Research Article |
Corresponding author: Peter Huemer ( p.huemer@tiroler-landesmuseen.at ) Academic editor: Bernard Landry
© 2021 Jürg Schmid, Peter Huemer.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Schmid J, Huemer P (2021) Unraveling a complex problem: Dichrorampha velata sp. nov., a new species from the Alps hitherto confounded with D. alpestrana ([Zeller], 1843) sp. rev. = D. montanana (Duponchel, 1843) syn. nov. (Lepidoptera, Tortricidae). Alpine Entomology 5: 37-54. https://doi.org/10.3897/alpento.5.67498
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Analysis of wing pattern, genital morphology and results of DNA barcoding indicates that the name Dichrorampha montanana sensu auct. actually comprises two species. D. alpestrana ([Zeller], 1843) sp. rev. is considered as senior synonym of D. montanana (Duponchel, 1843) syn. nov., and a lectotype is designated for the latter name to fix the identity. After thorough search for possible synonyms, Dichrorampha velata sp. nov. is described and differentiated morphologically and with DNA barcodes from D. alpestrana and adults and genitalia of both species are figured. Dichrorampha velata sp. nov. is restricted to the European Alps and adjacent regions whereas D. alpestrana is more widespread with likely arctic-alpine disjunction and records from the Alps and the northern part of Great Britain.
cryptic diversity, DNA barcoding, integrative taxonomy, nomenclature, revised synonymy
With currently 62 species records Dichrorampha Guenée, 1845 is the most diverse genus of European Tortricidae (
In 2006, the first author collected two superficially similar fresh male Dichrorampha specimens occurring syntopically at a site in the Engadine, Switzerland. They differed in their wing pattern in that one specimen displayed a distinct whitish triangular mark arising from the dorsal border of the forewing, while in the other, this mark was hardly apparent (Figs
Already more than a decade earlier the junior author found similar discrepancies in the context of material from Austria and Germany. However, there was no intensive processing of the problem at the time because these genital differences had already been described in detail and figured by
The history of this taxon, however, is characterized by confusion and misunderstandings:
Sometime before 1843, the Austrian entomologist Josef Emanuel Fischer von Röslerstamm captured an unknown tortricid at Gscheidt in the Rax mountain range, in the Austrian province of Styria. He sent it under the name “Montanana n.sp.” to the famous German entomologist Philipp Christoph Zeller who described the new species as “Grapholita alpestrana” ([
Herrich-Schäffer (1849) provided another description based on Zeller’s diagnosis, whereupon two mistakes occurred: he identified his illustration 193 as “montanana”, and in the text, while using Zeller’s name “alpestrana“, he attributed it erroneously to Fischer von Röslerstamm [=FR]. Probably in order to correct his first mistake, he added to the description the explanatory “montanana FR. olim” (Herrich-Schäffer [1849]). Finally, the taxon became known as Grapholita alpestrana HS.
Thus, in short: “montanana FR” in litt. became alpestrana [anonymous, but in fact Zeller], in error, then alpestrana FR, montanana HS and finally alpestrana HS. According to the International Code of Zoological Nomenclature’s Recommendation 51D, this taxon should bear the name Dichrorampha alpestrana ([Zeller], 1843), because the anonymous description can unambiguously be traced to Zeller (
Unfortunately almost simultaneously with [
As of today, Dichrorampha alpestrana ([Zeller], 1843) is considered an invalid junior synonym of Dichrorampha montanana (Duponchel, 1843), both in
We will subsequently prove that this view does not follow the regulations of
In the course of this study, a total of 104 male and 12 female specimens of “D. montanana sensu auct.” originating mainly from different parts of the Alps and the Jura mountains were investigated:
Specimen repositories:
The following specimen characteristics were analyzed:
The genital preparations were photographed with a Canon EOS 7D digital camera, using a Zeiss Primo Star microscope with a 4× and 10× plan-achromat lens.
Most of fresh specimens of both sexes could be grouped into two main categories: those with olive ground colour and irregular silvery lines on their forewings and those with more brownish ground colour and a more or less conspicuous bright triangular mark arising from dorsum and extending beyond mid-wing. A few specimens, however, presented a pattern/colour not easily assignable to either group.
The serial dissection of alpine “Dichrorampha montanana sensu auct.” specimens disclosed that the genital phenotype of all male specimens could unambiguously be assigned to two well defined groups with no intermediate forms:
DNA sequencing resulted in full barcode fragments for 35 specimens of "D. montanana s. auct" -complex. These grouped into two well delimited clusters, corresponding to the above mentioned groups I (18 specimens) and group II (17 specimens). Furthermore 248 full barcode sequences, 64 sequences > 500 bp and four shorter sequences < 500 bp from BOLD, covering 37 European species and two subspecies, were considered for analysis. Thirty-seven distinct DNA barcode clusters were observed, separating the vast majority of sequenced taxa, including the two suspected species in the D. montanana species group. However, DNA barcodes failed to separate closely related species of the D. plumbana species group (D. plumbana (Scopoli, 1763), D. sedatana Busck, 1906, D. aeratana (Pierce & Metcalfe, 1915), D. tarmanni Huemer, 2009), as well as D. infuscata (Danilevsky, 1960) and D. inconspiqua (Danilevsky, 1948).
