Nychiodes divergaria Staudinger, 1892

Wanke, Dominic, Hausmann, Axel, Krogmann, Lars, Petrányi, Gergely & Rajaei, Hossein, 2020, Taxonomic revision of the genus Nychiodes Lederer, 1853 (Geometridae: Ennominae: Boarmiini) with description of three new species-an integrative approach, Zootaxa 4812 (1), pp. 1-61 : 32-34

publication ID	https://doi.org/ 10.11646/zootaxa.4812.1.1
publication LSID	lsid:zoobank.org:pub:74FE6D6A-B41D-4FC9-BC26-B07DDEB9A0F6
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scientific name	Nychiodes divergaria Staudinger, 1892
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Nychiodes divergaria Staudinger, 1892 View in CoL

( figs 52–72, 113–119 View FIGURES 108-114 View FIGURES 115-118 View FIGURE 119 , 141, 142 View FIGURES 135-144 ; maps 4, 5)

Nychiodes divergaria divergaria Staudinger, 1892 View in CoL . Deutsche Entomologische Zeitschrift Iris 5, 171. Syntypes ♂ (Mesopotamia [ Turkey]: Mardin, Egin ) (in MNHU, examined).

Nychiodes divergaria achtyca Wehrli, 1939 View in CoL . Entomologische Rundschau 56 (33), 365. Syntypes 3♂ ([Transcaucasus]: Rjabov) (in ZFMK, examined). Hereby regarded as a new synonym of Nychiodes divergaria View in CoL based on morphological examination and sympatric occurrence of these forms.

Nychiodes divergaria elbursica Wehrli, 1937 . Lambillionea 37 (8-9), 187. Syntypes 11♂ 4♀ ([ Iran]: Elburs Mts.), (in NHRS, examined). Hereby regarded as a new synonym of Nychiodes divergaria based on morphological examination and sympatric occurrence of these forms.

Nychiodes divergaria fallax Wehrli, 1939 . Entomologische Rundschau 56 (33), 366. Holotype ♂ (Transcaucasus), (in ZFMK, examined). Hereby regarded as a new synonym of Nychiodes divergaria based on morphological examination and sympatric occurrence of these forms.

Nychiodes variabila Brandt, 1938 View in CoL . Entomologische Rundschau, 55 (51), 37. Syntypes ♂, ♀ ([ Iran]: Comèe , Fort Mian-Kotal , Fort Sine-Sefid), (in NHRS, examined). Hereby regarded as a new synonym of Nychiodes divergaria View in CoL based on morphological and molecular examination.

Nychiodes variabila opulenta Brandt, 1941 View in CoL . Mitteilungen der Münchner Entomologischen Gesellschaft, 31, 878. Syntypes ♂, ♀ ( Iran, Baloutchistan , Kouh i Taftan (Khach )), (in NHRS, examined). Hereby regarded as a new synonym of Nychiodes divergaria View in CoL based on morphological examination and sympatric occurrence of these forms.

Nychiodes variabilis . Incorrect subsequent spelling of Nychiodes variabila View in CoL syn. nov. of Nychiodes divergaria View in CoL in Wiltshire, 1943. Journal of the Bombay Natural History Society, Bombay, 43, 630.

Nychiodes variabilis . Incorrect subsequent spelling of Nychiodes variabila syn. nov. of Nychiodes divergaria in Wiltshire, 1957. The Lepidoptera of Iraq. Government of Iraq (Ministry of Agriculture), 111.

Type material examined. Nychiodes divergaria , Lectotype (herewith designated), ♂, Origin, [ Turkey], Mardin, g.prep. 2106/2017 H. Rajaei ; Paralectotype (herewith designated), 1 ♂, Origin, [ Turkey], Egin, g.prep. 2107/2017 H. Rajaei; all in MNHU .

Nychiodes divergaria achtyca . Syntype, 1 ♂, Dagestan, Ackty, 28.vii.[19]33, M. Rjabov, g.prep. 7272; in ZFMK .

Nychiodes divergaria elbursica Holotype, ♂, Persia, Elbursgebirge , Rehne-Demavend, 2600 m, 21.- 27.vii.[19]36, Schwingenschuss; in ZFMK.

MAP 4. Distribution pattern of N. divergaria . This species is widely distributed from Turkey to south-eastern Iran. MAP 5. Distribution pattern of male N. divergaria specimens indicating on a cline in the Middle East. The ampulla inferior is highly variable from absent to present in this species. An increase in the development of this structure is visible from Southern Iran to Turkey.

Nychiodes divergaria fallax Holotype, ♂, Transcaucasus, Arax, Darasham, v. 1934, M. Rjabov g.prep. 7273; in ZFMK.

Nychiodes variabila Holotype, ♂, Iran, Fars, Strasse Chiraz-Kazeroun, Fort Sine-Sefid , ca. 2200 m, 25.v.1937, coll. Brandt, g.prep. 10930 ; Paratypes, 1 ♀, same locality, ca. 2000 m, 8.-7.v.1937, coll. Brandt, (labeled as Al- lotype) , g.prep. 10931; all in NHRS. 1 ♂, same locality, ca. 2200 m, Sept.1937, coll. Brandt ; 1 ♀, same locality, 28.v.1937, coll. Brandt ; 1 ♂, same locality, Mai.1937, coll. Brandt ; 1 ♂, 1 ♀, same locality, 10.v.1937, coll. Brandt ; all in ZFMK.

