Taxonomic changes in the genus Dyscia ( Lepidoptera : Geometridae : Ennominae )

This paper presents a revision of the genus Dyscia Hübner, [1825] (Lepidoptera, Geometridae: Ennominae). Examination of types and additional material for most described taxa has resulted in several new synonyms: Dyscia ilivolans Wehrli, 1953 syn. n. and Dyscia duanjiao Yang, 1978 syn. n. are new synonyms of Dyscia fagaria (Thunberg, 1784); Dyscia karsholti Wiltshire, 1991 syn. n. is a new synonym of Dyscia galactaria Turati, 1934; Dyscia dagestana Wehrli, 1934 syn. n. is a new synonym of Dyscia malatyana Wehrli, 1934, and Dyscia rjabovi Wardikjan, 1957 syn. n. is a new synonym of Dyscia negrama Wehrli, 1953. One taxon, formerly treated as a species is reassigned to subspecific level, Dyscia conspersaria ssp. sultanica Wehrli, 1936 stat. rev. Dyscia innocentaria sicanaria (Oberthür, 1923) stat. n. and Dyscia malatyana senecai Wiltshire, 1990 stat. n. are downgraded to subspecies-level. On subspecies-level, Dyscia distinctaria perdistincta Herbulot, 1957 syn. n. is a new synonym of Dyscia distinctaria (Bang-Haas, 1910), Dyscia fagaria alvarensis Wahlgren, 1913 syn. n., Dyscia fagaria albescens Lempke, 1952 syn. n., Dyscia fagaria fusca Lempke, 1952 syn. n., and Dyscia fagaria postdelineata Lempke, 1952 syn. n. are new synonyms of Dyscia fagaria. Dyscia fagaria psoricaria (Eversmann, 1848) syn. n. is a new synonym of Dyscia fagaria favillacearia (Hübner, [1799]), Dyscia holli duponti Thierry-Mieg, 1910 syn. n. is a new synonym of Dyscia holli (Oberthür, 1910), Dyscia malatyana nachadira Brandt, 1941 syn. n. and Dyscia malatyana theodoraria Warnecke, 1941 syn. n. are new synonyms of Dyscia malatyana albersaria War­ necke, 1940 stat. n., and Dyscia penulataria naevata Wehrli, 1953 syn. n. is a new synonym of Dyscia penulataria (Hübner, [1819]). Lectotypes for 22 taxa and neotypes for three taxa Geometra conspersaria [Denis & Schiffermüller], 1775, Geometra fav­ illacearia Hübner, [1799], and Geometra emucidaria Hübner, [1813] are designated. One species is excluded from Dyscia, Thysanopyga serena (Dognin, 1906) comb. n. From a total of72 previously described taxa, 19 are recognized as species.


