Integrative revision of the Caryocolum schleichi species group – a striking example of a temporally changing species concept (Lepidoptera, Gelechiidae)

The taxonomy of the Palaearctic Caryocolum schleichi species group is revised, leading to a largely new species concept. Initially described as four different species (C. schleichi, C. arenariella, C. dianthella, C. improvisella), these taxa were later considered as subspecies. Recently the taxon C. arenariella was re-instated to species level without, however, revision of the remaining three subspecies. Analysis of DNA barcodes of the mtCOI (Cytochrome c Oxidase 1) gene of all four validly described taxa showed an unexpected pattern of genetic diversity. Careful re-examination of morphological traits, particularly male and female genitalia, fully supported this pattern, leading to the re-instatement of C. dianthella sp. rev. and C. improvisella sp. rev. as valid species and the description of three new species, all of them occurring in the Alps: C. messneri sp. nov. (Italy, Slovenia, Bulgaria, Greece), C. lamai sp. nov. (Italy, France) and C. habeleri sp. nov. (France, Switzerland, Germany). All species are described in detail and the adults and genitalia of both sexes are illustrated.


Introduction
With about 865 species the Gelechiidae are among the most diverse families of Lepidoptera in Europe, only exceeded by the Tortricidae, Geometridae and Noctuidae. Additionally, many undescribed species probably exist (Huemer and Karsholt 2020) and it seems not unlikely that the species number will rise to 1000. In contrast to several unrevised groups of Gelechiidae, the genus Caryocolum has received above-average attention with extensive revisionary work on a continental or intercontinental scale (Klimesch 1953(Klimesch -1954Huemer 1988;Huemer and Karsholt 2010). Collecting efforts in remote areas and the introduction of molecular traits has further contributed to an increase of previously overlooked diversity and the number of described species has reached 57 in Europe (Bella 2008;Grange and Nel 2012;Huemer and Nel 2005;Huemer and Karsholt 2010;Huemer et al. 2014;Nel and Requena 2017;Huemer and Karsholt 2020). However, Huemer et al. (2014) from extensive DNA barcoding of a large portion of the European fauna have already indicated several species groups with potentially cryptic species. One of these groups is that of Caryocolum schleichi s.l., revised in this paper. Following the taxonomic revision of Caryocolum (Huemer 1988) this species was considered as a morphologically variable taxon with several geographically separated subspecies in Europe and parts of Asia: C. schleichi schleichi (Christoph, 1872), C. schleichi dianthella (Chrétien, 1925), C. schleichi improvisella (Rebel, 1936) and C. schleichi arenariella (Benander, 1937), all of them initially described as species. However, Aarvik et al. (2017), without in-depth discussion, re-instated C. arenariella as a separate species, leaving the remaining taxa untouched. Extensive studies of barcode sequences of the mitochondrial COI gene (Huemer et al. 2014) had already indicated species level for this species and the remaining subspecies and the possibility of further cryptic species. These shortcomings are here formally resolved with two subspecies reinstated to species level and the unexpected discovery of three hitherto unnamed species.

Material and methods
Generic descriptions of Caryocolum have been published by Huemer (1988) and Huemer and Karsholt (2010) and are thus not repeated here.

Specimens
The study is based on about 200 specimens of the Caryocolum schleichi-group. Material was either traditionally set and dried or, particularly more recently, mainly spread or only pinned. Genitalia preparations usually followed standard techniques (Robinson 1976) adapted for the male genitalia of Gelechiidae and female genitalia of Caryocolum by the "unrolling technique" as described by Pitkin (1986) for males and Huemer (1987) for females.

DNA Barcodes
DNA barcode sequences are based on a 658 base-pair long segment of the mitochondrial COI gene (cytochrome c oxidase 1). DNA samples (dried legs) were prepared according to the prescribed standards. Legs from 33 specimens of the Caryocolum schleichi species group were successfully processed at the Canadian Centre for DNA Barcoding (CCDB, Biodiversity Institute of Ontario, University of Guelph) to obtain DNA barcodes using the standard high-throughput protocol described in deWaard et al. (2008), supplemented by a further 12 public sequences from BOLD. Sequences were submitted to GenBank, and further details including complete voucher data and images can be accessed in the public dataset "Lepidoptera of Europe -Caryocolum schleichi species group [DS-CARYSCHL]" in the Barcode of Life Data Systems (BOLD; Ratnasingham and Hebert 2007). Degrees of intra-and interspecific variation of DNA barcode fragments were calculated under the Kimura 2 parameter model of nucleotide substitution using analytical tools of BOLD systems v. 4.0. (http://www.boldsystems. org). A Neighbor-Joining tree of DNA barcode data was constructed using MEGA 6 (Tamura et al. 2013) under the Kimura 2 parameter model for nucleotide substitutions. A three-letter code (ISO 3166-1 alpha-3, https:// en.wikipedia.org/wiki/ISO_3166-1_alpha-3) was used to abbreviate country names.