Sequences of the COI barcode region in European Dichrorampha reveal moderately low intraspecific but significant interspecific genetic distances. Mean distances within species are 0.54% with a minimum of 0% and maximum of 4.54% in D. bugnionana (Duponchel, 1843) subspecies which likely represent distinct species. Mean distances to nearest neighbours are much higher with 3.45%, ranging from minimum 0% in the above mentioned species with barcode sharing to maximum 7.94% (Fig.
Neighbor-Joining tree of species in European Dichrorampha (Kimura 2 parameter, built with MEGA 6 cf.
Intraspecific mean K2P (Kimura 2 Parameter) divergences, maximum pairwise distances, nearest species, nearest neighbor and distance to nearest neighbor (%).
Species | Mean Intra-Sp % | Max Intra-Sp % | Nearest species | Nearest Neighbour | Distance to NN % |
---|---|---|---|---|---|
Dichrorampha acuminatana | 0.37 | 1.87 | Dichrorampha petiverella | BTLBP376-11 | 4.83 |
Dichrorampha aeratana | 1.32 | 5.08 | Dichrorampha sedatana | LON5763-17 | 0.46 |
Dichrorampha agilana | 0.39 | 0.64 | Dichrorampha teichiana | LEEUA626-11 | 4.18 |
Dichrorampha alpestrana | 0.72 | 2.5 | Dichrorampha velata | PHLAD201-11 | 3.96 |
Dichrorampha alpigenana | 1.54 | 3.1 | Dichrorampha inconspiqua | BTLBP451-11 | 1.4 |
Dichrorampha alpinana | 0.92 | 1.99 | Dichrorampha flavidorsana | FBLMT309-09 | 2.5 |
Dichrorampha bugnionana | 4.54 | 9.34 | Dichrorampha teichiana | LEEUA626-11 | 4.16 |
Dichrorampha cacaleana | 0.17 | 0.17 | Dichrorampha carpatalpina | LEASV661-19 | 7.94 |
Dichrorampha carpatalpina | N/A | 0 | Dichrorampha alpigenana | PHLAF568-11 | 5.65 |
Dichrorampha chavanneana | 0.78 | 1.87 | Dichrorampha consortana | FBLMZ645-12 | 4.92 |
Dichrorampha cinerascens | 1.23 | 2.44 | Dichrorampha acuminatana | LEAST411-17 | 6.4 |
Dichrorampha cinerosana | 0.86 | 1.24 | Dichrorampha plumbagana | CGUKC685-09 | 2.76 |
Dichrorampha consortana | 0.27 | 0.58 | Dichrorampha chavanneana | PHLAD710-11 | 4.92 |
Dichrorampha dinarica | 0.15 | 0.31 | Dichrorampha thomanni | LEASU278-18 | 4.47 |
Dichrorampha eximia | N/A | 0 | Dichrorampha petiverella | LASTS800-15 | 4.23 |
Dichrorampha flavidorsana | 0.58 | 1.44 | Dichrorampha teichiana | LEEUA626-11 | 2.05 |
Dichrorampha forsteri | N/A | 0 | Dichrorampha pentheriana | LEATJ1152-16 | 5.91 |
Dichrorampha gueneeana | 0.13 | 0.25 | Dichrorampha vancouverana | LPAB581-08 | 0.15 |
Dichrorampha heegerana | 0.23 | 0.46 | Dichrorampha teichiana | LEEUA626-11 | 3.1 |
Dichrorampha incognitana | N/A | 0 | Dichrorampha teichiana | LEEUA626-11 | 3.47 |
Dichrorampha inconspiqua | N/A | 0 | Dichrorampha infuscata | BTLBP373-11 | 0.93 |
Dichrorampha infuscata | N/A | 0 | Dichrorampha inconspiqua | BTLBP451-11 | 0.93 |
Dichrorampha ligulana | 1.17 | 1.76 | Dichrorampha petiverella | NOELE683-20 | 4.42 |
Dichrorampha melaniana | N/A | 0 | Dichrorampha sedatana | PHLAC013-10 | 0 |
Dichrorampha minutiana | N/A | 0 | Dichrorampha teichiana | LEEUA626-11 | 4.18 |
Dichrorampha obscuratana | 0.83 | 1.72 | Dichrorampha plumbagana | CGUKC685-09 | 3.6 |
Dichrorampha pentheriana | 0 | 0 | Dichrorampha teichiana | LEEUA626-11 | 2.82 |
Dichrorampha petiverella | 0.73 | 1.55 | Dichrorampha teichiana | LEEUA626-11 | 3.11 |
Dichrorampha plumbagana | 0.32 | 1.04 | Dichrorampha cinerosana | BTLBP382-11 | 2.76 |
Dichrorampha plumbana | 1.26 | 3.65 | Dichrorampha sedatana | PHLAC013-10 | 0 |
Dichrorampha podoliensis | 0.89 | 0.89 | Dichrorampha petiverella | TDAAT820-19 | 4.8 |
Dichrorampha rejectana | 0 | 0 | Dichrorampha inconspiqua | BTLBP451-11 | 3.64 |
Dichrorampha rilana | 0 | 0 | Dichrorampha teichiana | LEEUA626-11 | 3.82 |
