Taxonomic revision and zoogeographical patterns of the species of Gnopharmia Staudinger, 1892 (Geometridae, Ennominae) Author Sh, Hossein Rajaei Author Stüning, Dieter Author Trusch, Robert text Zootaxa 2012 3360 1 52 journal article 10.5281/zenodo.214977 77ee786b-c3de-46c0-a2e8-9247decffea5 1175-5326 214977 G. colchidaria objectaria Staudinger stat. rev. ( Figs. 18–24 & 35 ; Map 1) Gnopharmia colchidaria var. objectaria Staudinger, 1892 : 181 . Syntypes : 1 3, 1 Ƥ, coll. MNHU (examined). Type locality: (3) Akhal-Tekke region, Ashkhabad ( Turkmenistan ), (Ƥ) Mardin ( Turkey ) (= irakensis Wehrli ). Lectotype 3 designated here. Gnopharmia colchidaria objectaria : Staudinger, 1901 : 344 ; Prout, 1915: 384; Parsons et al., 1999 : 406 . Gnopharmia objectaria : Wehrli, 1953: 566 , pl. 47e; Viidalepp, 1988 : 137 . Gnopharmia objectaria objectaria : Parsons et al., 1999 : 406 . Gnopharmia colchidaria var. degeneraria Staudinger, 1892 : 181 . syn. nov. Syntypes : 4 3, 1 Ƥ, colls MNHU, ZFMK (examined). Type locality: Ashkhabad ( Turkmenistan ). Lectotype 3 designated here. Gnopharmia degeneraria : Viidalepp, 1988 : 136 . Gnopharmia objectaria degeneraria : Wehrli, 1953: 567 , pl. 47e. Gnopharmia colchidaria degeneraria : Parsons et al., 1999 : 406 . Gnopharmia inermis Wiltshire, 1967 :156 , pl. 2, figs 20, 21; pl. 13, fig. 54; pl.14, fig. 55. syn. nov. Holotype 3, paratypes 3 3, 1 Ƥ, coll. SMNK (examined). Type locality: Herat, Afghanistan . Gnopharmia inermis : Parsons et al., 1999 : 406 . Gnopharmia inermis vartianae Wiltshire, 1970 : 389 , pl. 1, fig. 4. syn. nov. Holotype 3, Paratype Ƥ, coll. NHMW (examined). Type locality: Jussufabad, near Mashhad, NE Iran . Gnopharmia inermis vartianae : Parsons et al., 1999 : 406 . Gnopharmia eberti Wiltshire, 1967 : 158 , pl. 2, fig. 17; pl. 14, fig. 59. syn. nov. Holotype 3, coll. SMNK (examined). Type locality: Arghandab river, 30 km N of Kandahar (SW. Afghanistan ). Gnopharmia eberti : Parsons et al., 1999 : 406 . Gnopharmia maculifera kasyi Wiltshire, 1970 : 389 , pl. 1, fig. 3. syn. nov. Holotype 3, NHMW (examined). Type locality: Jussufabad, near Mashhad, NE Iran . Gnopharmia maculifera kasyi : Parsons et al., 1999 : 406 . Type material examined. G. colchidaria var. objectaria Staudinger, 1892 , lectotype 3 (hereby designated in order to stabilize nomenclature), [ Turkmenistan ] ‘Askhabad, “Eyldt” [name of collector, hand-written, not correctly readable], ‘Origin’, ‘764’, ‘GlobInG specimen ID: 0023’, ‘[slide no] GU: 679/2008 R. Trusch’, ‘ Lectotype G. colchidaria objectaria Staudinger, 1892 | design. here’; in coll. Staudinger in MNHU ; paralectotype Ƥ, ‚Mardin, 88. “Sint.” [1888; name of collector, not correctly readable]’, ‚Origin.’, ‘[slide no] GU: 680/2008 R. Trusch’, ‘ paralectotype G. colchidaria objectaria Staudinger, 1892 | det. H. R. 2011’; in coll. Staudinger in MNHU . G. colchidaria var. degeneraria Staudinger, 1892 , lectotype 3 (hereby designated in order to stabilize nomenclature), [ Turkmenistan ] ‘Askhabad | 89 “Eyl.” ’, ‘Origin.’, ‘GlobInG specimen ID: 0022’, ‘gen. prep. 409/ 2008 H. R.’, ‘ Lectotype G. colchidaria var. degeneraria Staudinger, 1892 | design. here’; in coll. Staudinger in MNHU ; paralectotype Ƥ, [ Turkmenistan , Askhabad], ‘Origin’, ‘gen. prep. 410/2008 H. R.’, ‘ G. c o l c h i d a r i a objectaria Staudinger, 1892 | det. H. R., 2009’; in coll. Staudinger in MNHU . Further typical and topotypical material: paralectotype 3, [ Turkmenistan ] ‘ colchidaria v. degeneraria Stgr. , 3, 1896, Askhabad’, ‘ Gnopharmia degeneraria Stgr. , abgebildet Seitz IV. Suppl. fig.’ [pl. 47e, right specimen of the two degeneraria ], Wehrli gen. prep. no. 5139, G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; Ƥ, [ Turkmenistan ] ‘ Colchidaria v. degeneraria Stgr., Ƥ , Askhabad’, ‘ Gnopharmia degeneraria Stgr. , abgebildet Seitz IV. Suppl. fig.’ [pl. 47e, left specimen of the two degeneraria , erroneously with male antennae], G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; paralectotype 3, [ Turkmenistan ] ‘ colchidaria v. degeneraria Stgr. , 3, Askhabad’, ‘ cocandaria v. degeneraria ’, ‘ paralectotype G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’ [abdomen missing]; 3, Transkasp[ien] Askhabad [ Turkmenistan ], ‘655’, Wehrli gen. prep. no. 7219, „ G. objectaria Stgr , von ihm erworben“[i.e. purchased from Staudinger]; Gnopharmia objectaria Stgr. , abgebildet Seitz IV. Suppl. fig. [pl. 47e, right specimen of the two objectaria , erroneously labelled as female]; Ƥ, ‘Askhabad, v. cocandaria Stgr ,’ [ Turkmenistan ], Gnopharmia objectaria Stgr. , abgebildet Seitz IV. Suppl. fig. [pl. 47e, left specimen of the two objectaria , erroneously with male antennae and labelled as male]; gen. prep. 883/2009 H. R.; 3, ‘Tekke’ [ Turkmenistan ], ‘652’, „ G. objectaria Stgr. , von ihm erworben“, paralectotype G. colchidaria objectaria Staudinger, 1892 | det H. R., 2009’, Ƥ ‘Tekke’ [ Turkmenistan ], ‘653’, „ G. objectaria Stgr. , von ihm erworben“, ‘ paralectotype G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; all in ZFMK . Gnopharmia inermis , Holotype 3 (examined), ‘ Afghanistan | Herat, 970 m | 15.– 25.4.1956 | H. G. Amsel leg.’, ‘ Gnopharmia | inermis Wilts. | Holotype 3’, ‘gen. prep. 596/2009 H. R.’, in coll SMNK . Paratypes (examined): 1 Ƥ, ‘ Afghanistan | Herat, 970 m | 15.4.1956 | H. G. Amsel leg.’, ‘ Gnopharmia | inermis Wilts. | Allotype Ƥ’, ‘ G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; 2 3 ‘ Afghanistan | Herat, 970 m | 15.4.1956 | H. G. Amsel leg.’, ‘preparation MW. 153’, ‘ Gnopharmia | inermis Wilts. | Paratype 3’, ‘ G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; 1 3: ‘ Afghanistan | Herat, 970 m | 5.5.1956 | H. G. Amsel leg.’, ‘Preparation E.P. 1136’, ‘ Gnopharmia | inermis Wilts. | Paratype 3’, ‘ G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; both in SMNK . Gnopharmia inermis vartianae , Holotype 3, ‘ 23.VI.1963 | NO [NE]- Iran , W v. | Jussufabad | [leg.] Kasy & Vartian’, ‘Preparation WW. 78. 3’, ‘[a piece of blue paper]’, ‘Nat. Hist. Mus. | Wien | Gen. Praep. | MV 17306’, ‘ Gnopharmia | inermis Wilts. | ssp. vartianae Wilts. | holotype 3’, ‘ G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; Paratype Ƥ (examined), ‘ 21.VI.1963 | NO [NE]- Iran , | W v. Meshed [Mashhad] | [leg.] Kasy & Vartian’, ‘Preparation WW. 78. Ƥ’, ‘[a piece of blue paper]’, ‘Nat. Hist. Mus. | Wien | Gen. Praep. | MV 17306’, ‘ Gnopharmia | inermis Wilts. | ssp. vartianae Wilts. | allotype Ƥ’, ‘ G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; both in NHMW . Gnopharmia eberti , Holotype 3 ‘SW. Afghanistan | Fluß Arghandab 30 | km nördl. Kandahar | 1150 m | 23.V.1957 | leg. G. Ebert’, ‘Preparation WM. 110 [E. P. Wiltshire]’, ‘ Gnopharmia | eberti Wilts. 3 | Holotype’, ‘ G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; in SMNK . Gnopharmia maculifera kasyi , Holotype 3, ‘ 23.VI.1963 | NO [NE]- Iran , W v. | Jussufabad | [leg.] Kasy & Vartian’, ‘Preparation WW. 148’, ‘Nat. Hist. Mus. | Wien | Gen. Praep. | MV 17305’, ‘ Holotypus | Gnopharmia | maculifera Stgr. | ssp. kasyi Wilts.’, ‘ G. colchidaria objectaria Staudinger, 1892 | det. H. R., 2009’; in NHMW . Additional material studied: 93 3, 15 Ƥ, see appendix. Description. Wings and body ( Figs 18–24 ). Frons strongly extended, with a separate protrusion set into a distal depression. Genae distinctly pointed antero-ventrally. Free apical flagellomeres in male antennae 12. Wingspan 24–28 mm . Tibial spines long and acute. Wings on upperside light grey, sand-coloured, dusted with brown scales, transverse lines replaced by a few rather indistinct dark brown spots, most conspicuous those along costa. Postmedial row of dots more numerous, incurved in the middle; a broad, dark grey marginal band present, apical patch as ground colour, rather indistinct. Discal dots blackish brown, fairly visible on all wings. Under side with basal two thirds almost white, scattered with a few black scales, marginal band well contrasting, dark grey, including a white apical patch, discal dots large, almost black ( colchidaria objectaria , fig. 18-b). Variation. Even more abundant than the nominate form is a more or less uniform, dark grey infrasubspecific form with reduced pattern elements, described as var. degeneraria (of colchidaria ) by Staudinger (1892: 183) (fig. 19) in which also the under side is strongly suffused with brown or grey scales, the marginal band being only slightly darker, not well contrasting. In this form, the apical patches are almost absent, the