Pharmacis cantabricus, Kallies, Axel & Farino, Teresa, 2018

Kallies, Axel & Farino, Teresa, 2018, A new species of Pharmacis Huebner, 1820 from Spain with a brief review of the genera Pharmacis and Korscheltellus Boerner, 1920 (Lepidoptera, Hepialidae), Nota Lepidopterologica 41 (2), pp. 225-249 : 228-234

Pharmacis cantabricus sp. n. Figs 1, 5-8, 29-31, 32, 39, 41, 46

Type material.

Holotype (Fig. 5): 1 ♂, N Spain, Cantabria, Picos de Europa National Park, Sierra de Bejes, ca 800 m, 43.247N, 4.651W, 9. July 2016, day flying, leg. A. Kallies & T. Farino (CAK, in MfN). Paratypes: 2 ♂, same data as holotype (genitalia prep. AK790, Figs 6, 39, 46); 7 ♂, 1 ♀, same locality, 18. July 2017, day flying, leg. Y. Monasterio, T. Farino & R. Escobés (CAK, specimens will be lodged in MfN, ZSM and IBEB, Figs 7, 29-32); 2 ♀, N Spain, Asturias, Picos de Europa National Park, Lagos de Covadonga, ca 1375 m, 43°14 ’37.10” N, 4°59 ’44.23” W, 20. July 2007, during the day, leg. F. J. G. Estébanez (CFG, genitalia prep. AK861, Figs 8, 41).

Description.

Male (Figs 5, 6, 29, 30, 39, 46). Alar expanse 29-33 mm, forewing length 13-15.5 mm. Head and thorax ochre-brown to greyish, covered with soft and relatively long, hair-like scales. Scales between eye and antenna present. Antenna consisting of 30-31 segments that are short, wide and relatively flat, resulting in an overall flattened (lamellate) antenna morphology. Labial palps well-developed, protruding well beyond the frons; three-segmented, with the three palpomeres well separated, the middle palpomere about twice as long as the apical one, the apical palpomere globular; dorsally with short scales, ventrally with long, hair-like scales. Forewings dark brown, along the costa and the area of the radius suffused with light ochre-brown; with an ill-defined whitish streak close to base; with an irregular white patch anterior of vein A; with another white patch between veins A and CuA2 near the anal angle, sometimes extending to the area between CuA1 and CuA2; with a few very small, ill-defined white patches postmedially; with a small white spot in the area of the cell at the stem of M2 and another small white spot at the basal fork of M1; all white markings embedded into light brown areas. Hindwing uniformly dark brown. Fringes of both fore and hindwing short and uniformly dark. Fore tibia with a distinct epiphysis, which consists of an open tube-like structure, ending in a short and wide tooth. Abdomen dorsally ochre to grey, ventrally light ochre, posterior margins of tergites light ochre. Abdominal sclerites (Fig. 46) poorly sclerotized, tergites 7 and 8 simple, sternite 7 laterally with a membranous spot on each side; sternite 8 roughly triangular and relatively long.

Female (Figs 7, 8, 31). Alar expanses 30-35 mm, forewing length 15-17.5 mm. Similar to male, but wings narrower, with 26-29 antennal segments, the extent of the white forewing markings variable. The white forewing markings of the female paratype from the type locality are extensive and similar to the markings of the males from the same location. However, the female photographed at the same location (Fig. 1, right specimen) has reduced white marks. One female from the Lagos de Covadonga lacks any distinct white markings (Fig. 8).

Male genitalia (Fig. 39). Vinculum with long processes (vincular condyles sensu Grehan 2012) at posterior margin; saccus rounded; the paired apical processes of the pseudotegumen long, each with a small protrusion at the distal margin, not fused; ventral extension of the pseudotegumen (trulleum sensu Zilli 1988 and Simonsen 2018; spatulate mesosome sensu Wagner 1988 and Grehan 2012) large and well sclerotized; valva relatively straight, long and narrow, without sub-basal tooth at the ventral margin.