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

Molecular analysis
Successfully sequenced specimens originate mainly from Europe, with a few exceptions of Asian vouchers (Fig. 1). DNA sequencing resulted in a full barcode fragment for 38 specimens, 6 sequences > 600 bp and a single shorter sequence of 568 bp which were considered for analysis. Seven distinct DNA barcode clusters were observed, supporting the already known four taxa (irrespective of subspecies or species level) and another three hitherto unrecorded putative taxa. Subsequent morphological examination of these clusters resulted in a fully concordant pattern of seven morphospecies (see below).
Sequences of the COI barcode region of these morphospecies reveal moderately low intraspecific but significant interspecific genetic distances. Mean distances within species are 0.60% with a minimum of 0% and maximum of 1.71%. Mean interspecific distances are much higher with 4.71%, ranging from minimum 2.35% to maximum 6.47%. Barcode gaps range from 2.35% to 3.43% to the nearest neighbor (Fig. 2

Caryocolum schleichi species group
The Caryocolum schleichi species group belongs to the leucomelanella-group as defined by Huemer (1988) and which is characterized in the male genitalia by the strongly sclerotized transtilla, without spines, the broad valva with one or two processes, two pairs of digitate processes at the posterior margin of the vinculum and a straight phallus with minute apical cornuti. In the fe-male genitalia the group is characterized by the lack of specialized structures of segment VIII, the folds delimiting antrum and joined in medial part of segment VIII, the short antrum and the long lateral sclerotizations in the posterior part of the ductus bursae. The Caryocolum schleichi species group itself is characterized by the distally broad valva with two processes, a unique structure in Caryocolum.
Larvae of the Caryocolum schleichi species group are restricted to the genus Dianthus L. (Caryophyllaceae) as far as known. Dorso-apical process of valva longer than ventro-apical process, slender, valva with nearly straight outer edge sub-apically; posterior margin of vinculu with pair of short digitate processes (Figs 17,19,25,27 1 ♀, Prov.Konya, 20 km SE Hadim, 1800m, 14 Jul. 1986. Diagnosis. C. schleichi can be easily separated from other species of the group by the white head, thorax and tegulae (but see below for material from Turkey). The male genitalia are particularly characterized by the massive valva, with a short and broadly pointed dorso-apical process. In this character the species is most similar to C. habeleri with a more slender, spine-like dorso-apical process of the valva and to C. dianthella with a smaller and dorsally concave valva with shorter and more slender dorso-apical and ventro-apical processes. Further, the sacculus of C. schleichi is shorter and stouter than in C. dianthella and C. habeleri. In all other species of the C. schleichi species group, the valva is more slender with a much longer dorso-apical process and a more slender dorso-ventral process. The female genitalia are almost indistinguishable from other species of the C. schleichi species group, except for the apophysis anterior which distinctly exceeds the length of segment VIII by about one-quarter, whereas it is of about the same length in all other species except for C. dianthella from which it differs by the narrower signum-hook.
Description. Adult (Fig. 3). Forewing length ♂ 4.5-6.5 mm, ♀ 4.5-5.0 mm. Head cream-white; labial palpus predominantly cream-white, dark brown mottling on outer surface of segment one, light brown mottling on outer surface of segment two and dark brown mixed with cream on segment three; antenna black, weakly ringed paler. Thorax and tegula white, occasionally mottled with light brown, few dark brown scales anteriorly. Abdomen cream-white on ventral surface. Forewing dark brown, mottled with greyish white on dorsal margin; cream-white transverse fascia from fold to costa at one-fifth; cream-white medial area frequently extended towards costa and dorsal margin; cream-white costal and tornal spots usually separate; fringes basally dark brown, distal part lighter. Hindwing light grey.
Variation. Specimens from Turkey frequently exhibit a darker head and thorax (Huemer and Karsholt 2010) and the taxonomic status of such specimens should be re-evaluated in future from more extensive material and molecular data.