discal dots are weak. Transitional forms occur, also rarely specimens with a reddish-brown pattern on upperside. G. i n e r m i s Wiltshire is such a brownish form (fig. 20), while G. inermis vartianae (figs 21 & 22) is transitional between objectaria and degenararia . G. eberti (fig. 23) and G. maculifera kasyi (fig. 24) are, on the other hand, rather typical objectaria . Male genitalia and pre-genital abdomen ( Fig. 35 ). Distal projection of sacculus slightly shorter than proximal one. Aedeagus long (1.3–1.5 mm), with a long coecum penis and a small ventral fin, situated almost in the middle of the shaft. Distal subapical spines present, very small, tooth-like, single or arranged in a row of 2–3, rarely more (fig. 35-b). Proximal subapical spines variable, rarely completely absent, mostly a single small spine only, but sometimes larger, rarely even two spines. Distal and proximal subapical spines placed at a rather large distance from each other. Vesica with a small, multiple cornutus. Octavals short (0.25–0.4 mm), with a wide opening between them (fig. 35-d). Diagnosis. G. colchidaria objectaria is a highly variable subspecies (as the other two are). Generally, the typical objectaria has a light greyish-white ground colour and also in the form degeneraria , ground colour is rather grey than brown, but not always. Shape of frons and antennae are similar to those of sinesefida , but differences in male genitalia may help separating both: arrangement and size of subapical spines are different, sinesefida has a large, compound cornutus (small in objectaria ); octavals in G. c. sinesefida are longer, with a narrower opening between them (figs 35 & 36). Moreover, the areas of distribution of both subspecies are well separated (see map 1). G. irakensis and G. kasrunensis may occur in the same area together with G. c. objectaria , however, but both can be clearly separated from objectaria by characters of head (frons, antennae), octavals and genitalia (see diagnosis of sinesefida ). The closely related species G. cocandaria Erschoff has not been found close to the range of objectaria (diagnostic characters see under cocandaria ). Barcoding results clearly indicate that objectaria is conspecific with sinesefida and colchidaria (genetic distance about 1%), but not with cocandaria Erschoff (see fig. 54). Taxonomic notes. G. colchidaria var. objectaria was described by Staudinger (1892: 181–183) as a geographic variation (i.e. a subspecies of colchidaria ), based on one male specimen from “Tekke” (= Akhal-Tekke) or Ashkhabad, Turkmenistan . He added a female from Mardin ( Turkey ) to the types because of its very similar appearance, mainly of the under side with white base and black marginal bands. After closer examination (frons not extended, spine of foreleg tibia absent) this turned out to be a female of G. irakensis Wehrli ). A very similar male, also from the Akhal-Tekke-region, had been published and figured by Christoph (1885 : 121, pl. 6, fig. 2a, b) before, but he determined it as Boarmia cocandaria Erschoff. Staudinger , who did not agree with him, described this as var. objectaria of G. colchidaria (which also shows this contrasting under side), including the specimens figured by Christoph. Wehrli (1953: 566) considered objectaria to be different from colchidaria and raised it to species rank. He based this opinion on the different shape of the frons, the male genitalia and the shape of the octavals. DNA barcoding results, however, indicate that objectaria is conspecific with colchidaria and should be treated as a geographically as well as morphologically well separated subspecies, with G. c o l c h i d a r i a var. degeneraria Staudinger (1892: 183) being just a uniformly coloured, darker grey infrasubspecific form. The latter seems to be even more abundant, at least at the Akhal-Tekke-region. Wiltshire (1967 , 1970 ) described further new species and subspecies from NE. Iran and Afghanistan which are synonymised here with G. c. objectaria . These are (1): Gnopharmia inermis Wiltshire, 1967 , from Herat, W. Afghanistan (fig. 20): described by Wiltshire as “resembles best the degeneraria Stgr. forms from Ashkhabad, but differs in being even less marked, and also genitaliter”. The latter