Research Article |
Corresponding author: Axel Kallies ( axel.kallies@unimelb.edu.au ) Academic editor: Maria Heikkilä
© 2018 Axel Kallies, Teresa Farino.
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:
Kallies A, Farino T (2018) A new species of Pharmacis Hübner, 1820 from Spain with a brief review of the genera Pharmacis and Korscheltellus Börner, 1920 (Lepidoptera, Hepialidae). Nota Lepidopterologica 41(2): 225-249. https://doi.org/10.3897/nl.41.26835
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We here describe a new ghost moth (Hepialidae) species, Pharmacis cantabricus sp. n. from the Picos de Europa National Park, Cantabria, in northern Spain. The new species belongs to a group of mostly day-flying species that are restricted to the European Alps and some mountain ranges of southern Europe. Based on morphology and analysis of mitochondrial COI gene sequences, the new species is closely related to Pharmacis aemilianus (Constantini, 1911), an endemic of the Italian Apennines. However, Pharmacis cantabricus sp. n. can easily be distinguished from all related species based on both external and genitalic characters. We briefly review and illustrate all species of the genus Pharmacis Hübner, 1820 and discuss its relationship with the related genus Korscheltellus Börner, 1920. We reinstate Hepialus castillanus Oberthür, 1883 as a distinct species and transfer it to Korscheltellus (stat. rev., comb. n.).
Describimos una nueva especie de Hepialidae, Pharmacis cantabricus sp. n. del Parque Nacional de Picos de Europa, Cantabria, España septentrional. La nueva especie pertenece a un grupo de especies de vuelo diurno cuya distribución se limita a los Alpes europeos y otras cadenas montañosas de Europa meridional. Según estudios morfológicos y análisis de secuencias del gen mitocondrial COI, la nueva especie es más próxima a Pharmacis aemilianus (Constantini, 1911), un endemismo de los Montes Apeninos en Italia. Sin embargo, Pharmacis cantabricus sp. n. se distingue fácilmente de todas las especies relacionadas en sus caracteres externos y genitales. Evaluamos brevemente e ilustramos todas las especies del género Pharmacis Hübner, 1820 y comentamos su relación con el género emparentado Korscheltellus Börner, 1920. Restituimos Hepialus castillanus Oberthür, 1883 como especie y la trasladamos a Korscheltellus (stat. rev., comb. n.).
The ghost moths (Hepialidae) constitute a family of evolutionarily primitive moths that occur worldwide with >500 species (
In 2007, the second author photographed a mating pair of Hepialidae in the Picos de Europa Mountains in northern Spain that was initially considered to belong to Korscheltellus fusconebulosa (De Geer, 1778) (Fig.
In order to place the new species, we review all species of the genus Pharmacis and the related genus Korscheltellus Börner, 1920, and use morphological and molecular evidence to support species delineation. The results of these studies and the description of the new species, Pharmacis cantabricus Kallies & Farino, sp. n., are presented here.
1. P. cantabricus sp. n., mating pair, Spain, Cantabria, Picos de Europa, Sierra Bejes. 2. Type locality of P. cantabricus sp. n. at the Sierra Bejes at about 800 m altitude. 3. Habitat of P. cantabricus sp. n. above the Lagos de Covadonga, at about 1375 m altitude. Image 3, F. González.
The genus Pharmacis comprises a small group of medium-sized Palaearctic ghost moths.
The considerable list of synonyms and intrasubspecific names linked to some of these taxa can be found in
Pharmacis Hübner, 1820
P. carna ([Denis & Schiffermüller], 1775)
P. claudiae Kristal & Hirneisen, 1994
P. anselminae (Teobaldelli, 1977)
P. bertrandi (Le Cerf, 1936)
P. pyrenaicus pyrenaicus (Donzel, 1838)
P. pyrenaicus alticola (Oberthür, 1881)
P. aemilianus (Constantini, 1911)
P. cantabricus sp. n.
Korscheltellus Börner, 1920
K. fusconebulosa (De Geer, 1778)
K. castillanus (Oberthür, 1883), stat. rev., comb. n.
K. gracilis (Grote, [1865])
K. lupulina (Linnaeus, 1758)
COI sequences were downloaded from BOLD (http://www.boldsystems.org/index.php/) or generated using the methodology described in detail by
In order to place the new species confidently within the genus Pharmacis and to test whether currently recognized species in the genus were supported by molecular data, we utilized mitochondrial COI DNA barcodes. Analysis of the COI sequences revealed that all species analysed in this study formed well separated monophyletic entities, supporting the notion that they are valid taxa (Fig.