Female genitalia (Figs 29,36,43). Apophysis posterior about three times length of apophysis anterior; segment VIII without processes, smooth; ostium bursae with short lateral folds; apophysis anterior 1.2 times length of segment VIII; antrum short, about one-fifth length of apophysis anterior, funnel-shaped; posterior part of ductus bursae with pair of long lateral sclerites extending to about apex of apophysis anterior, membranous part of ductus bursae about length of segment VIII including apophysis anterior; signum on right side of entrance to sub-oval corpus bursae, with broad base and moderately short, strongly bent hook. Molecular data. BIN: BOLD:ADH6455. The intraspecific average distance of the barcode region is 0.8%, the maximum distance 0.8% (p-dist) (n = 2). The minimum distance to the nearest neighbor, C. arenariella, is 3.37%.
Distribution (Fig. 1). Confirmed records are known from south-western Russia to Turkey and parts of Syria. A record from Macedonia (Klimesch 1968) probably refers to C. arenariella. The taxonomy of some populations from the Near and Middle East requires further revisionary work. Huemer (1988) had already suspected an additional subspecies from northern Syria and Afghanistan but due to the lack of material and in absence of molecular data this problem has to be re-assessed in the future.
Bionomics. Host-plant and early stages are unknown, but it is suspected that the larva feeds on Dianthus similar to other species of the C. schleichi species group. C. schleichi is most probably restricted to warm and sunny habitats at low elevations to about 1800 m, but precisely documented observations to the habitat are missing. Moths have been collected from late May to October.
Remarks. Lita schleichi was described from an unspecified number of specimens from south-western Russia (Christoph 1872) with the lectotype selected by Huemer (1988). Caryocolum syriacum was described from Syria and the holotype fixed in the original description (Povolný 1977). (Chrétien, 1925)  Diagnosis. C. dianthella differs from C. schleichi by the brown rather than white head, thorax and tegulae but cannot be reliably separated from the other species of the complex by external characters. However, the male genitalia are characterized by the shape of the valva, particularly the convex dorsal margin with a sub-apical hump, the shortly pointed sub-dorsal process and the small digitate ventral process. A similarly short but broader dorso-apical process is only found in C. schleichi, whereas C. habeleri with a similar dorso-apical process differs in the large ventro-apical projection and the weakly curved dorsal margin of the valva. In all other species of the C. schleichi species group, the valva is more slender with a much longer dorso-apical process and a more slender dorso-ventral process. The female genitalia are almolst indistinguishable from other species of the C. schleichi species group except for the characteristic signum with a broad hook and a largely reduced base.

Caryocolum dianthella
Description. Adult (Fig. 4). Forewing length ♂ 4.0-5.5 mm, ♀ 4.0-5.0 mm. Head dark brown, frons cream-white to white; labial palpus dark brown, second segment cream-white on inner and upper surface, mottled with grey-brown on outer surface, third segment dark brown mottled with few white scales; antenna black, weakly ringed paler. Thorax and tegula rusty brown, anterior part dark brown. Abdomen predominantly grey on ventral surface, medially cream-coloured. Forewing dark brown with weak light mottling, distinct white markings: transverse fascia from fold to costa at one-fifth, distinct white medial spot, separate costal and tornal spots; fringes basally dark brown, distal part lighter. Hindwing light grey.
Variation. Weak variation in the extent of the white markings.
Female genitalia (Figs 30,37,44). Apophysis posterior about four times length of apophysis anterior; segment VIII without processes, smooth; ostium bursae with short lateral folds; apophysis anterior 1.3 times length of segment VIII; antrum short, about one-fifth length of apophysis anterior, funnel-shaped; posterior part of ductus bursae with pair of long lateral sclerites extending to about apex of apophysis anterior, membranous part of ductus bursae about length of segment VIII including apophysis anterior; signum on right side of entrance to sub-oval corpus bursae, with broad base lacking crescent-shaped lateral projections, and stout hook. Molecular data. BIN: BOLD:AAU1854. The intraspecific average distance of the barcode region is 1.43%, the maximum distance 1.83% (p-dist) (n = 3). The minimum distance to the nearest neighbor, C. lamai, is 3.05%. Distribution (Fig. 1). C. dianthella is restricted to the Western Mediterranean, with country records from Morocco, Portugal, Spain and France (Pyrenees).
Bionomics. The larva has been recorded in May and June feeding on Dianthus scaber toletanus (Boiss and Reuter) Tutin and D. deltoides L. and moths emerged in July (Chrétien 1925). In the Pyrenees it was also bred from Dianthus pungens L. (see material examined). In the field moths have been collected from late June to October up to altitudes of about 2600 m in Morocco.