INTRODUCTION
The palearctic Gnophini genus Dyscia Hübner, [1825] (Lepidoptera: Geometridae: Ennominae) contains, by pre sent knowledge, 19 species, which occur on steppe land and along the margins of deserts of North-Africa, Arabia, and Asia, and at the south of Europe.Only one species, Dyscia fagaria (Thunberg, 1784), occurs further north to Northwest-Europe as a stenoecic species of open and sandy heaths.The medium-sized moths are nocturnal, and usually of inconspicuous colour.Therefore, numerous species were discovered and described only in the 20th century in the course of the common application of the genital-examination.
The genus Dyscia was established by Hübner ([1825]: 314) for Dyscia conspersaria ( [Denis & Schiffermüller], 1775) and Dyscia emucidaria (Hübner, [1813]).For the first time in their catalogues Staudinger (1871) and Staud inger & Rebel (1901) arranged the species of the palaearctic fauna which are part of Scodiona Boisduval, 1840 (= Dyscia;Hulst, 1896).The first comprehensive representation of the group is found in Prout (1912Prout ( -1915)), who treated all known taxa in the context of his work on the palaearctic geometrid moths.Culot (1919Culot ( -1920) ) gave a summary of the European Dyscia species and, for the first time, made notes on the life his tory of this group.Oberthür (1923) finally dealt with this group.
Mainly taxonomic oriented works by Warnecke (1940Warnecke ( -1941Warnecke ( , 1941) ) and Albers & Warnecke (1940) fol lowed, in which, for the first time, male genital organs as well as a distribution overview were shown for the spe cies of the so called "Dyscia conspersaria F.[sic!]--group".Wehrli (1950) grouped the Dyscia species mainly with features of the male genitalia into altogether eight subgenera.Later he assumed this grouping in his substantial contribution on palaearctic Geometridae (Wehrli, 1953).The only zoogeographic studies which followed were a study by Wiltshire (1990) which contains a tabular survey of the Dyscia species of North African countries, and a study by Erlacher & Trusch (1999) which deals with the phylogeny and distibution of the "Dysciaconspersaria-group".Herbulot (1981) described a new Dyscia species from Morocco.Wiltshire (1991Wiltshire ( , 1994) ) worked on North African and Arabian Dyscia and described the last discovered species in 1986 from Saudi Arabia.
THE GENUS DYSCIA HÜBNER, [1825] The moths of the genus Dyscia can externally be recog nised by their characteristic shape.The individuals are medium-sized and coloured from pale grey to beige, with the exception of the reddish Dyscia nobilaria (Bang-Haas, 1906).Unlike other geometrid moths, the outer margin of the forewings is slightly concave.Another external character is the lack of the proboscis (cf.already Hulst, 1896: 366).In the male genitalia, the width to length ratio of the valvae distinguishes two types of genitalia, the high and the wide type (cf.Trusch & Erlacher, 2001).At the basis of the dorsal margins of the valvae, costal arms emerge medially.A further diagnostic feature is the loss of the uncus.The absence of an uncus occurs also in the genus Aspitates Treitschke, 1825 andSiona Duponchel, 1829.Therefore, this distinguishing mark is not an exclusive apomorphy to the genus Dyscia.
The female genitalia are small and often lacking differ entiating features.Its components are for the most part weakly sclerotized.There are two types of corpus bursae, one is weakly sclerotized and the other solid and sculp tured, i.e. with very fine tubercles on the inner side of the corpus bursae.The larvae of all species posses a dorsal process on their 9th abdominal segment, except Dyscia nobilaria.In Dyscia penulataria (Hübner, [1819]), this feature is polymorphic, i.e. there are specimens with and without this structure.Finally, in Dyscia lentiscaria (Donzel, 1837) the process is doubled, as an alteration to this character.
Traditionally, the genus Dyscia is placed within the tribe Gnophini (Pierce, 1914, as Gnophinae), which is part of the subfamily Ennominae.

Description of the type species Dyscia conspersaria ([Denis & Schiffermüller], 1775)
Male.Forewing length 16-21 mm; basal colour whiteyellowish to white-brownish, suffused with dark scales.Melanistic individuals occur, which are sometimes com pletely brown and lack all markings.Discal spot orbicular or oval, filled with a paler colour, occasionally absent; basal line and postmedian line vary considerably between specimens, ranging from some vein markings to a con tinuous wavy line; median shadow occurs particularly in specimens from Asia Minor, but occasionally also in the remainder of the species range; suffused spots at the hind margin frequent, in most cases only slightly darker than the basal colour.Hindwing has the same basal colour as forewing; postmedian line continuous on hindwing in the way described above; small central spot point to commashaped, unfilled.Body in basal colour, thorax with paler scales in melanistic individuals.Femora, tibiae, tarsi in basal colour.Underside of wings whitish, partly shiny, the 2/3 towards the anterior margin of forewing brownish.
Central spots at the ventral side of forewing and hindwing and the outer line marking clear.Antennae bipectinate.
Female.Antennae filiform, otherwise as male.Male genitalia.Of the high type (cf.Trusch & Erlacher, 2001).Valvae widened ventrally, especially specimens from Turkey have proximally somewhat lobusshaped outgrowths.Complex of valvae rotated towards the median line, appearing very broad, but distally not widened.Dorsally, the costal arm is broadly attached at the base of the valvae.It is horn-shaped and bent inwards, with one terminal short tooth.Gnathos almost absent, only a very delicate structure.Transtilla proximally con nected, the junction as a longitudinal fold, in preparation directed vertically.Saccus regulary rounded.Aedoeagus slender, s-shaped; with a bulge at ductus ejaculatorius; caudal end with two short teeth, otherwise without cornuti.
Female genitalia.Corpus bursae solid and sculptured, i.e. with very fine tubercles at the inner side of the corpus bursae.Ductus bursae tubular and short, about 1/4 length of corpus bursae and of constant diameter (approx.1/2 of its length).Sterigma of the lamella postvaginalis narrow and high; laterally of this structure a bigger longitudinal fold on both sides, that leads towards the sterigma.On both sides of the ostium bursae are another five smaller longitudinal folds.

