hauderi Huemer & Tarmann, 1992

Kessleria hauderi Huemer and Tarmann 1992: 31. Type locality: Austria, Steiermark, Eisenerzer Reichenstein. Type material: Holotype (designated by Huemer and Tarmann 1992: 32) (TLMF) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 21–33, figs 23–24 (adult), figs 132–137 (male genitalia), fig. 227 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is 0% (n=2). The minimum distance to the nearest neighbour K. burmanni is 7.61%.

Kessleria diabolica Huemer & Tarmann, 1992

Kessleria diabolica Huemer and Tarmann 1992: 33. Type locality: Spain, Prov. Avila, Sierra de Gredos. Type material: Holotype (designated by Huemer and Tarmann 1992: 33) (ZMUC) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 33, fig. 25 (adult), figs 138–140 (male genitalia).

Molecular data. Unavailable.

Remarks. Female unknown.

Kessleria brevicornuta Huemer & Tarmann, 1992

Kessleria brevicornuta Huemer and Tarmann 1992: 34. Type locality: Spain, Prov. Avila, Sierra de Gredos. Type material: Holotype (designated by Huemer and Tarmann 1992: 34) (coll. Arenberger, Vienna) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 34–35, figs 26–27 (adult), figs 141–143 (male genitalia), fig. 235 (female genitalia).

Molecular data. Unavailable.

Kessleria pyrenaea Friese, 1960

Kessleria pyrenaea Friese 1960: 76. Type locality: France, Dép. Pyrénées-Orientales, Mt. Canigou. Type material: Holotype (designated by Friese 1960: 76) (NHMV) [examined].

Redescription and diagnosis see Huemer and Tarmann 1992: 35, fig. 28 (adult), figs 144–146 (male genitalia).

Molecular data. The intraspecific divergence of the barcode region is unknown (n=1). The minimum distance to the nearest neighbour K. apenninica is 5.47%.

Remarks. Female unknown. The identity of the sequenced specimen is doubtful.

Kessleria brachypterella Huemer & Tarmann, 1992

Kessleria brachypterella Huemer and Tarmann 1992: 36. Type locality: France, Dép. Hautes-Pyrénées, Pic du Midi de Bigorre. Type material: Holotype (designated by Huemer and Tarmann 1992: 34) (BMNH) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 36–37, figs 29–30 (adult), figs 147–149 (male genitalia), fig. 228 (female genitalia).

Molecular data. Unavailable.

Kessleria zimmermanni Nowicki, 1864

Kessleria zimmermanni Nowicki 1864: 13. Type locality: Poland, Tatra mts., ?Kopa Magury. Type material: Lectotype (designated by Huemer and Tarmann 1992: 39) (SDEI) [examined].

Kessleria tatrica Friese 1960: 71. Type locality: [?Poland], Tatra mts. Type material: Holotype (designated by Friese 1960: 71) (NHMV).

Redescription and diagnosis see Huemer and Tarmann 1992: 39–42, figs 32–33 (adult), figs 162–167 (male genitalia), fig. 217 (8th abdominal segment), fig. 226 (female genitalia).

Molecular data. The intraspecific divergence of the barcode region is 0% (n=3). The minimum distance to the nearest neighbour K. petrobiella is 5.73%.

Kessleria albomaculata Huemer & Tarmann, 1992

Kessleria albomaculata Huemer and Tarmann 1992: 42. Type locality: France, Dép. Hautes-Pyrénées, Cauterets. Type material: Holotype (designated by Huemer and Tarmann 1992: 42) (MNCN) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 42–43, fig. 36 (adult), figs 153–155 (male genitalia), fig. 216 (8th abdominal segment).

Molecular data. The intraspecific divergence of the barcode region is unknown (n=1). The minimum distance to the nearest neighbour K. petrobiella is 6.76%.

Remarks. Female unknown.