Dichrorampha sedatana | 0.27 | 1.24 | Dichrorampha plumbana | PHLAJ255-14 | 0 |
Dichrorampha senectana | N/A | 0 | Dichrorampha sedatana | PHLAC013-10 | 0.15 |
Dichrorampha sequana | 0.44 | 0.92 | Dichrorampha teichiana | LEEUA626-11 | 5.31 |
Dichrorampha simpliciana | 1.29 | 3.46 | Dichrorampha teichiana | LEEUA626-11 | 4.92 |
Dichrorampha sylvicolana | 0 | 0 | Dichrorampha teichiana | LEEUA626-11 | 4.57 |
Dichrorampha tarmanni | 0.79 | 2.18 | Dichrorampha sedatana | PHLAC013-10 | 0 |
Dichrorampha teichiana | 0 | 0 | Dichrorampha flavidorsana | FBLMT309-09 | 2.05 |
Dichrorampha thomanni | N/A | 0 | Dichrorampha dinarica | PHLAE559-11 | 4.47 |
Dichrorampha vancouverana | 0.11 | 0.33 | Dichrorampha gueneeana | BTLBP389-11 | 0.15 |
Dichrorampha velata | 0.05 | 0.31 | Dichrorampha alpestrana | PHLAB566-10 | 3.96 |
Thus, based on male genital, wing pattern characteristics, and DNA barcode divergences the existence of two well defined species was postulated. However, the search for valid names of these two taxa was challenging.
Dichrorampha montanana (Duponchel, 1843)
Ephippiphora montanana Duponchel [1843]: 413, [1845]: pl. 83, fig. 7.
In the “Muséum national d’histoire naturelle” in Paris, Mr. Patrice Leraut kindly checked the D. montanana specimens of the collection. He found that J.D. Bradley (BMNH) had already dissected a syntype specimen of D. montanana Duponchel, a male, and that the genitalia slide still exists. Mr. Christian Gibeaux kindly forwarded us photographs of the genitalia preparation, the mounted specimen and the labels. The original description of the species does not give any indication to the number of examined specimens but according to
The lectotype of D. montanana obviously has a large-toothed phallus combined with a valval inner border bulge. Thus, specimens of the abovementioned group I represent D. montanana.
As synonyms of D. montanana Duponchel,
Grapholitha alpestrana [Zeller], 1843; Grapholitha alpestrana Herrich-Schäffer, 1851; Dichrorampha tanaceti Stainton, 1857; Dichrorampha herbosana Barrett, 1872; Hemimene blasiana Kennel, 1919; Hemimene modestana Müller-Rutz, 1922 [false 1992]
1) Grapholitha alpestrana [Zeller], 1843
Grapholitha alpestrana Herrich-Schäffer, 1851
Mr. Kevin Tuck from the British Museum-Natural History kindly informed us, that in the collection of his institution, there is no material of D. alpestrana originating from the Herrich-Schäffer collection (Herrich-Schäffer [1851]). One specimen of D. alpestrana comes from the collection of Zeller; it was however collected by Heinemann and cannot be considered as syntype.
In the Berlin Museum, Dr. Wolfram Mey allowed us to screen the Palaearctic Tortricidae collection for Herrich-Schäffer/Zeller material, unfortunately without success.
The Senckenberg Museum at Frankfurt, however, owns two specimens of Dichrorampha [alpestrana] montanana which obviously are from the time of discovery of this species. One male without abdomen is labelled “Gr.[apholita] Montanana FR, Styria, Fischer v.R.” with an additional minute red square label with nr. 7. A second male is labelled “Gr. Montanana FR. Styria Mann (FR vid.)” and therefore obviously has been identified by Fischer von Röslerstamm himself. Thus these specimen come closest to the original material Zeller must have based his description on but it cannot be proved that the material was studied by him. It is therefore not suitable for designation of a lectotype. Similarly two additional males from the Heyden collection in Frankfurt labelled as “Gr. Alpestrana” are not to be considered as syntypes as they have been collected by Mann at a different locality “Schneeberg” or just labelled insufficiently “Austr. Alp.”.