Remarks. Lita dianthella was described from an unspecified number of specimens from central Spain (Chré-tien 1925) and the genitalia of a syntype figured by Agenjo (1962) leave no doubt as to the identity. Caryocolum hackeri was described from northern Spain (Pyrenees) with the holotype fixed in the original description (Derra 1985).
Caryocolum improvisella (Rebel, 1936)  Diagnosis. C. improvisella differs from C. schleichi by the brown rather than white head, thorax and tegulae. It is usually larger than C. arenariella and C. lamai but otherwise cannot be reliably separated from the other species of the complex by external characters. The male genitalia are characterized in particular by the distally widened valva with a moderately long, pointed dorsal process, a character which is only shared with C. arenariella, C. messneri, and C. lamai. However, this process is distinctly shorter than in C. arenariella and C. lamai. The species differs from C. messneri in several characters such as the broader valva with a longer dorsal process and the distinctly shorter latero-medial processes of the vinculum. The female genitalia are hardly discernible from several other species of the C. schleichi species group though the short crescent shaped base of the signum in combination with the moderately slender signum hook seem characteristic and differ from all other species except for C. messneri. However, the individual variation of these characters is insufficiently documented due to lack of material.
Description. Adult (Figs 5, 6). Forewing length ♂ 5.5-7.0 mm, ♀ 4.5-6.0 mm. Head with dark grey-brown vertex, mottled with white, frons cream-white; first seg-Figures 9-12. Adults. 9 Caryocolum lamai, paratype, male, Italy; 10 C. lamai, paratype, male, Italy; 11 C. messneri, paratype, male, Italy; 12 C. habeleri, paratype, male, France. ment of labial palpus dark brown, second segment creamwhite on inner and upper surface, ventral and outer surface predominantly dark brown, dark brown mixed with few whitish scales on segment three; antenna black, weakly ringed paler. Thorax and tegula dark brown anteriorly, posterior half mixed cream with dark brown and some rusty brown mottling. Abdomen grey to whitish grey on ventral surface. Forewing dark brown, mottled with greyish white on dorsal margin; white transverse fascia from fold to costa at one-fifth; white medial spot occasionally extended towards costal and dorsal margin; white costal and tornal spots separate; fringes basally dark brown, distal part paler. Hindwing light grey.
Variation. The extension of white markings of the forewings varies considerably.
Male genitalia (Figs 15,16,23,24). Uncus broadly sub-quadrangular, posterior corners rounded; lateral sclerites of gnathos distinct, medial part with large minutely spined culcitula; tegumen weakly widened anteriorly, with slightly emarginated anterior margin; transtilla sclerotized, longitudinally folded; pedunculus large, suboval, with sclerotized inner edge; valva nearly straight, long, distal part broadly dilated with weakly curved dorsal and straight ventral edges, apex with two processes, moderately long and pointed dorsal process extended to about middle of uncus, broadly digitate ventral process at about right-angles to and about length of dorsal process; sacculus moderately long, slender, with parallel margins, apically pointed; posterior margin of vinculum deeply incised medially, with pair of moderately long digitate processes, pair of latero-medial processes broadly humplike; saccus slightly longer than valva, slender, gradually tapered to apex; phallus long, slender, nearly straight, apically with area of small cornuti.
Female genitalia (Figs 31, 38, 45). Apophysis posterior about four times length of apophysis anterior; segment VIII without processes, smooth; ostium bursae with short lateral folds; apophysis anterior about length of segment VIII; antrum short, about one-fifth length of apophysis anterior, funnel-shaped; posterior part of ductus bursae with pair of long lateral sclerites extending to about apex of apophysis anterior, membranous part of ductus bursae about length of segment VIII including apophysis anterior; signum on right side of entrance to sub-oval corpus bursae, with broad and shortly crescent-shaped base and moderately slender hook. Molecular data. BIN: BOLD:ACL3018. The intraspecific average distance of the barcode region is  0.31%, the maximum distance 0.46% (p-dist) (n = 3). The minimum distance to the nearest neighbor, C. messneri, is 3.05%.
Distribution (Fig. 1). The species is currently only known from a small section of the Central Alps of Switzerland, Italy and Austria.