Kessleria caflischiella (Frey, 1880)

Swammerdamia caflischiella Frey 1880: 344. Type locality: Switzerland, Wallis, ?Gamsen. Type material: Holotype (designated by Frey 1880: 344) (BMNH) [examined].

Redescription and diagnosis see Huemer and Tarmann 1992: 49–51, figs 45–46 (adult), figs 191–196 (male genitalia), fig. 229 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is low with 0.04%, ranging from a minimum of 0% to a maximum of 0.15% (n=8). The minimum distance to the nearest neighbour K. alpmaritimae is 6.39%.

Kessleria klimeschi Huemer & Tarmann, 1992

Kessleria klimeschi Huemer and Tarmann 1992: 47. Type locality: Italy, Prov. Udine. Montasio, Malga Pecol. Type material: Holotype (designated by Huemer and Tarmann 1992: 48) (TLMF) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 47–49, figs 43–44 (adult), figs 185–190 (male genitalia), fig. 218 (8th abdominal segment), fig. 233 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is low with 0.06%, ranging from a minimum of 0% to a maximum of 0.15% (n=5). The minimum distance to the nearest neighbour K. inexpectata is 8.83%.

Kessleria helvetica Huemer & Tarmann, 1992

Kessleria helvetica Huemer and Tarmann 1992: 46. Type locality: Switzerland, Wallis, Zermatt. Type material: Holotype (designated by Huemer and Tarmann 1992: 47) (TLMF) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 46–47, figs 41–42 (adult), Figs 179–184 (male genitalia), fig. 232 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is unknown (n=1). K. helvetica overlaps in the barcode with a haplogroup of topotypical K. inexpectata, but diagnostic morphological characters indicate species status. The minimum distance to a further haplogroup of K. inexpectata is 1.77%.

Kessleria inexpectata Huemer & Tarmann, 1992

Kessleria inexpectata Huemer and Tarmann 1992: 45. Type locality: France, Dep. Alpes Maritimes, Marguareis. Type material: Holotype (designated by Huemer and Tarmann 1992: 46) (TLMF) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 45–46, figs 39–40 (adult), figs 173–178 (male genitalia), fig. 231 (female genitalia).

Molecular data. K. inexpectata splits into two major haplogroups. The average intraspecific divergence of the barcode region within the haplogroup of topotypical specimens is low with 0.16%, ranging from a minimum of 0% to a maximum of 0.32% (n=4) whereas the average intraspecific variation within the second haplogroup is considerable with 0.84% (maximum 1.68%). The mean intraspecific divergence of the entire sample is 1.42% (maximum 2.18%). The haplogroup of the topotypical population overlaps with K. helvetica. The minimum distance to K. orobiae is 3.14%.

Kessleria albescens (Rebel, 1899)

Hofmannia albescens Rebel 1899: 177. Type locality: Italy, South Tyrol, Bozen. Type material: Lectotype (designated by Friese 1960: 72) (NHMV) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 43–44, figs 37–38 (adult), figs 168–172 (male genitalia), fig. 230 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is 0% (n=3). The minimum distance to the nearest neighbour K. orobiae is 2.66%.

Subgenus Hofmannia Heinemann & Wocke, 1877
Kessleria saxifragae (Stainton, 1868)

Zelleria saxifragae Stainton 1868: 139. Type locality: Austria, Nordtirol, Kaisergebirge. Type material: ?Syntypes (examined by Huemer and Tarmann 1992: 53) (BMNH) [examined].

Description and diagnosis see Huemer and Tarmann 1992: 51–54, figs 47–50 (adult), figs 197–202 (male genitalia), fig. 236 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is low with 0.43%, ranging from a minimum of 0% to a considerable maximum of 1.28% (n=20). The minimum distance to the nearest neighbour Zelleria celastrusella Kearfott, 1903, from North America is 6.22%, and the minimum distance to the congeneric K. fasciapennella is 7.21%.