Dr. Wolfgang Nässig kindly allowed us to examine these specimens which, judging from wing pattern, all clearly represent D. alpestrana.
Finally Mr. Daniel Bartsch informed us that there is no European type material from Herrich-Schäffer in the Stuttgart Museum.
2) Dichrorampha tanaceti Stainton, 1857
The description of this species mentions “...macula magna triangulari dorsali dilutiore..” [with a big triangular diluted dorsal patch], thus describing the obvious differentiating wing pattern character of D. montanana [alpestrana]. Obrastzov (1953) depicted the male genitalia of D. tanaceti, which are identical with D. montanana. Then, in 1958, the same author formally synonymized D. tanaceti with D. montanana [alpestrana].
3) Dichrorampha monticolana described in detail by
4) Dichrorampha herbosana Barrett, 1872
The description says: “dorsal blotch triangular, oblique, pointed at the apex” [...] “Readily distinguished from the allied species by its pointed wings and distinct, pointed dorsal blotch..”, thus, referring it to D. montanana [alpestrana]. D. herbosana was already synonymized by
5) Hemimene blasiana Kennel, 1919
In the description of this species, Kennel mentions: “... ohne dass ein scharfer, heller Dorsalfleck gebildet wird...” [without there being formed a sharp, bright dorsal blotch], but the concomitant illustration depicts a male with such a blotch.
6) Hemimene modestana Müller-Rutz, 1922
The description of this new species is very detailed and particularly mentions the special form of the end of phallus with its marked curled peak. Mr. Daniel Burckhardt of the Basel Museum allowed us to study the two males labelled as “types” in the Müller-Rutz collection. Their wing pattern corresponds well with D. alpestrana, so do the nice illustrations in
7) Dichrorampha alpestrana ab. schatzmanni Rebel, 1927
Rebel, in 1927 published a description and a photograph of his Dichrorampha alpestrana ab. schatzmanni both of which fit perfectly the characteristics of the taxon of group I. According to article 45.6.2 of
8) Dichrorampha alpestrana f. olivana Müller-Rutz, 1934
Specimens of the type series were examined at the Basel Museum (Müller-Rutz collection). These specimens, originating from the Zermatt region, correspond well with the taxon of group I. However, the name was used infrasubspecifically for an alpine form and subsequently not adopted as the valid name of a species or subspecies. Following article 45.6.4 of
9) Dichrorampha pseudoalpestrana Danilevsky (in Obraztsov, 1953)
According to Obraztsov, Danilevsky described this taxon in order to give a valid name to the unavailable name “alpestrana”. There is a figure of the male genitalia which proves that this taxon is neither D. alpestrana nor the taxon of group II.
Further additional synonyms of D. montanana [alpestrana] listed by
According to our inquiries Zeller´s description of D. alpestrana is both valid and also published in advance of Duponchel´s work on D. montanana. From the imprinted date of the relevant issue of Stettiner entomologische Zeitung the description was published in May 1843. Duponchel´s description was published in fascicle 26 of Godart and Duponchel (1842–[1844]) which according to
Dichrorampha alpestrana ([Zeller], 1843) (Grapholitha) sp. rev.
= D. montanana (Duponchel, 1843) (Ephippiphora) syn. nov.
= D. alpestrana (Herrich-Schäffer, 1851) (Grapholitha)
= D. tanaceti Stainton, 1857 syn. rev.
= D. monticolana Heinemann, 1863, unjustidied emendation
= D. herbosana Barrett, 1872 syn. rev.
= D. blasiana (Kennel, 1919) (Hemimene) syn. rev.
= D. modestana (Müller-Rutz, 1922) (Hemimene) syn. rev.
Dichrorampha velata sp. nov.
= D. alpestrana ab. schatzmanni Rebel, 1927 (infrasubspecific, unavailable)
= D. alpestrana f. olivana Müller-Rutz, 1934 (infrasubspecific, unavailable)
Conclusion: Despite the fact that several authors introduced names within the “montanana/alpestrana”-complex, no valid name for the taxon of group II could be ascertained. Therefore, this species is newly described.
Dichrorampha velata is in overall appearance very similar to D. montanana from which it differs in wing pattern mainly by its more olive ground colour (in fresh specimens), by its markedly weaker dorsal blotch and by its slightly larger wingspan.
In male genitalia, the most obvious and constant differences are found in the shape of the phallus and the inner lobal line of the cucullus which both allow the unambiguous separation of the two taxa. In female genitalia, no clear differences could be ascertained.
“velatus” Latin, meaning “veiled” with respect to its confused history.