Species of the genus Pharmacis occur in the mountains of central, southern and south-western Europe, with one species, P. carna, also occurring in Fennoscandia. Some of the species show an extremely limited distribution, including P. bertrandi, P. anselminae and P. claudiae, which occur exclusively in small, high-altitude areas in the French and Italian Alps. Although both P. pyrenaicus and P. aemilianus have larger ranges, they too are restricted to relatively small regions covering the Pyrenees and Abruzzi, respectively. Notably, several of the Pharmacis species, including P. anselminae, P. bertrandi and P. pyrenaicus, have brachypterous females, probably an adaptation to the harsh conditions of the alpine environment in which they occur. Two of these species, P. anselminae and P. pyrenaicus, appear to be strictly day-flying, a feature shared by many moths that occur in alpine environments or high latitudes. The third species with brachypterous females, P. bertrandi, was found to be active in the second half of the night and in the morning. In contrast, P. claudiae and P. aemilianus have fully winged females and fly during the night time. Pharmacis carna on the other hand occurs from the foothills to the alpine regions of the Alps and other mountain ranges. It has been observed at night in some areas, whereas it was found to be active predominantly during the day in other areas (
Abbreviations.
Holotype (Fig.
Male (Figs
Female (Figs
Male genitalia (Fig.
Female genitalia (Fig.
Pharmacis species. 5–8. P. cantabricus sp. n., Spain, Picos de Europa. 5. Male, holotype, alar expanse 33 mm (
Pharmacis cantabricus sp. n. shows considerable variability in the extent of the white markings on the forewing. Specifically, the spots between A and CuA2 can be reduced or confluent, the white streak in the base of the forewing can be more or less extensive. Similarly, the extent of the light brown areas of the forewing varies. The female of a mating pair figured on
Males of P. cantabricus sp. n. can be differentiated from males of all other species by the white forewing markings, which are usually reduced to an often ill-defined white streak near the base, an irregular white patch just anterior of vein A, and a third white patch between veins A and CuA2 near the anal angle. The extent of these white markings is, however, variable. In some specimens they are further reduced or even absent (as in the female in Fig.
Based on DNA barcoding, morphology of the genitalia and labial palps and the number of antennal segments, the new species appears to be closely related to P. aemilianus (Figs
Pharmacis pyrenaicus (Figs
Finally, P. cantabricus sp. n. differs from all species of Korscheltellus in the shape of ventral extension of the pseudotegumen (spatulate and simple in Korscheltellus, three-dimensional and furrowed in P. cantabricus sp. n.) and the forewing pattern. From both K. fusconebulosa and K. castillanus stat. rev., comb. n. the new species also differs by the monochrome brown fringes of fore and hindwings (chequered in the two species compared).
Pharmacis species. 13. P. bertrandi, male, Italy, Cuneo, alar expanse 38 mm (CAM). 14–16. P. carna. 14. Male, Austria, Oberösterreich, alar expanse 33 mm (CEF). 15. Male, Austria, Steiermark, alar expanse 35 mm (CEF). 16. Female, Italy, Lombardia, alar expanse 40 mm (CAK). 17. P. anselminae, male, Italy, Valle de Aosta, alar expanse 33 mm (CAK). 18. P. claudiae, male, Valle de Aosta, alar expanse 41 mm (CAK). Images 13, H. Deutsch; 14–15, E. Friedrich.
Pharmacis cantabricus sp. n. inhabits open grassy woodlands and rich meadows and pastures in montane and subalpine parts of the Picos de Europa. In the Sierra de Bejes, the authors found P. cantabricus in open Quercus petraea (Matt.) Liebl. woodland with Corylus avellana L. on a north-west-facing ridge of Carboniferous limestone at about 800 m altitude (Fig.
The second known locality is considerably different. It is situated above the Lagos de Covadonga at about 1375 m (Fig.
Males of P. cantabricus sp. n. were seen flying rapidly over the vegetation between the late morning and afternoon. Mating was observed in the afternoon. Detailed observations were made in the Sierra de Bejes on the 18 July 2017. On arrival at the site at 12.45 pm, the weather was sunny, 24 °C, with light winds. The cloud cover was 40–50% initially, but increased to about 90% by 4.40 pm, with only a few sporadic sunny intervals later. A total of 7 males were caught between 1.35 and 5.05 pm. One male was caught at 3.00 pm, when it was attracted to a female hanging in grasses by the side of the track (Fig.
The new species is currently known only from a small area that includes the Picos de Europe and the surrounding area in the provinces of Asturia and Cantabria in north-western Spain.