Kessleria fasciapennella (Stainton, 1849)

Zelleria fasciapennella Stainton 1849: 80. Type locality: GB, Scotland, Edinburgh, Pentland hills. Type material: Lectotype (designated by Huemer and Tarmann 1992: 54) (BMNH) [examined].

Kessleria longipenella Friese 1960: 83. Type locality: Russia, Karelia, S Petrosawodsk. Type material. Holotype (designated by Friese 1960: 83) (SDEI).

Description and diagnosis see Huemer and Tarmann 1992: 54–56, figs 52–54 (adult), figs 203–208 (male genitalia), fig. 237 (female genitalia).

Molecular data. The average intraspecific divergence of the barcode region is low with 0.04%, ranging from a minimum of 0% to a maximum of 0.15% (n=8). The minimum distance to the nearest neighbour Zelleria celastrusella Kearfott, 1903, from North America is 6.58%, and the minimum distance to the congeneric K. saxifragae is 7.21%.

Discussion

Our study proves the advantage of an integrative taxonomic approach, initially based on morphology, with molecular data supplemented as an additional tool for delimitation of cryptic species. Even within genera of European Lepidoptera which had been considered as well explored, cryptic diversity seems much more widespread than hitherto estimated. Recent molecular studies have proven the existence of a remarkable amount of cryptic species in several genera or species-groups, e.g. Callisto (Kirichenko et al. 2015), Stigmella (Nieukerken et al. 2012), Olethreutes (Segerer et al. 2010), Elachista (Mutanen et al. 2013), Eulamprotes (Huemer et al. 2013), Sattleria (Huemer and Hebert 2011, Huemer and Timossi 2014), Caryocolum (Huemer et al. 2014) and Coleophora (Baldizzone et al. 2014, Tabell and Baldizzone 2014). Similarly, the proportion of unnamed species in Kessleria is high, adding about 20% to the hitherto described species diversity, not including several further yet unresolved possible candidates of cryptic diversity. We expect to find such additional overlooked taxa in e.g. K. alpicella and K. albanica, but additional material is needed to resolve this. Outside Europe, the species diversity of Kessleria cannot even be estimated at the present time, with the Chinese K. nivosa (Meyrick, 1938) as the only known congeneric species from Asia, and K. parnassiae (Braun, 1940), a close relative of K. fasciapennella, from North America. Particularly the Asian fauna of Kessleria may prove diverse, e.g. indicative of which is an extraordinary diversity of potential Kessleria host-plants in China with 216 out of about 450 worldwide known species of Saxifraga (139 endemic) and 63 out of 70 Parnassia spp. (49 endemic) (Jintang et al. 2001).

Most of the newly described species belong to complexes of closely related species with strictly allopatric distribution patterns. K. cottiensis, K. dimorpha and K. alpmaritimae are morphologically and genetically most similar to K. alternans and K. wehrlii, forming a separate species group in Kessleria (Fig. 32). Similarly, K. orobiae along with K. albescens, K. klimeschi, K. helvetica and K. inexpectata belong to a species-group of cryptic allopatric taxa (Fig. 67). All these taxa are extremely similar in external and internal morphology. This is a phenomenon well known from the related genus Yponomeuta, namely the Y. cagnagellus species-complex which includes morphologically virtually indistinguishable species (Bakker et al. 2008), which furthermore often share DNA barcodes. Evolution and reproductive isolation in this genus was likely driven by specific host-plant associations and sex pheromones (Menken 1981, Menken et al. 1992, Menken 1996, Löfstedt 1991, Turner et al. 2010). Unlike Yponomeuta, barcode sharing seems to be a rare exception in Kessleria, only observed in K. helvetica and K. inexpectata so far, and indicating possible introgression or recent speciation. DNA barcode divergence to the nearest neighbour is considerable in Kessleria with roughly 2–3% distance in sister species, rising to about 6–9% between morphologically well separated taxa (Table 1, Fig. 1). If reflected by at least one supplementing morphological character stage we consider barcode divergence of roughly 2% as taxonomically relevant. These taxa are described as cryptic species and not subspecies, although such decisions are prone to subjectivity (Hausmann and Huemer 2011, Huemer and Mayr in press, Mutanen et al. 2012).