Nychiodes variabila opulenta Holotype ♂, Iran, Baloutchistan , Kouh i Taftan (Khach), 2500 m, 26.iv.1938, coll. Brandt, g.prep. 10928 ; Paratypes, 1 ♂, Iran, Baloutchistan , Kouh i Taftan (Khach), 2500 m, 3.v.1938, coll. Brandt, (labeled as Allotype) g.prep. 10929; NHRS . 1 ♂, same locality, 3000 m, 20.vi.1938, coll. Brandt ; Paratype 1 ♂, same locality, 2500 m, 17.vi.1938, coll. Brandt; all in ZFMK .

Additional material studied: 232 ♂, 84 ♀ (see appendix).

Note. Nychiodes divergaria is the most variable species within this genus, in sense of wing pattern and male genitalia (see figs 52–72). This extreme variability caused the description of several species and subspecies, which are regarded as synonymous taxa based on the extensive morphological and molecular examinations in the current study. In male genitalia, the most variable structure is the ampulla inferior of the valva. Examination of 180 genitalia slides and plotting these on a geographic map reveals a clinous charater distribution of the size of the ampulla inferior, starting from southern Iran (without ampulla inferior) to south-western Turkey (with well-developed ampulla inferior) (see map 5). The female genitalia show stable diagnostic characters with little variation between populations.Another reliable differential character is the costa of valva in the male genitalia. Therefore, we strongly recommend dissection for secure species identification of N. divergaria .

Diagnosis. Wingspan ♂ 25–40 mm, ♀ 29–44 mm (forewing length ♂ 15–22 mm, ♀ 17–25 mm) ( figs 52–72). Ground colour of wings highly variable, differing from sandy yellow and light brown to dark, almost black specimens. N. divergaria can be confused with most sympatric congeners ( N. farinosa , N. rayatica , N. subfusca , N. leviata , N. subvirida , N. eberti sp. nov.). Diagnostic characters of male and female genitalia in comparison to the above mentioned sympatric species are presented below.

Male genitalia ( figs 113–119 View FIGURES 108-114 View FIGURES 115-118 View FIGURE 119 ) with costa of valva wide, strongly sclerotized, reaching the apex of valva (in rare cases exceeding apex, figs 119 View FIGURE 119 J–P) (costa of valva exceeding tip of valva, slightly curved in N. farinosa ; apically extremely dilated, exceeding tip of valva in N. rayatica ; curved towards uncus in N. subfusca ; curved in N. leviata ; medially humped in N. subvirida ; costa of valva narrow, digitiform in N. eberti sp. nov.) (see figs 98 View FIGURES 96-102 , 105, 106 View FIGURES 103-107 , 108, 110, 111 View FIGURES 108-114 , 119 View FIGURE 119 , 120, 122 View FIGURES 120-126 ). Ampulla superior long, broad, usually tubular; ampulla inferior highly variable, from absent state to thin and long in size, if present always much narrower than ampulla superior (see fig 119 View FIGURE 119 ) (ampulla superior apically dilated, ampulla inferior slightly curved in N. farinosa ; ampulla superior conical, broad, ampulla inferior strongly curved in N. rayatica ; ampulla superior narrow, apically dilated, ampulla inferior short and broad in N. subfusca ; ampulla superior broader than ampulla inferior, both with similar length and curved in N. leviata ; both ampullae slightly conical in N. subvirida ; both ampullae digitiform, characteristically located in the distal half of valva in N. eberti sp. nov.) (see figs 98 View FIGURES 96-102 , 105, 106 View FIGURES 103-107 , 108, 110, 111 View FIGURES 108-114 , 119 View FIGURE 119 , 120, 122 View FIGURES 120-126 ).

Female genitalia N. divergaria with strongly reduced apophyses anteriores, and two strongly sclerotized spherical patches on sternite A9 which cannot be confused with any other species of the amygdalaria species-group (see figs 141, 142 View FIGURES 135-144 ).

Phenology. Three generations (as mentioned also by Wiltshire 1957), flying from March to November.

Biology. In Iraq feeding observations on trees and shrubs of Rosaceae ( Prunus , Amygdalus and cultivated apricot) ( Wiltshire 1957).

Habitat. Occurring in the woodland zone of the mountains and in apricot orchards in lower plains; in the Zagros range inhabiting scrub woods ( Wiltshire 1957). In altitudes from 400 up to 3000 m.

Distribution. Anatolian-Iranian, from southern Turkey, southern Armenia, to western, northern and south-eastern Iran (maps 4, 5).

DNA barcoding. Clustering with N. subvirida in a ‘ divergaria - subvirida -complex’, but showing diagnostic morphological characters, supporting the taxonomic validity of both taxa at species rank. Nearest species (minimum pairwise distances): N. eberti sp. nov. (3.1%) ( fig. 145 View FIGURE 145 ).