Dyscia conspersaria conspersaria ([Denis & Schiffermüller], 1775)
Geometra conspersaria [Denis & Schiffermüller] Horn et al., 1990).The characters differentiating Dyscia conspersaria and the other Dyscia species are described in Trusch & Erlacher (2001), and the Neotype is figured there, pl. 1, Figs D,E and pl. 4,Fig. C. The relationships between the taxa con spersaria and sultanica are explained in the following.Hübner, 1790: 38, pl Remarks.Dyscia sultanica Wehrli, 1936 is treated here as a subspecies of Dyscia conspersaria.Already Albers & Warnecke (1940:122) had stated that in Asia Minor Dyscia conspersaria occurs, but in the form sultana[sic!].Wehrli (1953: 659) gave a number of charac ters which led him to regard sultanica as a species, all of which are, however, gradual and not clearly definable.He indicated a difference in the shape of the costal arm of the valva as well as a wider gnathos and a differently-shaped saccus as characters distinguishing sultanica from con spersaria.Wehrli (1953) subsequently gave the bifurca tion at the tip of the costal arm as the only qualitative dif ference in the genitalia, a feature which is not clear from an examination of his material nor of material examined by the present authors.

Ph[alaena] Noct[ua] cuniculina
This facts are particularly true of animals from Asia Minor which show a wide range of variation (cf.Trusch & Erlacher, 2001).Thus, in Anatolia there occur not only whitish-brown moths on which the description of sul-tanica is based, but also brightly coloured specimens which resemble the European ones and, rarely, com pletely melanic individuals.Separation of this group of forms from European representatives on the basis of genital differences does not appear to be justified, although in males from Asia Minor the ventral side of the valva shows a small lateral step which has not so far been seen in European material.This structure is very variable but can be stated, according to current knowledge, as a character separating sultanica as a subspecies, unless fresh data calls for a new interpretation, together with the more contrasting pattern of the forewings which also fre quently have a median shade.The absence of findings in the Balkans indicates a strong geographical isolation between the populations of Anatolia and Europe.Less is known, however, about the degree of separation of those from the Transcaucasus and Lower Volga.
Remarks.According to Article 75 and 76 ICZN, a Neotype is designated with the purpose of clarifying the taxonomic status of Dyscia fagaria favillacearia (Hübner, [1799]), particularly regarding the status of the taxon psoricaria Eversmann, 1848.It is common knowledge that the Lepidoptera-types of Hübner were destroyed in Vienna in 1848 (e.g.Horn et al., 1990) Horn et al., 1990).A melanic male, infrasub specific, not available.Wehrli, 1953: 662 Remarks.Of Dyscia ilivolans Wehrli, 1953, the female lecto-and the male paralectotype were examined (see above).Further specimens of the taxon have not been identified up to now but may have been mixed up with Dyscia malatyana (cf.Viidalepp, 1979 and1988: 142).The two type specimens have been determined as Dyscia fagaria.The differential features given by Wehrli (1953: 662), "gnathos as in malatyana but somewhat shorter and more slender", "saccus as is malatyana" and "costal arm of valva rather 'intermediate'" particularly are not correct.Wehrli indicates that "the structure of the genitalia is intermediate between Dyscia malatyana and Dyscia fagaria", which cannot be confirmed here.