Bionomics. The larva has been recorded feeding within shoots of Dianthus sylvestris Wulfen (Klimesch 1953;Burmann 1990; pers. obs.) and moths emerged from early July to the beginning of August. The species has been recorded along steep rock-surface on siliceous soil in the montane zone.  Remarks. Lita improvisella was described from 1 ♂ and 1 ♀ collected at two different localities in Switzerland (Graubünden) by Thomann, and a further in-correctly identified specimen probably of C. leucomelanella from Valais, with the genitalia figured in the original description (Rebel 1936 figured the male genitalia of a syntype from Remüs, which is here designated as the lectotype in order to fix the identity. The second specimen from Landquart, 1.vii.1922, leg. Thomann, probably belongs to C. habeleri, a species which has been collected nearby in the Swiss Rhine valley. (Benander, 1937)  Diagnosis. C. arenariella is, besides C. lamai, the smallest and darkest species of the C. schleichi species group, with the white markings and brown scales reduced in many specimens. It differs from C. schleichi by the brown rather than cream-white head, thorax and tegulae but cannot be reliably separated from the other species of the complex by external characters. However, the male genitalia are characterized by the shape of the valva, particularly the long, pointed dorsal process, only shared with C. improvisella, C. messneri, and C. lamai. This process is more slender and longer than in C. improvisella and C. messneri whereas the male genitalia differ from C. lamai by the broader sacculus and the more slender and longer digitate medial processes of the vinculum. The female genitalia are hardly discernible from other species of the C. schleichi species group, except for the signum with a slender and long crescent-shaped base, a character only shared with C. lamai, and the particularly short apophysis posterior. However, the individual variation of this character is insufficiently documented due to lack of material.
Variation. Specimens from Macedonia are larger (6.0 mm) and together with parts of Russian material exhibit some light mottling on the thorax and extended creamwhite mottling of the forewing, particularly the dorsal area.
Male genitalia (Figs 17, 25). Uncus broadly sub-quadrangular, posterior corners rounded; lateral sclerites of gnathos distinct, medial part with large minutely spined culcitula; tegumen weakly widened anteriorly, with slightly emarginated anterior margin; transtilla sclerotized, longitudinally folded; pedunculus large, sub-triangular, with sclerotized inner edge; valva nearly straight, long, distal part moderately slender, weakly dilated with straight dorsal and ventral edges, apex with two processes, long and pointed dorsal process extended to posterior third of uncus, broadly digitate ventral process at about right-angles to dorsal process; sacculus moderately long, slender, with parallel margins, apically pointed; posterior margin of vinculum deeply incised medially, with pair of moderately short digitate processes, pair of latero-medial processes broadly projected; saccus slightly longer than valva, slender, gradually tapered to apex; phallus long, slender, nearly straight, apically with area of small cornuti.
Female genitalia (Figs 32, 39, 46). Apophysis posterior about 3.5 times length of apophysis anterior; segment VIII without processes, smooth; ostium bursae with short lateral folds; apophysis anterior about length of segment VIII; antrum short, about one-fifth length of apophysis anterior, funnel-shaped; posterior part of ductus bursae with pair of long lateral sclerites extending to about apex of apophysis anterior, membranous part of ductus bursae about length of segment VIII including apophysis anterior; signum on right side of entrance to sub-oval corpus bursae, with slender and long crescent-shaped base and long slender hook. Molecular data. BIN: BOLD:AAE9479. The intraspecific average distance of the barcode region is 0.6%, the maximum distance 1.31% (p-dist) (n = 10). The minimum distance to the nearest neighbor, C. messneri, is 2.25%. Distribution (Fig. 1). Only known from scattered localities in Sweden, Finland, the Baltic countries (Aarvik et al. 2017), North Macedonia, and from southern Siberia.
Bionomics. The larva has been recorded in May, feeding in the stem of Dianthus arenarius L. which becomes gall-like stout and swollen (Benander 1926). Pupation takes place outside the gall on the ground surface. However, according to Mutanen (in litt.) larvae live within the stems, without causing a conspicuous swollen gall. In the field moths have been collected from late June to early September at artificial light sources.  Remarks. Lita arenariella was described from an unspecified number of specimens and no syntypes have been traced. However, the detailed description of the moth, genitalia and biology (Benander 1926) leave no doubt about the identity of this species. The taxon was re-instated at species level by Aarvik et al. (2017).  1 ♀, Epirus, Mitsikeli Mt. near Joannina, 1400m, 39°46'15"N, 20°48'20"E, 8 Jul. 2005.