he describes as “without exterior thorn”. Without “exterior thorn” (= without proximal subapical spines) or with a small spine (a condition we found in the genitalia of the holotype of inermis ) is the most abundant condition in the genitalia of G. c. o b j e c t a r i a. Appearance of inermis indeed falls within the range of variation of the degeneraria Stgr. forms, as Wiltshire mentioned in the description. The type locality Herat is situated rather near to the centre of distribution of G. c. objectaria . Therefore we synonymise Gnopharmia inermis Wiltshire, 1967 , with Gnopharmia colchidaria objectaria Staudinger, 1892 . (2): Gnopharmia inermis vartianae Wiltshire, 1970 , from Jussufabad, NE Iran (figs 21 & 22): Type locality close to that of inermis inermis , but in Eastern Iran . Type specimens are even more looking like typical G. c. objectaria , with a contrasting dark marginal band on under side, upperside suffused with grey (a transitional form between objectaria and degeneraria ). Genital structures are said to be ( Wiltshire, 1970: 389 ) the same as in inermis , therefore we synonymise Gnopharmia inermis vartianae Wiltshire, 1970 with Gnopharmia colchidaria objectaria Staudinger, 1892 . (3): Gnopharmia eberti Wiltshire, 1967 from Arghandab River, 50km from Kandahar, “South-west “ Afghanistan (rather South-east Afghanistan , see map 1). The description was based on one, rather worn male specimen, the holotype (examined). Externally, it resembles typical objectaria on upper- and under side, genitalia are described as “aedeagus without large exterior thorns or even conical smaller thorns; instead, the right side has a few distal scobinations, widely spaced, distally on the sclerotized sheath”. This (and other characters mentioned in the description) perfectly agrees with an abundant condition of the genitalia of objectaria . Wiltshire (1970: 390) subsequently –invalidlydesignated a female allotype and 29 male and female paratypes . Ebert (1965: 19, 21–22) already mentioned this material ( 11 males , 21 females from Arghandab river, 30 km northern of Kandahar, 1100 m , 23./ 24.v. 1957 , leg. G. Ebert) as “ Gnopharmia objectaria Staudinger ssp.?”. He also described the appearance of this material and the variability of the male genitalia characters (which exactly describes the variabilty of the objectaria male genitalia we also found). According to circumstances unknown to us, Wiltshire (1967) described this material as G. eberti , but based it on one male only. He later (1970) added the rest of the material, as mentioned above. We here synonymise Gnopharmia eberti Wiltshire, 1967 with Gnopharmia colchidaria objectaria Staudinger, 1892 . (4): Gnopharmia maculifera kasyi Wiltshire, 1970 , from Jussufabad, NE Iran . Type locality and time of collecting are the same as those of inermis vartianae . Indeed, the single specimen ( holotype ) of kasyi is very similar to the latter, just without the grey suffusion and thus exactly agreeing with typical objectaria . Also the aedeagus, with two small proximal subapical spines and two very small, tooth-like distal subapical spines, with a wide gap between both groups, and the small, multiple cornutus is typical for objectaria . Therefore, we here synonymise Gnopharmia maculifera kasyi Wiltshire, 1970 , with Gnopharmia colchidaria objectaria Staudinger, 1892 . Life history and habitat. ( Fig. 50 ). According to the ample material studied here, the flight period starts at mid-April ( Afghanistan , Herat, 970 m ) and lasts until end of August. Specimens are attracted to light and it seems that the species is exclusively night-active. Further early records are from Golestan (North Iran , Moghadam, 8th May , altitude 1150 m ). The latest record is the 26th August at an altitude of 1300 m (Shahrud, North Iran ). The larva was reared by H. R. on Prunus ( Amygdalus ) scoparia and Prunus ( Amygdalus) spinosissima (Rosaceae) , two species of “Bitter Almonds” ocurring, for example, on the northern and southern slopes of the Kopet-Dagh mountains. Distribution (Map 1). North and Northeast of Iran to South Turkmenistan , Tajikistan , East and South of Afghanistan and Pakistan (new records for Pakistan ). It is collected at altitudes between 400 m and 2500 m a.s.l.