Material examined (specimens identified from genitalia preparations and/or DNA barcodes). Holotype. ♂, wingspan: 15.2 mm. CH-La Punt GR, God Arscheida [46.5864°N, 9.91928°E], 1820 m; 26.VI.2006; GP 108; BOLD: BC TLMF Lep 04060. Deposited in TLMF. Paratypes. Switzerland: 1♂: Felsberg GR, Sand [46.84541°N, 9.47091°E], 590 m, 24.6.2005, Schmid, BOLD 4061; 1 ♂: Avers, Innerferrera GR, [46.51657°N, 9.45399°E],1750 m, 29.6.2009, Schmid BOLD 4057; 1♂: Bergün GR, Pentsch [46.63913°N, 9.73708°E], 1500 m, 1.7.2009, Schmid, DNA Barcode ID TLMF Lep 04056; 2♂: Laax GR, Nagens [46.85122°N, 9.24287°E], 1820 m, 28.7.2005, Schmid, DNA Barcode ID TLMF Lep 0 4062; 2♂: La Punt GR, God Arscheida [46.5864°N, 9.91928°E], 1820 m, 26.6.2006, Schmid, DNA Barcode IDs TLMF Lep 04060, TLMF Lep 04063; 1♂: Avers-Cresta GR [46.47579°N, 9.50389°E], 1880 m, 15.7.2009, Schmid; 1♂: Tarasp-Fontana GR, Val Zuort [46.77299°N, 10.25755°E], 1440 m, 10.7.2008 Schmid; 1♂: La Punt GR, God Arscheida [46.5864°N, 9.91928°E], 1980 m, 30.6.2007, Schmid; 3 ♂: Tujetsch GR, Selva [46.66171°N, 8.2043°E], 1600 m, 20.6.2005, Schmid; 1♂, 1♀: Pigniu GR, Lag [46.82367°N, 9.11229°E], 1430 m, 21.6.2003, Schmid; 1♂: Tujetsch GR, Oberalp-Canals [46.65123°N, 8.68557°E], 1900 m, 1.8.2005, Schmid; 1♀: Sedrun, Bugnei [46.69012°N, 8.78537°E], 1700 m, 31.7.2004, Schmid; 1♂: Pigniu GR, Alp [46.82784°N, 9.10864°E], 1460 m, 10.7.2001, Schmid; 1♂: Medel GR, Acla [46.63106°N, 8.83805°E], 1520 m, 12.7.2001, Schmid; 1♂: Cormoret BE, Métairie de Morat [47.14394°N, 7.06376°E], 1500 m, 10.6.2000, Bryner; 1♀: Cormoret BE, Métairie de Morat [47.14394°N, 7.06376°E], 1500 m, 10.6.2000, Bryner; 1♂, 1♀: Villeret, Chasseral, Krete west [47.12969°N, 7.04969°E], 1550 m, 23.7.1994, Bryner. 1♂: Cormoret BE, Métairie de Morat [47.14394°N, 7.06376°E], 1500 m, 10.7.2003, Bryner; 1♂: Nods BE, Chasseral, Les Roches [47.13657°N, 7.07567°E], 1520 m, 15.7.2004, Bryner; 1♂: Nods BE, Chasseral, Piste [47.12568°N, 7.06243°E], 1285 m, 1.7.2013, Bryner; 1♂: Nods BE, Chasseral, sous les Roches [47.13398°N, 7.07345°E], 1380–1480 m, 13.6.2001, Bryner; 1♂: Villiers NE, Métairie de l’Ile [47.10633°N, 7.01728°E], 1350–1470 m, 14.6.2002, Bryner; 1♂: Ayers VS, Zinal, Pti Mountet [46.10759°N, 7.63174°E], 1800 m, 8.7.2015, Bryner; 1♂: Château-d’Oex, Béviau-Le Crinson [46.51371°N, 7.16527°E], 1260–1540 m, 22.6.2013, Bryner; 1♂: Zinal VS [46.13081°N, 7.62554°E], 1600–1850 m, 6.7.2020, Wittland; 1♂: Avers-Cröt GR [46.47681°N, 9.48857°E], 1750 m, 5.8.2011, Wittland; 1♂: Zinaltal VS, Le Vichiesso [46.09454°N, 7.63751°E], 1950–2140 m, 1.7.2014, Wittland; 3♂: Zinaltal VS, Le Vichiesso [46.11087°N, 7.63369°E], 1700–1850 m, 1.7.2014, Wittland: 1♂: Turtmanntal VS, Augstbordregion [46.2°N, 7.7666°E], 2400 m, 12.7.2010, Wittland; 1♂: Leuk VS, Guttet-Tschärmilong [46.47681°N, 9.48857°E] 1800 m, 8.8.2016, Wittland; 1♂: Leuk VS, Erschmatt-Brentschen [46.32883°N, 7.69125°E], 1550 m, 28.6.2017, Wittland; 1♂: Villeret BE, Chasseral [47.13194°N, 7.05477°E], 1430 m, 25.6.2020, Wittland.