Female genitalia (Fig. 41). Papillae anales narrow and setose; lamella antevaginalis shortly setose, with two submedial invaginations forming ventricle-like structures; each a very small sclerotized plate on either side of the ostium; corpus bursae ovoid, membranous; ductus bursae long and simple.

Variability.

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 Biodiversidadvirtual (2018, as Pharmacis sp.) has dark grey instead of brown forewings. The corresponding male is ochre-brown, and both specimens have the white mark reduced to the discal spot. In some females the white marks can be absent altogether (Fig. 8).

Diagnosis.

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. 8), in others they form an almost continuous white line (as in female in Fig. 7). Pharmacis cantabricus sp. n. differs from most other species of Pharmacis , with the exception of P. aemilianus , by the well-developed labial palps. Although the labial palps of all Pharmacis species consist of three segments, they markedly differ in their structure. In P. cantabricus sp. n. all segments are well-separated, and while the middle palpomere is very long, the apical palpomere is very short and distinctly bulbous. In all other species, with the exception of P. aemilianus , the labial palps are much shorter, and the palpomeres are of similar length, with the apical one pointed and partially fused to the middle segment. Furthermore, in P. cantabricus sp. n. and P. aemilianus , the labial palps are covered with short scales dorsally and long and thin scales ventrally, while they are clothed in long hair-like scales dorsally and ventrally in all other species. Diagnostic characters can also be found in the abdominal sclerites, which are poorly sclerotized in P. cantabricus sp. n. (Fig. 46) and P. aemilianus (not shown), while they are well sclerotized in the other species examined, P. anselminae (Fig. 47) and P. pyrenaicus (Fig. 48). Sternite 7 is relatively wide in P. cantabricus sp. n. and P. aemilianus while it is narrow in the distal half in other species examined ( P. pyrenaicus and P. anselminae ). Sternite 8 is roughly triangular and relatively long in P. cantabricus sp. n. and P. aemilianus , while it is quadrangular and short in P. anselminae (Fig. 47) and very wide and short in P. pyrenaicus (Fig. 48).

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 9, 10) from Italy. Both species have about 29-30 antennal segments, well-developed labial palps and a well-developed ventral extension of the pseudotegumen of the male genitalia. However, these two species can be easily separated by external characteristics such as wing coloration (yellow-brown to ochre-brown in P. aemilianus ; dark brown with light brown markings in P. cantabricus sp. n.), white forewing markings (typically reduced and often with just a white mark prominent in the cell in P. aemilianus ; typically with larger white markings along the posterior section of the forewing in P. cantabricus sp. n.), the size (substantially larger, alar expanse in males 30-36 mm and in females 36-52 mm in P. aemilianus ), and wing shape (in P. aemilianus the forewings longer and narrower in males, broader in females). The genitalia of both species are very similar. Males of both species display a well-developed ventral extension of the pseudotegumen and large vincular condyles. Similarly, the ventricle-like invaginations in the lamella antevaginalis of the female genitalia is present in both species. However, P. cantabricus sp. n. differs from P. aemilianus in a number of details: the apical part of the valva is shorter and narrower in P. cantabricus sp. n. (longer and expanded in P. aemilianus ), the paired ventral processes of the pseudotegumen are longer, and the apical margins of pseudotegumen have a small protrusion (absent in P. aemilianus ), the saccus is more rounded (tapering to a point in P. aemilianus ) and the processes of the vinculum are tapering and bent outwards (wider and bent inwards in P. aemilianus ). Furthermore, both species differ in their behaviour, with P. aemilianus being active at dusk and during the night, whereas P. cantabricus sp. n. appears to be strictly day-flying. Finally, females of P. aemilianus are able to fly freely, which does not appear to be the case for female P. cantabricus sp. n.