A similar extent of interspecific divergence in allopatric sister species is also known from other Lepidoptera with geographically restricted alpine distribution patterns, e.g. Sattleria (Huemer and Hebert 2011) and Sciadia (Huemer and Hausmann 2009). The timing of radiation of these and Kessleria is unknown, but estimations of substitution rates of COI indicate that divergences of 1.0–2.5% correspond to divergence times of roughly one million years (Kandul et al. 2004, Hausmann et al. 2011). Even though such estimations should be considered with caution, they indicate that several well separated species of Kessleria may have diverged already in the lower Pliocene (5.3 mya–1.8 mya) while others, such as four out of five newly described species, are possibly of younger origin. Speciation in these taxa was likely reinforced by climatic oscillations in the late Pliocene and during the Pleistocene, with unglaciated but highly isolated refugial areas, particularly in the southern Alps, an area well documented as a hotspot for endemic Lepidoptera (Huemer 1998). The widespread female flightlessness may have been crucial for reducing dispersal and interrupting gene flow, particularly in maternally inherited mitochondrial DNA, thereby expediting the speciation processes. Host-plant relationship itself seems of limited importance for speciation processes in Kessleria as host specificity is moderately pronounced and host-plants are regularly much more widespread than their consumers. E.g. Saxifraga paniculata as one of the major host-plants of species of the K. albescens-group is widely distributed in the Alps and other European mountain systems, but the Kessleria spp. are allopatrically distributed in a small section of the southern Alps. Summing up, further in-depth phylogenetic studies will be necessary to finally identify drivers of speciation in Kessleria.

Acknowledgments

We are particularly grateful to Paul Hebert and his team at the Canadian Centre for DNA Barcoding (Guelph, Canada), whose sequencing work was enabled by funding from the Government of Canada to Genome Canada through the Ontario Genomics Institute. We are also grateful to the Ontario Ministry of Research and Innovation and to NSERC for their support of the BOLD informatics platform.

Stefan Heim (TLMF) is acknowledged for his kind assistance with photographic work. Toni Mayr (Feldkirch, Austria), Jürg Schmid (Illanz, Switzerland), Zdenko Tokár (Michalovce, Slovakia) and Christian Wieser (Klagenfurt, Austria) provided material for our examination. Furthermore, we thank all the institutions and private persons who supported earlier work on the genus, and in this context particularly PH´s former co-author Gerhard Tarmann (Innsbruck, Austria) for various kinds of help.

PH is particularly indebted to the Promotion of Educational Policies, University and Research Department of the Autonomous Province of Bolzano - South Tyrol for helping to fund the project “Genetic biodiversity archive - DNA barcoding of Lepidoptera of the central Alpine region (South, East and North Tyrol)”, and to the Austrian Federal Ministry of Science, Research and Economics for funds received in the framework of ABOL (Austrian Barcode of Life).

We thank two anonymous reviewers and the subject editor Erik van Nieukerken for several helpful comments on the manuscript. Last, but not least, we thank Marianna Teräväinen (Helsinki, Finland) for linguistic improvement of the manuscript.

