Pyralis cardinalis, a charismatic new species related to P. regalis [Denis & Schiffermüller], 1775, first recognized in Finland (Lepidoptera, Pyralidae)

The informal Pyralis regalis complex, including species of the genus Pyralis Linnaeus, 1758 (Pyralidae), with a bright white or silvery pattern on the forewing, is reviewed, supplemented by observations of the externally distinguished P. perversalis (Herrich-Schäffer, 1849), which also exhibits similarities in genitalia and DNA barcodes. We describe Pyralis cardinalis Kaila, Huemer, Mutanen, Tyllinen & Wikström, sp. nov., based on specimens ranging from Denmark and Sweden in the West to Japan and South Korea in the East. A neotype is designated for the predominantly South European P. regalis [Denis & Schiffermüller], 1775. Lectotypes are designated for Asopia kacheticalis Christoph, 1893 and Pyralis princeps Butler, 1889. Pyralis regalis ssp. sagarrai Leraut, 2005 is considered a valid species, stat. nov.


Introduction
Pyralis Linnaeus, 1758 is a diverse genus of predominantly Palearctic, South-East Asian and African moths, covering 86 described species some of which remain unrevised (Nuss et al. 2003(Nuss et al. -2019. The European fauna with only seven species recognized is comparatively poor. It includes the cosmopolitan, well known pest species meal moth P. farinalis Linnaeus, 1758, P. regalis [Denis & Schiffermüller], 1775, P. perversalis (Herrich-Schäffer, 1849), P. lienigialis (Zeller, 1843), P. kacheticalis (Christoph, 1893), and exceptional records of the tropical pests P. manihotalis (Guenée, 1854) and P. pictalis (Curtis, 1834)  . The allegedly transpalearctic P. regalis can be regarded as the flagship species of the genus in Europe due to its external appearance, which is one of the most striking in the Pyralidae. Pyralis kacheticalis (Christoph, 1893) is a species closely related to P. regalis, described from Transcaucasus (now Azerbaijan). It is reported to be distributed in Turkey, eastern Greece and Ukraine in Europe. Other than the recently described P. regalis ssp. sagarrai Leraut, 2005 no taxonomic changes have been found for more than 100 years on this continent. It was therefore surprising that during DNA barcoding campaigns (FinBOL, Lepidoptera of the Alps) the European P. regalis was found to divide into four clusters: a northern, transpalearctic group; a central-southern European group; a south-western cluster; and a somewhat heterogeneous eastern Mediterranean cluster corresponding to P. kacheticalis. The taxonomy of P. regalis and, indeed, most of the taxa involved, has been somewhat unclear as the type specimens seemingly have not been studied for a long time, if ever after their original descriptions. Nor have suspicions regarding the taxonomy been suggested (but see Leraut 2005). Particularly, the taxonomy of P. regalis from northern Europe proved to be a problem that could only be resolved in a larger geographic context.

Specimens
In this paper we only treat those species of Pyralis that are characterized by a distinctive, white or silvery white pattern on the forewing. In addition to those species treated here, we are aware of one further species from Greece: Crete, and seemingly more than one SE Asian species that match this characterization, all preserved in ZMUC. We refrain from treating them further due to the paucity of material available. Leraut (2005) also included P. perversalis into this taxon complex; this interpretation is supported by genital morphology and common pattern of DNA barcode clustering. Therefore, we illustrate this species here as well even though its forewing pattern has no clearly white pattern.
We examined ca. 660 specimens of Pyralis spp. from various European collections (see below). Data for the specimens and their repositories are given for each taxon treated. The spellings of locality and collector names follow those recorded on the original labels. We list under 'Distribution' only those country records either verified by us or by other trustworthy sources. 1 Forewing ground colour pale greyish brown; pale markings yellowish (Figs 20,21 Diagnosis. Pyralis cardinalis is unique among other focal species as having two cornuti in the male genitalia; their structure is detailed below. It differs externally from P. regalis, P. sagarrai, P. kacheticalis, P. princeps and P. joannisi by the shape of the inner, silvery fascia: its outer margin is somewhat broadened medially, while the fascia is entirely or nearly parallel-sided in the other species. It can sometimes also be outwards slightly broadened in P. regalis and may thus taken alone not always be adequate for correct identification. Unlike other species the outer area of the hindwing of P. cardinalis is deep purple, that of other species is either dark brown (P. regalis, P. princeps, P. joannisi, P. regalis may also have brown or purple tinge), or pale yellowish grey, or suffused with pale grey (P. sagarrai, P. kacheticalis), but without purple. The inner silvery fascia is narrow and reaches the dorsal margin in P. kacheticalis, unlike other species. The median area of the hindwing is paler in P. kacheticalis and P. sagarrai than in the other species, as noted in the key. In the male genitalia, P. cardinalis differs from all other related species by its vesica which is devoid of coarse spines. It shares with P. kacheticalis and P. sagarrai the presence of one long, prominent, more or less straight cornutus, but has another smaller, curved cornutus formed of fused coarse spines.