Pharmacis pyrenaicus (Figs 11, 12, 35) can easily distinguished by the light yellow to white fringe of the hindwings (uniformly dark in P. cantabricus sp. n.) and the usually much more extensive white markings of the forewings. Furthermore, the males of both species differ in their antennae, which are not flattened, appear somewhat serrate and have only 22-23 segments in P. pyrenaicus . Both species also differ profoundly in their male genitalia. The ventral extension of the pseudotegumen and projections at the posterior margin of the vinculum are absent, the valva is short and arched, and the saccus is square in appearance in P. pyrenaicus . Finally, the female of P. pyrenaicus (Fig. 35) is brachypterous, while the female of P. cantabricus sp. n. is fully winged. Pharmacis bertrandi (Figs 13, 26) has very regular and extensive white forewing markings, usually also in the termen and close to the costa, markedly embedded into reddish brown areas (reduced white markings, absent in the termen and costal area in P. cantabricus sp. n.). Males have 34-35 antennal segments. Females of P. bertrandi are brachypterous. Pharmacis carna (Figs 14-16, 27, 33) has more extensive yellow to white forewing markings; the hindwing fringe is mixed yellow and dark brown (dark brown throughout in P. cantabricus sp. n.). Males have 25-27 antennal segments ( Buser et al. 2000 and our observations). Females of P. carna are fully winged and able to fly. Pharmacis anselminae (Figs 17, 25, 34) differs substantially by the light markings on its forewing, which are not distinctly edged or embedded into light brown scales, and by the presence of a dirty yellow patch subapically at the forewing costa. Males have 25-27 antennal segments ( Buser et al. 2000). Females of P. anselminae are brachypterous. Pharmacis claudiae (Fig. 18) has extensive and evenly distributed white markings on the forewings (reduced in P. cantabricus sp. n.) and the hindwing fringe is white, somewhat mixed with dark brown at the veins. Males have 26-28 antennal segments ( Buser et al. 2000). Females of P. claudiae are fully winged and able to fly.

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).

Habitat and biology.

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. 2). The ground layer is dominated by Brachypodium P. Beauv. species, with accompanying Aquilegia vulgaris L., Geranium sanguineum L., Astrantia major L. and Thalictrum minus L. Here, P. cantabricus sp. n. shares its habitat with Lopinga achine (Scopoli, 1763) ( Nymphalidae ) and Carterocephalus palaemon (Pallas, 1771) ( Hesperiidae ), two butterfly species with a very restricted distribution in Spain.

The second known locality is considerably different. It is situated above the Lagos de Covadonga at about 1375 m (Fig. 3). Here the specimens were found in open pastures interspersed with limestone outcrops, used for cattle grazing ( Estébanez, personal communication). Another mating pair was photographed by Antonio Rodriguez at low altitude (<200 m) west of the Picos de Europe on the 30th of July 2016 (Biodiversidadvirtual, 2018; as ' Pharmacis sp.'). Finally, specimens that may belong to P. cantabricus sp. n. were observed by Marcos Toribio in Robledo de Anayo (Asturia) at about 550 m altitude (Biodiversidadvirtual, 2018; as ' Pharmacis lupulina '). These specimens were attracted to lights over several days in early September 2014 (Toribio, personal communication). However, there were no specimens available to conclusively test their identity. Thus, while confirmed records suggest that P. cantabricus sp. n. is limited to the Picos de Europa and flies only during the day in July, additional unconfirmed records suggest that it may fly well into September and may also be active at night.

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. 31), although it made no attempt to copulate. The female was then placed in a cage. Her abdomen, although bulky, was not extended much below the wings in resting posture at this point. At 5.00 pm, two males were placed in the cage with the female. Shortly after this, copulation took place, which lasted less than 35 minutes (Fig. 32). During mating, the abdomen of the female became much more extended (compare Fig. 1). By approximately 6.30 pm the female had laid a copious quantity of eggs and her abdomen had shrunk considerably. On one occasion, on the 9 July 2016, on the same day when multiple males were observed flying during day time, light trapping was undertaken at the type locality. However, no specimens were attracted to the light, suggesting that the main period of activity is during the day. This notion is also supported by the observation that mating occurred in the afternoon.

Distribution.

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.