Diagnosis. C. messneri differs from C. schleichi by the brown rather than cream-white head, thorax and tegulae. It is usually larger than C. arenariella and C. lamai and the medial spot is less sharply delimited than in C. dianthella. Otherwise, C. messneri cannot be reliably separated from the other species of the complex by external characters, though the usually well developed rusty-brown mottling seems unique. The male genitalia are characterized in particular by the moderately long, pointed dorsal process of the valva, a character which is only shared with C. improvisella, C. arenariella, and C. lamai. However, this process is distinctly shorter than in C. arenariella and C. lamai. The species further differs from C. improvisella in several characters such as the more slender valva with parallel outer edges and a distodorsal bulge, a shorter dorsal process, and the distinctly longer latero-medial processes of the vinculum. The female genitalia are hardly discernible from several other species of the C. schleichi species group though the shortly crescent-shaped base of the signum in combination with the moderately slender signum hook seem characteristic and differ from all other species except for C. improvisella. However, the individual variation of these characters is insufficiently documented due to lack of material.
Description. Adult (Fig. 11). Forewing length ♂ 5.5-6.0 mm, ♀ 5.0 mm. Head with dark grey-brown vertex, mottled with white, frons cream-white; first segment of labial palpus dark brown, second segment cream-white on inner and upper surface, ventral and outer surface predominantly dark brown, dark brown mixed with few whitish scales on segment three; antenna black, weakly ringed paler. Thorax and tegula dark brown anteriorly, posterior half cream with rusty brown mottling. Abdomen cream-white with some grey on ventral surface. Forewing dark brown, mottled with white on dorsal margin; basal two thirds with rusty-brown mottling, particularly white markings; white transverse fascia from fold to costa at one-fifth; white medial spot occasionally extended towards costa and dorsal margin; white costal and tornal spots separate; fringes basally dark brown, distal part paler. Hindwing light grey.
Variation. In the limited available material, the extent of rusty-brown mottling shows some variation.
Male genitalia (Figs 18, 26). Uncus broadly sub-quadrangular, posterior corners rounded; lateral sclerites of gnathos distinct, medial part with large minutely spined culcitula; tegumen weakly widened anteriorly, with slightly emarginated anterior margin; transtilla sclerotized, longitudinally folded; pedunculus large, sub-triangular, with sclerotized inner edge; valva nearly straight, long, moderately slender distal part with parallel edges, except for bulged disto-dorsal edge, apex with two processes, stout and moderately long and pointed dorsal process extended to about middle of uncus, broadly digitate ventral process at about right-angles to and slightly longer than dorsal process; sacculus moderately long, slender, with parallel margins, apically pointed; posterior margin of vinculum deeply incised medially, with pair of moderately long digitate processes, pair of latero-medial processes moderately broad, digitate, distinctly projected; saccus slightly longer than valva, slender, gradually tapered to apex; phallus long, slender, nearly straight, apically with area of small cornuti.
Female genitalia (Figs 33, 40, 47). Apophysis posterior almost five times length of apophysis anterior; segment VIII without processes, smooth; ostium bursae with short lateral folds; apophysis anterior about length of segment VIII; antrum short, about one-fifth length of apophysis anterior, funnel-shaped; posterior part of ductus bursae with pair of long lateral sclerites extending to about apex of apophysis anterior, membranous part of ductus bursae about length of segment VIII including apophysis anterior; signum on right side of entrance to sub-oval corpus bursae, with broad and shortly crescent-shaped base and moderately slender hook. Molecular data. BIN: BOLD:ACT3307. The intraspecific average distance of the barcode region is 1.18%, the maximum distance 1.77% (p-dist) (n = 4). The minimum distance to the nearest neighbor, C. arenariella, is 2.25%.
Etymology. The species is named in honour of Reinhold Messner, the first climber to ascend all fourteen peaks above 8000 metres sea level and living in the neighborhood of the type locality of the new species.
Distribution (Fig. 1). The species is currently only known from a restricted area in northern Italy (region of Trentino-Alto Adige) and from specimens collected in Slovenia, Bulgaria and Greece.
Bionomics. Host-plant and early stages are undescribed but it seems most likely that the species shows a similar behaviour as related taxa with a hostplant restriction to Dianthus spp. At the type-locality Dianthus sylvestris Wulfen has been recorded. The adults have been found from mid to late August nearby rocks on siliceous soil where they were attracted to artificial light sources.