Korscheltellus species. 19–21. K. fusconebulosa. 19. Male, Italy, Vintschgau, alar expanse 36 mm (CEF). 20. Male, Germany, Schmalkalden (CEF). 21. Female, Germany, Schmalkalden (CEF). 22. K. castillanus, female, alar expanse 32 mm (
The highly restricted distribution of most of the Pharmacis species is probably a result of frequent glaciation events during the Pleistocene or earlier ice ages. During these periods, Pharmacispopulations may have withdrawn into isolated refugia in the mountains of southern Europe and undergone speciation. Such a scenario is exemplified by the distribution of the three species of the P. carna species group (P. anselminae, P. bertrandi and P. claudiae), each of which is represented by small isolated populations in the French and Italian Alps. The hypothesis of glaciation-driven speciation in Pharmacis may be supported by the relatively low pairwise sequence differences between closely related species (2–2.7%) observed here. Based on the estimated 1.0–2.3% per million years (Mya) average substitution rate for COI (
The new species, P. cantabricus sp. n., is currently known only from within and around the Picos de Europa, a small mountain range at the convergence of the provinces of Asturias, Cantabria and León in northern Spain. In contrast to the high alpine environment inhabited by other highly localized Pharmacis species, P. cantabricus sp. n. was found in open oak (Quercus) woodland in the montane zone and in subalpine pastures. However, additional observations suggest that it also occurs at much lower altitude, and it remains to be seen how widely P. cantabricus sp. n. is distributed in northern and north-western Spain. Pharmacis aemilianus, the closest relative of the new species, occurs in Italy in a range of different habitats ranging from about 200 to 2000 m altitude (
The restricted distribution of several of the Pharmacis species is likely to be supported by the limited potential for dispersal owing to the brachyptery of the females. Interestingly, it appears that brachyptery in Pharmacis females has arisen at least twice independently, as it can be found in the P. carna group, specifically in the sister species P. anselminae and P. bertrandi, and in P. pyrenaicus, which according to our molecular analyses belong to different clades within the genus. Although female P. cantabricus sp. n. have well-developed wings, they are short in comparison to the abdomen (Fig.
We have here followed the generic division of Pharmacis and Korscheltellus proposed by
Finally, the generic placement of some species from Central Asia and the Eastern Palaearctic that are currently assigned to the genus Thitarodes should be reassessed as they show similarities to Korscheltellus. This includes T. nebulosus (Alphéraky, 1889), a species considered to be closely related to K. fusconebulosa by
Notably,
The hepialid fauna of Spain was last reviewed by
Live Pharmacis specimens. 25. P. anselminae, male, Italy, Valle de Aosta. 26. P. bertrandi, male, France, Haute Alps. 27. P. carna, male, Poland, Tatra Mts,. 28. P. aemilianus, male, Italy, Central Apennines. 29–31. P. cantabricus sp. n., males (29, 30), female (31), Spain, Picos de Europa. Images 25, R. Bryner; 26, A. Longieras; 27, J. Wenta; 28, P. Mazzei.
P. aemilianus (Constantini, 1911)
Figs
This species is an Italian endemic, which occurs in most parts of the mainland, ranging from about 200 to 2000 m altitude (
Material examined. 1 ♂, 1 ♀, Italy, Abbruzi, Pescocostanzo (
P. pyrenaicus (Donzel, 1838)
Figs
This species is endemic to the Pyrenees (Spain, France and Andorra), where it occurs at altitudes between 1800 and 2800 m (
Material examined. P. pyrenaicus pyrenaicus: 1 ♂, France S, Pyrenees-O., Fort Rome NW, Col de Puymorens, 2000–2500 m, 13.vii.2003, leg. Andree Salk (CAK, genitalia prep. AK820, DNA AK350, Figs
Live Pharmacis specimens. 32. P. cantabricus sp. n., mating pair, Spain, Picos de Europa. 33. P. carna, mating pair, Poland, Tatra Mts. 34. P. anselminae, mating pair, Italy, Valle de Aosta. 35. P. pyrenaicus, female, Spain, Pyrenees. Images 33, J. Wenta; 34, R. Bryner; 35, E. Beltrán.
P. bertrandi (Le Cerf, 1936)
Figs
This species was described from the French Alps (
Material examined. 15 ♂, Italy, Prov. Cuneo, Gias Valcavera, 2050 m, 44°22.6’N, 07°08.2’E, 23. and 27.vii.2009, 17.vii.2012, 22.vii.2013, leg. A. Mayr (CAM, CAK, Fig.
P. carna ([Denis & Schiffermüller], 1775)
Figs
This species is distributed throughout the higher central European mountains, including the Alps and Carpathian Mts., also in Fennoscandia. Records from the Ural Mts (
Material examined. 1 ♂, Austria, Oberösterreich, Weyer, Küpfern, train station, 400 m, 11.vii.1999, at light, leg. E. Friedrich (CEF, Fig.