Dyscia ilivolans
There is a wide range of variation in colouring and pat tern in Dyscia fagaria.All external characters of the taxon ilivolans given by Wehrli (1953) are described from characteristically aged collection-specimens of Dyscia fagaria, the only material, which was available to Wehrli at the time of his description.Dyscia ilivolans Wehrli, 1953 is therefore synonymized with Dyscia fagaria (Thunberg, 1784).The consubspecific assignment of the taxon ilivolans Wehrli, 1953 to Dyscia fagaria fav illacearia Hübner, [1799] is based on the outer phenotype of the specimens, especially the grey-brownish basal colour of the wings and the lack of the elements rich in contrast typical in this subspecies.Yang, 1978: 392, pl. 20, Fig. 13.Syn.n.
Remarks.Dyscia duanjiao Yang, 1978 was described from the north-east of China by Yang (1978: 392).Com paring Dyscia duanjiao and Dyscia fagaria, the only one diagnostic character mentioned by Yang (1978) is the shorter antennae of the taxon described by him.The name, duanjiao, is derived from a Chinese word, meaning "short horn", referring in the case to "short antenna".So, the antenna in Dyscia fagaria should be about 2/3 of the length of the forewing costa.In contrast, the antenna in Dyscia duanjiao measures only half this length.The material examined here shows, however, that the length of the antenna in Dyscia fagaria generally is about half the length of the forewing costa (cf.Trusch & Erlacher, 2001).Furthermore, a smaller discal spot and a different shape of the postmedian line, insignificant differences, are given by Yang (1978) for his material, two males.These supposed differences are within the range of varia tion of Dyscia fagaria, which varies greatly across its huge distribution area from Western Europe to the Amur.Even the shape of Yang's illustration (1978: pl. 20, Fig. 13) appear identical with Dyscia fagaria, especially in specimens from Mongolia and China examined here (cf.Trusch & Erlacher, 2001: 81-82).Therefore Dyscia duanjiao Yang, 1978 must be regarded as a junior synonym of Dyscia fagaria (Thunberg, 1784).The con subspecific assignment of the taxon duanjiao Yang, 1978 to Dyscia fagaria favillacearia Hübner, [1799] is based on the outer phenotype of the specimens, see above.Remarks.According to Art. 75 ICZN, a neotype for the taxon Geometra emucidaria Hübner, [1813] is desig nated, in order to clarify the taxonomic status of the above cited Fig. 425 of Jacob Hübner.His Lepidopteratypes were destroyed in Vienna in 1848 (e.g.Horn et al., 1990).After a long period, during which the taxon emuci daria was considered to be a good species, a controversy concerning its status began in 1950.Dyscia emucidaria was then synonymised with Dyscia fagaria (Thunberg, 1784) by Lucas (1950: 94), but at that time he did not designate a neotype.Warnecke (1959: 4) then interpreted the questionable illustration of Hübner as a synonym of Dyscia penulataria (Hübner, [1819]).As a result, the common name Dyscia penulataria would become a junior synonym of Dyscia emucidaria.Therefore Warnecke (1959) suggested the conservation of the name penula taria, but he did not invoke the plenary power of the Commission according to article 79 ICZN to suppress the older name emucidaria.Since the type material no longer exists, a further discussion about its identity lacks any sci entific basis.Therefore, according to Lucas (1950), Dyscia emucidaria is seen here as representing specimens of Dyscia fagaria from Central and Southwest France, especially since an opinion regarding the identity of Dyscia fagaria and Dyscia emucidaria has already been given in the literature (Hackray et al., 1984).Furthermore, the pointed forewing shape of the individual of Fig. 425 strongly supports the opinion that there has been illustrated a specimen of Dyscia fagaria, and not of Dyscia penulataria, which has more rounded tips to the forewings.The characters which differentiate Dyscia fagaria and the other Dyscia species are described in Trusch & Erlacher (2001), and the valva of the Neotype of Geometra emucidaria Hübner, [1813]  Remarks.Described from the Libyan Cyrenaika, the female holotype of Dyscia galactaria Turati, 1934 was the only one known specimen of this species until 1998.From the western neighbouring country Tunisia, Dyscia karsholti, of which up to 1997 only two males were known, was described in 1991 by Wiltshire.In 1997, three further males, and in 1998 at last also a female, were captured as a result of two expeditions to Tunisia, done in the context of this work.The comparison of the genital-armature of the female with the holotype of Dyscia galactaria showed, that Dyscia karsholti Wiltshire, 1991 is a junior synonym of Dyscia galactaria Turati, 1934.Altogether, the specimens referred to here are the only ones known to date (cf.Trusch & Erlacher, 2001).Additional material examined.3 k k , 2 2 2 , see appendix in Trusch & Erlacher (2001).

Dyscia innocentaria sicanaria
Remarks.The synonymy of Dyscia sicanaria (Ober thür, 1923) and Dyscia innocentaria (Christoph, 1885) was made here, had already been suggested by Warnecke (1941: 209, 232).Later, however, Wehrli (1953: 665) gave the opinion that the two forms were specifically dif ferent.His reasons, derived exclusively from external fea tures and genitalia of the males examined by him, were 1.A different wing: body proportion and a "marking similar to Perconia".-More than 300 specimens exam ined during the present study show a wide range of variation in pattern and shape of the wings, and indicate that these are not diagnostic characters (cf.Trusch & Erlacher, 2001).