Diagnosis. C. lamai is, together with C. arenariella, the smallest and darkest species of the C. schleichi species group with reduced white markings and brown scales. It differs from C. schleichi by the brown rather than white head, thorax and tegulae but cannot be reliably separated from the other species of the complex by external characters of the forewings. However, the male genitalia are characterized by the shape of the valva, particularly the long, sharply pointed dorsal process, only shared with C. improvisella, C. messneri and C. arenariella. This process is more slender and longer than in C. improvisella and C. messneri, whereas the male genitalia differ from C. arenariella in particular in the more slender sacculus and the broader and the digitate medial processes of the vinculum. The female genitalia are hardly discernible from other species of the C. schleichi species group, except for the signum with a slender and long crescent-shaped base, a character only shared with C. arenariella, from which it differs by the longer apophysis posterior. However, the individual variation of this character is insufficiently documented due to lack of material.
Description. Adult (Figs 9, 10). Forewing length ♂ 4.5-5.5 mm, ♀ 4.0 mm. Head dark grey-brown, mottled with light grey, frons greyish white; labial palpus dark brown, second segment cream-white on inner surface, third segment blackish, mottled with white; antenna black, weakly ringed paler. Thorax and tegula dark greybrown, with few rusty brown scales. Abdomen dark grey on ventral surface. Forewing dark brown with some light mottling, particularly on dorsum; distinct white markings: narrow transverse fascia from fold to costa at onefifth, irregular white medial spot, separate costal and tornal spots; fringes basally dark brown, distal part lighter. Hindwing light grey.
Variation. The extension of light mottling of the forewing varies slightly.
Male genitalia (Figs 19,27). Uncus broadly sub-quadrangular, posterior corners rounded; lateral sclerites of gnathos distinct, medial part with large minutely spined culcitula; tegumen weakly widened anteriorly, with slightly emarginated anterior margin; transtilla sclerotized, longitudinally folded; pedunculus large, sub-triangular, with sclerotized inner edge; valva nearly straight, long, distal part moderately slender, weakly dilated with straight dorsal and ventral edges, apex with two processes, particularly long and pointed dorsal process extended to posterior third of uncus, slender digitate ventral process at about right-angles to dorsal process; sacculus long, slender, with parallel margins, apically pointed; posterior margin of vinculum deeply incised medially, with pair of short digitate processes, pair of latero-medial processes broadly projected; saccus slightly longer than valva, slender, gradually tapered to apex; phallus long, slender, nearly straight, apically with area of small cornuti.
Female genitalia (Figs 34,41,48). Apophysis posterior about four times length of apophysis anterior; segment VIII without processes, smooth; ostium bursae with short lateral folds; apophysis anterior about length of segment VIII; antrum short, about one-fifth length of apophysis anterior, funnel-shaped; posterior part of ductus bursae with pair of long lateral sclerites extending to about apex of apophysis anterior, membranous part of ductus bursae about length of segment VIII including apophysis anterior; signum on right side of entrance to sub-oval corpus bursae, with long and slender crescent-shaped base and long slender hook. Molecular data. BIN: BOLD:AAE9478. The intraspecific average distance of the barcode region is 0.04%, the maximum distance 0.15% (p-dist) (n = 7). The minimum distance to the nearest neighbor, C. dianthella, is 3.05%.
Etymology. The species is named in honour of David Lama (1990-2019, one of the most famous Austrian alpinists, who was tragically killed by an avalanche in Banff National Park (Canada) on the 1 st of April 2019. David supported earlier work on landscape conservation in Tyrol with enthusiasm.
Distribution. The species is currently only known from a restricted area in northern Italy (region of Piedmont) and France (Alpes-Maritimes, Alpes-de-Haute-Provence).
Bionomics. Host plant and early stages are unknown but it seems most likely that the species shows a similar behaviour as related taxa with a hostplant restriction to Dianthus spp. The adults have been found from late July until late August near rock and scree at altitudes of about 1800 to 2600 m on calcareous soil (Fig. 50), where they were attracted to artificial light sources.  Diagnosis. C. habeleri differs from C. schleichi by the brown rather than white head, thorax and tegulae, it is generally larger than C. arenariella and C. lamai but otherwise cannot be reliably separated from the other species of the complex from external characters. However, the male genitalia are characterized by the shape of the valva, particularly the short pointed sub-dorsal process and the broad ventral process. A similarly short but broader dorso-apical process is only found in C. schleichi which furthermore has a broader shovel-shaped valva, and in C. dianthella with a shorter and dorsally distinctly bulged valva. In all other species of the C. schleichi species group, the valva is more slender with a much longer dorso-apical process and smaller dorso-ventral process. The female genitalia are hardly discernible from those of the other species of the C. schleichi species group though the moderately long antrum, the distinctly crescent-shaped base of the signum  in combination with the moderately slender signum hook seem characteristic. However, the individual variation of these characters is insufficiently documented due to lack of material.