Germany: 2♂: Immenstadt, Mittag [47.53833°N, 10.21861°E], 1450 m, 8.7.1983 Süssner/TLMF; 3♂: Schwäbische Alb, Urach, Nägeles Fels [48.50639°N, 9.38861°E], 700 m, 19.6.1970, Süssner/TLMF; 4♂: ditto, but 19.6.1968, Süssner/TLMF; 1♂: Schwäbische Alb, Urach 4 km SSW [48.48278°N, 9.37833°E], 630 m, 24.6.1975, Süssner/TLMF; 1♀: Schwäbische Alb, Urach [48.48278°N, 9.37833°E], 8.8.1954, Groschke/TLMF; 1♂: 2ditto, but 5.7.1955, Groschke/TLMF; 3♂: Schwäbische Alb, Neuffen-Hohenneuffen, Randweg [48.55833°N, 9.39167°E], 700 m, 19.6.1968, Süssner/TLMF; 1♀: ditto, but 30.6.1967, Süssner/TLMF; 1♀: Schwäbische Alb, Hohenneuffen [48.55833°N, 9.39167°E], 700 m, 8.7.1956, Süssner/TLMF; 1♀: ditto, but 30.6.1967 Süssner/TLMF.
Austria: Tirol, Umhausen [47.14027°N, 10.9290°E], 20.6.48, Burmann/TLMF; 2♂: Tirol, Umhausen N, unt. Farst [47.15694°N, 10.92278°E], 1100 m, 26.6.2010, Huemer/TLMF, DNA Barcode IDs TLMF Lep 03368, TLMF Lep 03607; 1♂: ditto, but 2.8.2014, Huemer/TLMF, DNA Barcode ID TLMF Lep 15225; 1♂: Tirol, Sölden [46.978°N, 11.002°E], 1600 m, 25.7.1956, Süssner/TLMF; 1♂: Salzburg, Grossglockner [47.0289°N, 6.40417°E], 1900 m, 1.7.1976, Zürnbauer/TLMF.
Italy: 1♂: Gr. St. Bernhard [45.883°N, 7.191°E], 2350 m, 1.7.1967, Zürnbauer/TLMF; 1♂: Südtirol, Vinschgau, Graun, Rojental [46.80722°N, 10.47889°E], 1970 m, 7.7.2013, Huemer/TLMF, DNA Barcode ID TLMF Lep 12339; 1♂: ditto, but 1860–1880 m, 1.7.2014, Huemer/TLMF, DNA Barcode ID TLMF Lep 15529.
France: 1♂: Rhône-Alpes, Le Corbier [45.23722°N, 6.26029°E], 1650 m, 17.7.2008, Nel/TLMF, DNA Barcode ID TLMF Lep 03370; 1♂: Auvergne-Rhône-Alpes, La Ville des glaciers [45.73555°N, 6.75555°E], 2200 m, 12.7.2007, Nel/TLMF, DNA Barcode ID TLMF Lep 03369; 2♂: Provence-Alpes-Côte d’Azur, Col du Lautaret [45.0289°N, 6.40417°E], 2058 m, 20.7.2006, Nel/TLMF, DNA Barcode IDs TLMF Lep 03376, TLMF Lep 03377.
Wingspan 12.8–16.5 mm (n = 25) mean: 14.6 mm. Forewing length: 6–8 mm. Head light grey, mixed with ochreous scales. Labial palpi dark grey, conspicuously ochreous at base. Proboscis pale yellow, antennae ochreous. Thorax and tegulae yellowish grey mixed with ochreous scales. Legs and abdomen grey with ochreous scales. Forewing ground colour olive brown or beige brown. Costal fold about one fourth of costal length. Costal strigulae darker brown alternating with creamy-white marks. Along termen variable number (3–5) of dark dots. Dorsal blotch faintly brighter than ground color, usually inconspicuous, pyramidal, with faint irregular darker strigulae. Silvery lines irregular, usually two more pronounced lines running parallel to termen. Cilial area composed of a line of short dark scales in front of a line of longer dark-tipped creamy-white scales. Hindwing grey, paler at base, with a dark-white-dark banded cilial line.
Male genitalia (Figs
Dichrorampha velata sp. nov., variation in wing pattern. 11. CH-Disentis, 1500 m, 9.7.2006; 12. CH-Avers-Cresta, 1880 m, 15.7.2009; 13. CH-La Punt, 1820 m, 26.6.2006; 14. CH-Tujetsch, 1600 m, 20.6.2005; 15. CH-Avers-Cresta, 1880 m, 15.7.2009; 16. CH-La Punt, 1980 m, 30.6.2007, all coll. J. Schmid.
Female (Fig.