The ventral process of the valva in Dyscia innocentaria
(the eastern form is meant), is shorter, wider, and with more spines on the distal area.-These features appear, however, also in animals examined by us from Italy, the original area of Dyscia sicanaria and the central Asian animals cannot be separated on these characters.3.In the comparison of Dyscia innocentaria and Dyscia sicanaria, the notch of valva is more shallow, roundedrectangular, not acute-angled.Gnathos shorter and more slender.Aedeagus distally broader, with a more slender distal clasp.Saccus "less deeply concave", its margin nearly straight, caudal with a much weaker lead.
-These differences have to be put down to the indi vidual differences in the slides of the two males, exam ined by Wehrli (artefacts by pressure, cf.slide No. 7375 and 7376 Wehrli in coll.ZFMK).
Further the examination of a number of animals in both sexes from Turkmenistan, Turkey and Italy has also failed to yield specific differences between these animals.None of the alleged differentiating features for the taxon inno centaria given by Wehrli (1953) are diagnostic.Conse quently Dyscia innocentaria (Christoph, 1885) is a senior synonym of Dyscia sicanaria (Oberthür, 1923).Because of its more bright basal colour and some differences in the marking of the wings, Dyscia innocentaria sicanaria (Oberthür, 1923) can be treated as a subspecies of Dyscia innocentaria, its range is South Europe.Dannehl, 1933: VII, 146.

Dyscia lentiscaria lentiscaria (Donzel, 1837)
Crocallis lentiscaria Donzel, 1837: 13, pl.Remarks.In the framework of Wehrli's treatment of the Dyscia-group within the supplementary-volume of "Die GroBschmetterlinge der Erde", it had become clear that Dyscia plebejaria (Oberthür, 1910) and Dyscia ple bejaria dagestana Wehrli, 1934, originally described without examination of their genital morphology, are not conspecific.However, after dissection of one male, Wehrli (1953: 661;slide No. 7372 Wehrli in coll.ZFMK) elevated dagestana to specific rank, and gave numerous conjectural differences from Dyscia malatyana: 1. male genitalia of dagestana considerably bigger, although the moths smaller, 2. different form of valva, basal half of valva slender, distal part of valva widened and approx.1/3 larger as in Dyscia malatyana, 3. tegumen of dagestana "distinctly wider", 4. process of valva more strong, 5. ter minal thorn longer, 6. gnathos more slender, and 7. saccus anteriorly more flattened as in Dyscia malatyana.However, they are inappropriate or already decline within the variability of the material examined by Wehrli himself respectively, as shown by his slides (cf.Dyscia malat yana slide No. 7371, 7393; Dyscia malatyana nachadira slide No. 7410; Dyscia malatyana theodoraria slide No. 7384, and Dyscia malatyana dagestana slide No. 7372, all Wehrli in coll.ZFMK).The characters mentioned above are not clearly definable.With them, no group for mation can be carried out in connection with broader fea tures of the animals or their geographical distribution.The high variability of the colouring and pattern of body and wings in Dyscia malatyana does not allow a delimita tion on the basis of external characteristics (vs.Wehrli, 1953: 661).As shown from the data of the type material of Dyscia dagestana, these are specimens of the second generation of Dyscia malatyana.

Dyscia malatyana nachadira
Remarks.As a result of the material examined here, Dyscia senecai Wiltshire, 1990 should no longer be regarded as a good species.In his description, Wiltshire (1990: 355) gave only slight differences in the structure of the aedeagus as differentiating characters, in separating his species from the extremely variable Dyscia malatyana.His arguments could not be supported, either by examination of the type material or of the further material studied here (cf.Trusch & Erlacher, 2001).The sand-yellow coloured animals from Libya, which were available to Wiltshire (1990) for his description of Dyscia senecai, do not show any differences from animals of the "Klapperich-gain" from Jordan examined by Hausmann (1991: 112), for example.These have been determined by Hausmann (1991) as Dyscia malatyana, in agreement with the determination carried out here of the same mate rial.Also, new material collected here from Tunisia, having the phenotype of Wiltshire's senecai, had to be determined as Dyscia malatyana.If the mentioned sandyellow colouring of the specimens from North Africa and the Middle East is seen as an evidence for a good subspe cies, senecai can be treated as a subspecies of Dyscia malatyana of this region.

subspecies of Scodiona hispanaria Milliere, 1866 and elevated to species rank by
7.

.
Dyscia fagaria favillacearia is the geographic race from Central Europe to East Asia.