Description. Adult (Fig. 12). Forewing length ♂ 4.5-5.5 mm, ♀ 4.0-5.0 mm. Head with dark grey-brown vertex, mottled with white, frons cream-white; first segment of labial palpus dark brown, second segment cream-white on inner and upper surface, ventral and outer surface predominantly dark brown, dark brown mixed with few whitish scales on segment three; antenna black, weakly ringed paler. Thorax and tegula dark brown anteriorly, posterior half mixed cream with dark brown and some rusty brown mottling. Abdomen greyish cream on ventral surface. Forewing dark brown, basal half with some rusty-brown mottling, dorsal margin mottled with greyish; white transverse fascia from fold to costa at onefifth; white medial spot occasionally extended towards costa and dorsal margin; white costal and tornal spots separate; fringes basally dark brown, distal part paler. Hindwing light grey.
Variation. The extension of white marking on the forewing varies considerably and particularly worn specimens look paler.
Female genitalia (Figs 35,42,49). Apophysis posterior about 4.5 times length of apophysis anterior; segment VIII without processes, smooth; ostium bursae with short lateral folds; apophysis anterior about length of segment VIII; antrum short, about one-quarter length of apophysis anterior, funnel-shaped; posterior part of ductus bursae with pair of long lateral sclerites extending to about apex of apophysis anterior, membranous part of ductus bursae about length of segment VIII including apophysis anterior; signum on right side of entrance to sub-oval corpus bursae, with moderately broad crescent-shaped base and moderately slender hook. Molecular data. BIN: BOLD:ACB5067. The intraspecific average distance of the barcode region is 0.39%, the maximum distance 0.98% (p-dist) (n = 12). The minimum distance to the nearest neighbor, C. messneri, is 3.29%.
Etymology. The species is named in honour of Peter Habeler who, along with Reinhold Messner, completed the first solo ascent of Mount Everest without supplemental oxygen. Along with the author, Peter is currently involved in the "Blühendes Österreich" conservation foundation. Incidentally, his cousin Heinz Habeler  acquired one of the largest collection of Lepidoptera from the south-eastern Alps, stored at TLMF.
Distribution. The species is currently only known from widely separated localities, ranging from the French Pyrenees to south-eastern Bavaria.
Bionomics. Host-plant and early stages are undescribed but it seems most likely that the species shows a similar behaviour as related taxa with a hostplant restriction to Dianthus spp. The adults have been found from late July to late September at artificial light sources near rock and scree on calcareous soil ranging from lower altitudes of the submontane zone up to about 1000 m in the Alps.

Discussion
The Caryocolum schleichi species group is an impressive example of the frequently subjective taxonomic assessment of problematic taxa with an allopatric distribution pattern (Mutanen et al. 2012). The different classification of the involved taxa at the species or subspecies level has so far been based solely on morphological features, but the differentiation with molecular methods (Hebert et al. 2003) used here for the first time is a considerable advance. Mutanen et al. (2012) call for at least two mutually independent, diagnostically relevant feature complexes for allopatric taxa in order to arrive at a more objective species delimitation. Divergences in the DNA barcode between the individual taxa of the Caryocolum schleichi species group, which were described a considerable time ago, are such an important feature, and independent of morphological discrimination, that supports the species status of the taxa involved. Huemer et al. (2014) from first molecular data had already presumed that named taxa in the Caryocolum schleichi species groupshould be raised to species level, however, without formal implementation of this assumption.
In addition, DNA barcode distances are also an important tool for recognizing possible cryptic diversity. The basic taxonomic structure of the Caryocolum schleichi species group -apart from the species or subspecies congestion -was undisputed after the revision of the genus by Huemer (1988). As can now be seen, genetic sequences particularly of alpine populations show a much more complex structure. Instead of the previously postulated species or subspecies improvisella, four species are now recognized in the Alps, some with limited distribution (Fig. 50), some with a larger area outside Central Europe. In the area of the Central Alps between Austria, Switzerland and Italy, three species collide spatially and even an occasional sympatric occurrence seems possible. All these taxa could only be morphologically separated after the discovery of genetic distances. Former reports of the taxon improvisella have thus become obsolete and need to be checked thoroughly. The present study shows once again that a broad screening of samples using DNA barcoding can lead to surprises even in Central Europe. Blindly following earlier work must be strongly warned against, even in the case of classic, purely morphologically based revisions.