Forewing ground color dark brown, suffused with ochreous scales. Markings like in male but darker and more contrasting. Dorsal blotch variably conspicuous.
Female genitalia (Figs
BIN: BOLD:AAE0715. The intraspecific average distance of the barcode region is very low with only 0.05%, the maximum distance 0.31% (p-dist) (n = 18). The minimum distance to the nearest neighbor, D. alpestrana, is 3.96%.
(Fig.
Contrarywise D. alpestrana occurs only in the southern part of the Swiss Alps, i.e. in a region south of the Rhine-Rhône line. It is widely distributed in Austria but the only species of the group in the eastern part of the country (type locality of D. montanana and D. alpestrana). Further proved records come from the Italian Alps, Northern Macedonia and from the United Kingdom, indicating a highly disjunct arctic-alpine distribution pattern.
In some localities in Switzerland and Italy both species have been observed in sympatry.
Dichrorampha alpestrana (specimens identified from genitalia preparations and/or DNA barcodes). Switzerland: 12♂ 4♀: La Punt GR, God Arscheida [46.5864°N, 9.91928°E], 1820 m, 26.6.2006, Schmid, 1♂ DNA Barcode ID TLMF Lep 01366; 1♂: La Punt GR, God Arscheida [46.5864°N, 9.91928°E], 1820 m, 19.6.2006, Schmid; 3♂: Val Müstair GR, Sta Maria [46.60764°N, 10.41349°E], 1760 m, 30.6.2008, Schmid; 1♂ Avers, GR, Innnerferera, Starlera [46.5167°N, 9.43308°E], 1750 m, 30.6.2008, Schmid, DNA Barcode ID TLMF Lep 04058; 3♂: Berninapass Süd GR [46.41390°N, 10.03627°E], 2300 m, 16.7.2008, Schmid; 1♂ DNA Barcode ID TLMF Lep 04059; 1♂: Valposchiavo GR, Bernina Ospizio [46.41056°N, 10.02278°E], 2350 m, 13.07.2009, Schmid; 1♂: Pontresina GR, Lagalb, 2100 m, 10.8.2010, Schmid; 1♂: Ardez GR [46.77352°N, 10.20102°E], Thomann TLMF; 2♂: Simplonpass VS, Bistinealp [46.25803°N, 8.03124°E], 2000–2200 m, 15.7.2010, Wittland; 1♂: S-chanf GR, Alp Chaschauna, 2250–2800 m, 25.7.2012, Wittland. Austria: 1♂: Salzburg, Katschberg [47.00605°N, 13.60972°E], 1750–1850 m, 2.8.1991, Huemer & Karsholt/TLMF; 1♂: Osttirol, Virgen-Obermauern [47.00645°N, 12.43286°E], 1410 m, 8.7.1993, Tarmann/TLMF; 1♂: Osttirol, Rieserfernergruppe, Patschertal [46.92477°N, 12.18629°E], 2080 m, 158.1989, TLMF; 1♂: Osttirol, Matrei [46.99966°N, 12.54313°E], 1200 m, 2.6.1963, Burmann/TLMF; 1♂: Nordtirol, Vennatal [47°N, 11.55°E], 2000 m, 2.8.1969, Kapeller/TLMF; 1♂: Nordtirol, Stanzach [47.39527°N, 10.56722°E], 920 m, 5.7.1989, Huemer/TLMF; 1♂: Osttirol, Dorfertal [47.04297°N, 12.33361°E], 1880–2100 m, 28.7.1988, Tarmann/TLMF; 1♂: Osttirol, Kals, Tauernhaus [47.0707°N, 12.6233°E], 1700 m, 5.7.1960, Süssner/TLMF; 1♂: Osttirol, St. Jakob in Defereggen [46.91722°N, 12.33083°E], 1380 m, 12.7.2002, Deutsch/TLMF, DNA Barcode ID TLMF Lep 03366; 1♂: Osttirol, Ködnitztal, Greiwiesen [47.01944°N, 12.6819°E], 2100–2300 m, 21.7.2002, Deutsch/TLMF, DNA Barcode ID TLMF Lep 03367; 1♂: Steiermark, Turracher Höhe NW, [46.92805°N, 13.86805°E], 1750–1850 m, 4.7.2009, Huemer/TLMF, DNA Barcode ID TLMF Lep 00818; 1♂: Kärnten, Saualpe, Umg. Wolfberger Hütte [46.83333°N, 14.65°E], 1500–1800 m, 19.6.2000, Wimmer/TLMF; 1♂: Niederösterreich, Seeau S Hollenstein/Ybbs [47.75°N, 14.78°E], 600 m, 1.6.1986, Lichtenberger /TLMF. Italy: 1♂: Südtirol, Vinschgau, Graun, Rojental [46.80722°N, 10.47889°E], 1970 m, 7.7.2013, Huemer/TLMF, DNA Barcode ID TLMF Lep 12340; 1♂: ditto, but 1860–1880 m, 1.7.2014, Huemer/TLMF, DNA Barcode ID TLMF Lep 15528; 1♂: Südtirol, Ridnauntal [46.938°N, 11.256°E], 1500–1580 m, 8.6.2016, Wittland. Northern Macedonia: 2♂: NP Mavrovo, Korab, Korabska jezero, Kobilino pole [41.77833°N, 20.58194°E], 2080–2180 m, 28.7.-1.8.2011, Huemer & Tarmann/TLMF, DNA Barcode IDs TLMF Lep 05060, TLMF Lep 05061.