References

  • Burmann K (1980) Eine neue Kessleria-Art aus dem Monte Baldogebiet in Italien (Lepidoptera, Yponomeutidae). Nachrichtenblatt der Bayerischen Entomologen 29: 105–108.
  • Bakker AC, Roessingh P, Menken SBJ (2008) Sympatric speciation in Yponomeuta: No evidence for host plant fidelity. Entomologia Experimentalis et Applicata 128: 240–247. doi: 10.1111/j.1570-7458.2008.00684.x
  • Baldizzone G, Nel J, Landry J-F (2014) Coleophora nepetellae Baldizzone & Nel, a new species of the C. lixella group (Lepidoptera, Coleophoridae) from France and Italy. ZooKeys 459: 119–135. doi: 10.3897/zookeys.459.7983
  • Collins RA, Cruickshank RH (2013) The seven deadly sins of DNA barcoding. Molecular Ecology Resources 13: 969–975.
  • Cracraft J (1989) Speciation and its ontology. The empirical consequences of alternative species concepts for understanding patterns and processes of differentiation. In: Otte D, Endler JA (Eds) Speciation and its Consequences. Sinauer Associates, Sunderland, MA, 28–59.
  • de Queiroz K (1998) The general lineage concept of species, species criteria, and the process of speciation: A conceptual unification and terminological recommendations. In: Howard DJ, Berlocher SH (Eds) Endless forms: Species and speciation. Oxford University Press, New York, 57–75.
  • deWaard JR, Ivanova NV, Hajibabaei M, Hebert PDN (2008) Assembling DNA Barcodes: Analytical Protocols. In: Cristofre M (Ed.) Methods in Molecular Biology: Environmental Genetics. Humana Press Inc., Totowa, USA, 275–293. doi: 10.1007/978-1-59745-548-0_15
  • Frey H (1880) Die Lepidopteren der Schweiz. Leipzig, xxvi + 454 pp.
  • Friese G (1960) Revision der paläarktischen Yponomeutidae unter besonderer Berücksichtigung der Genitalien. Beiträge zur Entomologie 10: 1–131.
  • Hausmann A, Haszprunar G, Hebert PDN (2011) DNA barcoding the geometrid fauna of Bavaria (Lepidoptera): Suc-cesses, surprises, and questions. PLoS ONE 6(2): e17134. doi: 10.1371/journal.pone.0017134
  • Hausmann A, Huemer P (2011) Taxonomic decision as a compromise: Acasis appensata (Eversmann, 1832) in Central Italy – a case of conflicting evidence between DNA barcode and morphology (Lepidoptera: Geometridae). Zootaxa 3070: 61–68.
  • Hebert PDN, Cywinska A, Ball SL, deWaard JR (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London B, 270: 313–321. doi: 10.1098/rspb.2002.2218
  • Hebert PDN, deWaard JR, Landry JF (2009) DNA barcodes for 1/1000 of the animal kingdom. Biology Letters 6: 359–362. doi: 10.1098/rsbl.2009.0848
  • Herrich-Schäffer GAW (1847–1855) Systematische Bearbeitung der Schmetterlinge von Europa, zugleich als Text, Revision und Supplement zu Jakob Hübner´s Sammlung europäischer Schmetterlinge. 5. Die Schaben und Federmotten, 394 pp. Tineides pl. 1–124, Pterophorides pl. 1–7, Micropteryges pl. 1. Regensburg.
  • Huemer P (1998) Endemische Schmetterlinge der Alpen - ein Überblick (Lepidoptera). Stapfia 55: 229–256.
  • Huemer P, Mayr T (in press) Eine neue Unterart von Colostygia kitschelti aus den Bergamasker-Alpen, mit Bemerkungen zum Artkonzept im C. austriacaria-Komplex (Lepidoptera: Geometridae). Wissenschaftliches Jahrbuch der Tiroler Landesmuseen.
  • Huemer P, Tarmann G (1992) Westpaläarktische Gespinstmotten der Gattung Kessleria Nowicki: Taxonomie, Ökologie, Verbreitung (Lepidoptera, Yponomeutidae). Mitteilungen der Münchner Entomologischen Gesellschaft 81(1991): 5–110.