The large cornutus can sometimes also be outwards slightly broadened in P. regalis and may thus alone not always perfectly support correct identification. The vesica of P. regalis has one tiny cornutus that is often hard to decipher among spines (see, however, Remarks under that species). In the female genitalia, the inception of the ductus seminalis is broadened and somewhat sclerotized; this trait distinguishes it from all other species treated. The ductus bursae widens evenly towards the corpus bursae, it is similar only in the externally different P. perversalis, Description. External appearance (Figs 2, 4-7). Forewing length ♂: 8-9.5 mm, ♀: 8-11 mm. Labial palpus ascending, length of second segment 1.3 as long as diameter of eye; third segment short. Maxillary palpus 1/3 as long as labial palpus; head and these appendices, scape and pecten yellow. Head rough-scaled; collar yellow, intermixed with purple. Flagellum purple, male with thin cilia length of which is twice diameter of shaft; female antenna with very short ciliation. Thorax purple; abdomen varying from purple to lead-grey. Legs pale ochre. Forewing: basal third, distal quarter, and often area of distal quarter between fold and dorsal margin purple, near margin sometimes variably intermixed with dark grey scales; at 2/5 of wing length white fascia extending from costa to fold towards which it narrows, medially outwardly widened; similarly colored, elongate spot from distal 4/5 wing length from costa to 1/3 wing width towards middle; from both white areas to costal margin narrow, purple string both inwardly and outwardly narrowly bordered with dark grey. Between white areas wing colour orange brown, costa variably annulated by yellow, grey or alternating spots of both colour. Fringe brown, except in dorsal corner dark grey. Hindwing: divided by off-white stripes into three almost equally wide areas; stripes approaching each other towards anal margin. Basal and median area purple, sometimes variably intermixed with dark grey scales; distal area somewhat paler, purple. Fringe brown, except in anal corner silvery. Underside: forewing yellowish grey, area corresponding with outer white area of upper side pale ochre. Costal margin annulated by black and yellow, somewhat elongate spots; hindwing lead-grey, with evenly bent pale grey band along outer 2/3.
Male genitalia (Fig. 22). Uncus hat-shaped, weakly sclerotized medially with triangular, distally tapered as a sparsely setose, blunt-tipped lobe; uncus articulated from tegumen; tegumen narrow, broadest laterally, tapered as narrow band touching uncus. Gnathos articulated from tegumen, formed of arms being near base and medially broad, somewhat narrowed between, mesially formed as narrow, triangular lobe terminating with abruptly bent, narrow, fused hook-shaped apex, median part of gnathos twice as long as uncus. Valva 1.5-2 times as long as wide, width varying to some extent, valvae connected to each other anteriorly; dorsal margin of basal part of valva sclerotized, distal area formed of articulated square lobe; costa convex, weakly sclerotized. Transtilla laterally shortly sclerotized, median part articulated, membranous. Juxta scobinate, convex, broad tongue-shaped. Anellus connected to vinculum, nearly membranous, in lateral view bent as s-shaped, posteriorly connected to apex of phallus; easily severed during dissection from genital capsule and attached to phallus. Vinculum short, broad, v-shaped. Phallus 3 times as long as wide apart from distal 1/4 which is narrowed towards apex, and incised with ventrally directed distal opening. Vesica with two cornuti: one narrow spine, length of which is roughly half of the narrowed distal part of phallus, and another small, curved cornutus-like group formed of dense group of coarse, partly fused spines; vesica otherwise indistinctly spinose.
Female genitalia (Fig. 29). Papillae anales dorsally fused, relatively narrow, somewhat bent; densely setose. Ostium bursae situated at posterior margin of sternum 8, membranous, bowlshaped, width ¾ distance between posterior apophyses in dorsoventral projection. Posterior apophysis straight and slender; anterior apophysis somewhat longer, bent and basally stouter than apophysis posterioris. No antrum present between ostium and the distinctly sclerotized colliculum; colliculum as long as wide. Ductus seminalis incepted anterior to colliculum, at the inception ductus somewhat broadened and weakly sclerotized. Otherwise ductus bursae membranous, evenly widened towards corpus bursae without clear limit to the latter. Length of ductus bursae and corpus bursae about 7.5 times length of posterior apophysis. No signum or granulation in corpus bursae, but membrane with minute lens-shaped formations.