Furthermore, published genitalia preparations prove the occurrence of D. alpestrana in:
Austria: Niederösterreich, Sonnwendstein [47.63030°N, 15.86097°E], 1500 m, 15.7.2005, Buchner/Lepiforum; Steiermark, Ponigl bei Weiz [47.27077°N, 15.63122°E], ca. 800 m, 6.6.2020, Pichler/Lepiforum; Steiermark, Fischbacher Alpen, St. Kathrein am Offenegg [47.3°N, 15.56666°E], 1400 m, 24.7.2019, Pichler/Lepiforum; Kärnten, Heiligenblut/Gößnitztal, Innere Ebenalm, , Strutzberg [47.02653°N, 12.78645°E], 1800 m, 27.7.2019, BOLD (ABOL-Bioblitz 2019 19–1740/Lepiforum; Steiermark, Graz, Schöckl [47.19858°N, 15.46567°E], 1400 m, 2.7.2020, Pichler/Lepiforum. United Kingdom: Shropshire, Chapel Lawn, 10.6.2006, Clement/mothdissection.co.uk; RIS TrapTregaron, Ceredigion vc-46, 7.2013, Tillotson/mothdissection.co.uk; Bettws GG, North Wales, 6.2019, Graham/mothdissection.co.uk. A male adult from the Polish Tatra mts. figured by
Dichrorampha velata sp. nov. occupies a wide variety of biotopes from dry lowland pastures to alpine grassland above the treeline. It would seem, however, that the species prefers montane to subalpine borders of fertile meadows at woodland edges. The moths have been observed flying during daytime in the vicinity of Leucanthemum sp. and Achillea sp., in the roots of the latter, caterpillars of Dichrorampha vancouverana McDunnough, 1935 and Dichrorampha ligulana (Herrich-Schäffer, 1851) were found topotypical. However, attempts at finding caterpillars of D. velata/alpestrana were unsuccessful so far, though D. alpestrana is recorded from the rootstocks of Achillea ptarmica (
Nearly 30 years ago it became evident that “Dichrorampha montanana sensu auct.” morphologically is a complex of probably two species, a hypothesis later supported by genetic data. However, the subsequent search for type material became extremely tedious due to confounded taxonomy, scattered type material and cumbersome institutional bureaucracy.
The initial claim by
While wing phenotype is a good indicator of taxon identity in most fresh specimens, in single cases and especially when specimens are worn, this trait may be doubtful. Thus, genitalia preparations and/or DNA barcode results are needed for unambiguous identification.
In male specimens, shape and number of phallus end-thorns are very good differentiating characteristics. Unfortunately, in female genitalia, no clear distinguishing features could be detected so far; clearly more material will be necessary for future research.
The present geographical distribution, as far as it is known, suggests one or several south-eastern glacial refugia of D. alpestrana, while D. velata sp. nov. likely is of north-western peri-alpine glacial provenance (Fig.
The authors would like to thank Kevin Tuck, Natural History Museum London for very valuable information, Wolfram Mey, Museum für Naturkunde Berlin for his kind hospitality in his museum, Wolfgang. Nässig, Senckenberg Musum Frankfurt for allowing us to see the collections, Daniel Burckhardt, Naturhistorisches Museum Basel for his hospitality and the loan of material, Daniel Bartsch, Staatliches Museum für Naturkunde Stuttgart; Patrice Leraut, Muséum national d’histoire naturelle Paris for information, Christian Gibeaux for providing photographs of the D. montanana lectotype, Rudolf Bryner, Biel/Bienne and Wolfgang Wittland, Wegberg-Dalheim for data and loan of material; Paul D.N. Hebert and the entire team at the Canadian Centre for DNA Barcoding (Guelph, Canada) for continuous support with sequencing work. The study was furthermore supported by the Promotion of Educational Policies, University and Research Department of the Autonomous Province of Bolzano - South Tyrol with funds to the projects “Genetische Artabgrenzung ausgewählter arktoalpiner und boreomontaner Tiere Südtirols” and “Erstellung einer DNA-Barcode-Bibliothek der Schmetterlinge des zentralen Alpenraumes (Süd-, Nord- und Osttirol)”.