  • Huemer P, Tarmann G (1993) Wissenschaftliche Ergebnisse der lepidopterologischen Forschungen des Tiroler Landesmuseum Ferdinandeum im Alpenraum. 2. Eine neue Kessleria-Art aus den Alpi Orobie (Insecta, Lepidoptera, Yponomeutidae). Veröffentlichungen des Tiroler Landesmuseums Ferdinandeum 73: 41–46.
  • Huemer P, Hausmann A (2009) A new expanded revision of the European high mountain Sciadia tenebraria species group (Lepidoptera: Geometridae). Zootaxa 2117: 1–30.
  • Huemer P, Hebert PDN (2011) Cryptic diversity and morphology of high alpine Sattleria – a case study combining DNA barcodes and morphology (Lepidoptera: Gelechiidae). Zootaxa 2981: 1–22.
  • Huemer P, Elsner G, Karsholt O (2013) Review of the Eulamprotes wilkella species-group based on morphology and DNA barcodes, with descriptions of new taxa (Lepidoptera, Gelechiidae). Zootaxa 3746(1): 069–100.
  • Huemer P, Timossi G (2014) Sattleria revisited: unexpected cryptic diversity on the Balkan Peninsula and in the south-eastern Alps (Lepidoptera: Gelechiidae). Zootaxa 3780(2): 282–296. doi: 10.11646/zootaxa.3780.2.4
  • Huemer P, Karsholt O, Mutanen M (2014) DNA barcoding as a screening tool for cryptic diversity: an example from Caryocolum, with description of a new species (Lepidoptera, Gelechiidae). ZooKeys 404: 91–111. doi: 10.3897/zookeys.404.7234
  • Jintang P, Cuizhi G, Tsue-chih K, Shumei H, Shu-mei H, Zhaofen W et al. (2001) Saxifragaceae. In: Wu ZY, Raven PH (Eds) Flora of China. Vol 8 (Brassicaceae through Saxifragaceae). Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis, 269–452.
  • Kandul NP, Lukhtanov VA, Dantchenko AV, Coleman JWS, Sekercioglu CH, Haig D et al. (2004) Phylogeny of Agrodiaetus Hübner, 1822 (Lepidoptera: Lycaenidae) inferred from mtDNA sequences of COI and COII and nuclear sequences of EF1-α: Karyotype diversification and species radiation. Systematic Biology 53: 278–298. doi: 10.1080/10635150490423692
  • Kirichenko N, Huemer P, Deutsch H, Triberti P, Rougerie R, Lopez-Vaamonde C (2015) Integrative taxonomy reveals a new species of Callisto (Lepidoptera, Gracillariidae) in the Alps. ZooKeys 473: 157–176. doi: 10.3897/zookeys.473.8543
  • Löfstedt C, Herrebout WM, Menken SBJ (1991) Sex pheromones and their potential role in the evolution of reproductive isolation in small ermine moths (Yponomeutidae). Chemoecology 2: 20–28. doi: 10.1007/BF01240662
  • Menken SBJ (1981) Host races and sympatric speciation in small ermine moths, Yponomeutidae. Entomologia Experimentalis et Applicata 30: 280–292. doi: 10.1111/j.1570-7458.1981.tb03111.x
  • Menken SBJ (1996) Pattern and process in the evolution of insect-plant associations: Yponomeuta as an Example. Entomologia Experimentalis et Applicata 80: 297–305. doi: 10.1111/j.1570-7458.1996.tb00940.x
  • Menken SBJ, Herrebout WM, Wieber JT (1992) Small Ermine Moths (Yponomeutidae): Their Host Relation and Evolution. Annual Review of Entomology 37: 41–66. doi: 10.1146/annurev.en.37.010192.000353
  • Meregalli M, Menardo F, Klass K-D, Cervella P (2013) Phylogeny of the Saxifraga-associated species of Dichotrachelus (Insecta: Coleoptera: Curculionidae), with remarks on their radiation in the Alps. Arthropod Systematics & Phylogeny 71(1): 43–68.