Molecular data. BIN: BOLD:AAF5806. The intraspecific mean divergence of the barcode region is 0.21%, the maximum divergence 0.70% (N = 21). The minimum distance to the nearest European neighbour, P. sagarrai, is 7.43%. However, an unrevised species from China is only 4.23% apart from P. cardinalis.
Biology. Life history is not known. Adults have been observed at sugar bait as well as in light traps over a long period, spanning from the end of June to the end of August. The peak of the flight period is during July, and no evidence of more than one yearly generation seems to exist.
Male genitalia (Fig. 23). Uncus hat-shaped, weakly sclerotized, medially with triangular, distally tapered as sparsely setose, blunt-tipped lobe; uncus articulated from tegumen; tegumen narrow, broadest laterally, tapered as narrow band touching uncus. Gnathos articulated from tegumen, formed of arms being near base and medially broad, somewhat narrowed between, mesially formed as narrow, triangular lobe terminating with abruptly bent, narrow, fused hook-shaped apex; median part of gnathos twice as long as uncus. Valva 1.5-2 times as long as wide, width varying to some extent, valvae connected to each other anteriorly; dorsal margin of basal part of valva sclerotized, distal area formed of articulated square lobe; costa convex, weakly sclerotized. Transtilla laterally shortly sclerotized, median part articulated, membranous. Juxta scobinate, convex, broad tongueshaped. Anellus connected to vinculum, nearly membranous, in lateral view bent as s-shaped, posteriorly connected to apex of phallus; easily severed during dissection from genital capsule and attached to phallus. Vinculum short, broad, v-shaped. Phallus 3 times as long as wide apart from distal 1/4 which is narrowed towards apex, incised with ventrally directed distal opening. Vesica usually devoid of cornuti (see Remarks); coarsely granulose.
Female genitalia (Fig. 30). Papillae anales dorsally fused, relatively narrow, somewhat bent; densely setose. Ostium bursae situated in posterior margin of sternum 8, membranous, bowlshaped, width ¾ distance of that between posterior apophyses in dorsoventral projection. Posterior apophysis straight and slender; anterior apophysis somewhat longer, bent and stouter. No antrum present between ostium and distinctly sclerotized colliculum; colliculum 1.5 times as long as wide. Ductus seminalis incepted anterior to colliculum with no swelling or sclerotization; posterior third to half of the length of ductus bursae braid-shaped, granulose; otherwise membranous, evenly widened towards corpus bursae which it joins indistinctly. Corpus bursae elongate, oval. Length of ductus bursae + corpus bursae about 6 times length of posterior apophysis. No signum in corpus bursae, but the membrane of both ductus and corpus bursae with minute lens-shaped formations.
Molecular data. BIN: BOLD:AAP5668. The intraspecific mean divergence of the barcode region is 0.44% and the maximum divergence 1.83% (N = 28). The minimum distance to the nearest neighbour, P. sagarrai, is 3.96%.
Distribution. Widely distributed in South Europe at least to Austria and Switzerland in the north. As the southwestern limit of the distribution range of P. cardinalis is not very far from the closest records of P. regalis the identity of some records from, e.g., Germany, require re-examination. Pyralis regalis and P. cardinalis are sympatric at least in Russia: Belgorod.
Remarks. The collection of Denis & Schiffermüller was deposited in the "Hof-Naturalien-Kabinett" in Vienna but destroyed by fire during the Vienna Rebellion of 1848 (Hoffmann 1952). Despite a personal search by PH we found no potential type material and conclude that the syntypes are lost or destroyed. As this taxon is part of a species complex, designation of a neotype is necessary to preserve the stability of nomenclature (ICZN 1999). The type locality is presumably in Lower Austria, where the majority of the material of Denis & Schiffermüller was collected, and as indicated by the title of the original description (Denis and Schiffermüller 1775). We therefore designate as neotype a female specimen from this province with label data mentioned above and figured in Fig. 3.
According to collecting dates ranging from late May to October, it seems likely that at least two generations occur in southern Europe.
We follow Leraut (2005)  Diagnosis. Distal area of the hindwing is distinctly pale without a purple tinge, and the hindwing in general is also somewhat paler than in P. regalis. In P. kacheticalis only the basal third of the hindwing is pale. The phallus is similarly coarsely granulose as in P. regalis and the externally quite different P. perversalis, the vesica of which is devoid of a cornutus. There is a distinctive spine-like cornutus in the vesica of P. sagarrai, P. kacheticalis and P. cardinalis, and a minute one in P. regalis; it is considerably smaller in P. sagarrai than in P. kacheticalis and P. cardinalis, and basally distinctly formed as a fusion of smaller spines, while the cornutus is entirely smooth in the other species; in P. cardinalis there is another, smaller and curved cornutus as well. The caecum of the phallus is wider, and joined with an obtuse angle to the distal part of phallus, which is a diagnostic difference with P. regalis. This difference, however, is easily hidden depending on the position of the phallus. The female of P. sagarrai seems to be readily differentiated from P. regalis as having a long and evenly widening ductus bursae that joins the corpus bursae without limit. Together with the diagnostic external characters, the differences in the female genitalia and the significant difference in the barcodes we consider this taxon to merit status of a valid species even though the female genital differences are not firmly established. The separation of this species from P. princeps and P. joannisi is explained in the key.