P. anselminae (Teobaldelli, 1977)
Figs
This species is only known from the Aosta valley in NW Italy at altitudes between 1800 and 2500 m. It inhabits alpine meadows and pastures. Males fly rapidly in the sun, starting in the morning and continuing until late afternoon, with peak activity between 10 am and noon. They stop flying when the sun disappears behind clouds (
Material examined. 15 ♂, 2 ♀, Italy, Val d’Aosta, Val Valleile, Cogne, 1950 m, 11.vii.1990, leg. A. Kallies (CAK, genitalia prep. AK863, Figs
P. claudiae Kristal & Hirneisen, 1994
Fig.
This species was described relatively recently from the Aosta Valley in north-western Italy (
Material examined. 1 ♂, Italy, Val d’Aosta, Valtourrenche, A.-St. Andre, Torgnon, Alpi Clogne, 2100–2200m, 25.vii.1994, leg. P.M. Kristal & J. Roth (CAK, Fig.
K. fusconebulosa (De Geer, 1778)
Figs
This species is widespread and occurs from Western Europe to Japan (
Material examined. 1 ♂, Germany, Schmalkalden, JH ‘Ebertswiese’, 730 m, at light, 30.vi.–2.vii.1989, leg. E. Friedrich (CEF, Fig.
K. castillanus (Oberthür, 1883), stat. rev., comb. n.
Fig.
This taxon is insufficiently known. It was treated as a member of the genus Pharmacis by
Korscheltellus castillanus is similar to K. fusconebulosa but smaller and less vividly coloured, with the wingspan ranging from 27–28 mm in males to 32 mm in females (males 28–35 mm, females 35–46 mm in K. fusconebulosa). According to the figures provided by
K. castillanus is known from only a small number of specimens that have been collected in a small area north-west of Madrid (La Granja de San Ildefonso, Balsain, El Escorial, in the Sierra de Guadarrama), and no new material has been found in the last 70 years. According to
Material examined. 1 ♀, [Spain] Sn Ildef [San Ildefonso], 84, m / 1/6 / ♀ [unreadable] Stgr. (
K. gracilis (Grote, 1865)
This is a North American species, which has been discussed in detail elsewhere (
K. lupulina (Linnaeus 1758)
Figs
This species is widespread in Europe and well recorded from the northern part of Spain (
Material examined. 1 ♂, Spain, Leon, La Una, 15.vi.2013, leg. González Estébanez (CAK, genitalia prep. AK816, Fig.
Pharmacis and Korscheltellus, male genitalia (42–45) and terminal abdominal sclerites (46–48). 42. P. pyrenaicus pyrenaicus (pseudotegumen flattened). 43. P. pyrenaicus alticola (pseudotegumen in natural position). 44. P. anselminae. 45. K. lupulina. 46. P. cantabricus sp. n. paratype (stained). 47. P. anselminae (unstained). 48. P. pyrenaicus pyrenaicus (unstained). Images 46–48, Q. Wang.
We would like to express our gratitude to the large number of individuals who have contributed to this work. Only through their support was this study made possible in its current form. We thank John Grehan for alerting the first author to the images of P. cantabricus sp. n., and thereby initiating this study. We thank Yeray Monasterio León, Ruth Escobés Jiménez and Félix Javier González Estébanez for their invaluable help in observing and collecting type specimens of P. cantabricus sp. n. We thank Joël Minet, Vadim Zolotuhin, John Grehan, Axel Steiner, Thomas Witt and Josef de Freina for information regarding type material and for help collating relevant literature. We express our gratitude to Andree Salk and Tony Mayr for donating material used in this study. We thank Antoine Longieras, Mauro Gianti, Giovanni Delmastro and Tony Mayr for information concerning the distribution and habits of P. bertrandi, Antoine Longieras, Rudolf Bryner and Helmut Deutsch for permission to use their images, Enrique Murria Beltrán for images and specimens of P. pyrenaicus, Jarosłav Wenta for images and information relating to P. carna, Paolo Mazzei for images and information relating to P. aemilianus, Egbert Friedrich for images, and Yeray Monasterio León, Marcos Toribio, Colin W. Plant, Zdeněk Laštůvka, Martin F. V. Corley and Roger Vila for information concerning the distribution of Iberian Hepialidae. We are grateful to Axel Hausmann and Peter Huemer for providing sequences used in this study, and to Wolfram Mey for access to type material under his care. Furthermore, we wish to thank Tom McConville and Liz Milla for performing COI sequencing and analysis, and Qike Wang for his help with imaging. We would like to thank John Grehan, Thomas Simonsen and Carlos Mielke for helpful comments on the manuscript. Finally we would like to thank the Consejería de Medio Rural, Pesca y Alimentación del Gobierno de Cantabria and the Parque Nacional de Picos de Europa for providing us with the necessary licences to carry out the field work.