  • Mutanen M, Hausmann A, Hebert PDN, Landry J-F, de Waard JR, Huemer P (2012) Allopatry as a Gordian knot for taxonomists: Patterns of DNA barcode divergence in arctic-alpine Lepidoptera. PLoS ONE 7(10): e47214. doi: 10.1371/journal.pone.0047214
  • Mutanen M, Kaila L, Tabell J (2013) Wide-ranging barcoding aids discovery of one-third increase of species richness in presumably well-investigated moths. Scientific Reports 3: 2901. doi: 10.1038/srep02901
  • van Nieukerken EJ, Mutanen M, Doorenweerd C (2012) DNA barcoding resolves species complexes in Stigmella salicis and S. aurella species groups and shows additional cryptic speciation in S. salicis (Lepidoptera: Nepticulidae). Entomologisk Tidskrift 132: 235–255.
  • Nowicki M (1864) Microlepidopterorum Species novae. Cracoviae, 32 pp., 1 pl.
  • Padial JM, Miralles A, De La Riva I, Vences M (2010) The integrative future of taxonomy. Frontiers in Zoology 7: 16. doi: 10.1186/1742-9994-7-16
  • Rebel H (1899) Zweiter Beitrag zur Lepidopteren-Fauna Südtirols. Verhandlungen der kaiserlich-königlichen zoologisch-botanischen Gesellschaft in Wien 49: 158–185, pl. 4.
  • Robinson GS (1976) The preparation of slides of Lepidoptera genitalia with special reference to the Microlepidoptera. Entomologist’s Gazette 27: 127–132.
  • Sattler K (1991) A review of wing reduction in Lepidoptera. Bulletin of the British Museum of Natural History (Entomology) 60: 243–288.
  • Segerer AH, Haslberger A, Grünewald T (2010) Olethreutes subtilana (Falkovich, 1959): Unexpected occurrence of an ‘eastern’ leaf roller in Central Europe, uncovered by DNA barcoding (Tortricidae: Olethreutinae). Nota lepidopterologica 33: 197–206.
  • Stainton HT (1851) A supplementary catalogue of the British Tineidae & Pterophoridae. London, iv + 28 pp.
  • Stainton HT (1868) A few Observations on the Synonymy of Tinea (?) alpicella, and Zelleria saxifragae, (n. sp.). The Transactions of the Entomological Society of London 1868: 137–139. doi: 10.1111/j.1365-2311.1868.tb00620.x
  • Staudinger O (1871) Beschreibung neuer Lepidopteren des europäischen Faunengebiets (Schluß). Berliner Entomologische Zeitschrift 14: 273–330.
  • Tabell J, Baldizzone G (2014) Coleophora mareki Tabell & Baldizzone, sp. n., a new coleophorid moth of the serpylletorum species-group (Lepidoptera: Coleophoridae). SHILAP Revista de Lepidopterología 42(167): 399–408.
  • Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011) MEGA5: Molecular Evolutionary Genetics Analysis using Maximum Likelihood, Evolutionary Distance, and Maximum Parsimony Methods. Molecular Biology and Evolution 28: 2731–2739. doi: 10.1093/molbev/msr121
  • The Angiosperm Phylogeny Group (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: 105–121. doi: 10.1111/j.1095-8339.2009.00996.x
  • Turner H, Lieshout N, Van Ginkel WE, Menken SBJ (2010) Molecular Phylogeny of the Small Ermine Moth Genus Yponomeuta (Lepidoptera, Yponomeutidae) in the Palaearctic. PLoS ONE 5(3): e9933. doi: 10.1371/journal.pone.0009933
  • Zeller PC (1868) Beitrag zur Kenntniss der Lepidoptern-Fauna der Umgegend von Raibl in Oberkärnthen und Preth im angrenzenden Küstengebiet. Verhandlungen der kaiserlich-königlichen zoologisch-botanischen Gesellschaft in Wien 18: 563–628.