Molecular data. BIN: BOLD:ADS1090. The intraspecific mean divergence of the barcode region is 0.18% and the maximum divergence 0.31% (N = 3). The minimum distance to the nearest neighbour, P. regalis, is 3.96%.

Diagnosis.
Pyralis kacheticalis differs from other related species apart from P. sagarrai by the pale hindwings, and the shape of the white fascia on the forewing being very narrow, straight and extended to the costal margin. The male genitalia are close to those of P. sagarrai from which they are distinguished by the shape of the cornutus: it is entirely smooth in P. kacheticalis, basally wide and formed as fusion of coarse spines in P. sagarrai. The female genitalia of these species seem to be identifiable with some reservations (see key), but may be indistinguishable from P. regalis. The separation of this species from P. princeps and P. joannisi is explained in the key. Molecular data. BIN: BOLD:ABA8506, BOLD:ABA9291. Genetically variable species clustering into two BINs and a third distinct cluster presently lacking the BIN assignment. The intraspecific mean divergence of the barcode region is 2.18% and the maximum divergence 3.67% (N = 9). The minimum distance to the nearest neighbour, P. sagarrai, is 5.5%.

Remarks.
In the original description of P. kacheticalis it is indicated that the number of specimens on which the species is described is more than one, so no holotype exists. We herewith designate a lectotype for the taxon (Fig. 34) in order to fix the identity of the species and conserve nomenclatural stability. Pyralis imperialis Caradja was originally described based on two specimens. Although Popescu-Gorj (1991) designated a lectotype for this taxon, the specimen (Fig. 35) does not differ in any way from P. kacheticalis. We therefore follow Leraut (2005) in regarding P. imperialis with P. kacheticalis as synonyms.
Specimens from Cyprus, Lebanon and Ukraine/Greece form three distinct barcode clusters. Externally, there is some variation between specimens, but the variation observed in, e.g., the shape of median area of hindwing, seems not to correlate with barcode clusters. Also, the genitalia are identical among all specimens that we have examined. We refrain from making taxonomic conclusions due to the paucity of material at our disposal, and with evidence from barcodes alone.
An endemic population, externally closest to P. kacheticalis that occurs in Crete, seems to form its own entity, possibly deserving species status. This note is based on an unpublished barcode that deviates strongly from others, as well as external features (genitalia not studied). Due to the paucity of material we exclude this taxon from the present contribution. Diagnosis. The separation of this species from other species is explained in the key. In the genitalia, the dense bush of long spines at the posterior end of vesica is characteristic, and the genital characters readily separate it from the externally similar but smaller P. joannisi; see the key for further details.

Pyralis princeps
Molecular data. BIN: BOLD:ABA8505. The intraspecific mean and maximum divergence of the barcode region is 0% (N = 2). The minimum distance to the nearest neighbour, P. kacheticalis, is 8.26%.
Distribution. India, Nepal. Remarks. Pyralis princeps was described from four specimens collected in India (Himachal Pradesh, Dharamshala [Dharmsala]) and two from "Yezo" [region from Northern Japan to Kam- chatka] (Butler, 1889). However, the latter two are misidentifications of P. cardinalis, this interpretation also supported by our examination of some Japanese and South Korean specimens. Following ICZN (1999), a lectotype from Dharamshala [Dharmsala], already labelled as such by M. Shaffer in NHMUK, is here designated in order to fix the identity of the species and conserve nomenclatural stability.We have examined photographs of the specimen and its labels which confirm  our interpretation. Outside Europe, the name P. princeps Butler has been somewhat inconsistently used for the East Palearctic populations. E.g. Shibuya (1928), in his treatment of Japanese species of Pyralis and related genera, only recognized P. regalis of the relevant taxa to occur there. Inoue et al. (1982), plate 46: Figs 4, 5 illustrated Japanese specimens that seem identical to P. cardinalis and whose occurrence in Japan is verified in the present study. Neither of these authors mention other species, but they consider P. princeps a synonym of P. regalis.
Diagnosis. The separation of this species from others is explained in the key. Molecular data. Unavailable. Distribution. Vietnam. Remarks. We have seen in ZMUC single specimens of further SE Asian taxa close to species included in this paper, which appear to be currently unnamed. We refrain from treating them more closely due to the paucity of material available.