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18 March 2022 Taxonomy and systematics of the enigmatic emerald moth Xenochlorodes graminaria (Kollar, 1850) (Lepidoptera: Geometridae), and its assignment to a new genus
Dominic Wanke, Shamsi Feizpour, Axel Hausmann, Jaan Viidalepp, Hossein Rajaei
Author Affiliations +
Abstract

A new genus, Sabzia Wanke & Rajaei, gen. n., is described based on the Iranian species Phorodesma graminaria Kollar, 1850, which in recent taxonomy was combined with the genus Xenochlorodes Warren, 1897. Moreover, Xenochlorodes albicostaria Brandt, 1938, syn. n. is synonymized with Sabzia graminaria (Kollar, 1850), comb. n. The new genus is established based on external and internal characters of the adult specimens. Morphological traits including wing pattern, wing venation and male and female genitalia are described and illustrated.

Introduction

According to recent taxonomy, the genus Xenochlorodes Warren, 1897 consisted of nine species including X. graminaria (Kollar, 1850) and X. albicostaria Brandt, 1938, both described from, and endemic to, southern Iran (Scoble 1999; Scoble & Hausmann 2007). Viidalepp (1988) described the genus Hissarica, which was subsequently regarded as a subgenus of Xenochlorodes by Hausmann (1996). New results have demonstrated that Hissarica shows differences in wing shape, wing venation and genitalia characters, and it was therefore reinstated as a valid genus (Viidalepp & Kostjuk 2021).

In the framework of a revision of the subfamily Geometrinae in Iran (Feizpour et al., unpublished), we were faced with the unclear case of X. graminaria and X. albicostaria. Examination of specimens and of the relevant literature showed that X. graminaria and X. albicostaria most likely represent the same species. Moreover, this species differs greatly from other Xenochlorodes species in morphological characters (wing pattern and venation, male and female genitalia structures), as already suggested by Hausmann (1996), who questioned the generic assignment of X. albicostaria. In this study, we propose the synonymy of X. albicostaria with X. graminaria and assign graminaria to a new genus.

Material and methods

Type material and additional specimens examined in this study are deposited in the following collections: NHRS—Naturhistoriska Riksmuseet, Stockholm, Sweden; PCJM—Private collection of Jörg-Uwe Meineke, Kippenheim, Germany; SMNK—Staatliches Museum für Naturkunde Karlsruhe, Karlsruhe, Germany; SMNS—Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany; ZSM—Zoologische Staatssamm-lung München, Munich, Germany (SNSB).

Morphological examination

Type material and the original descriptions served for the identification and comparison of specimens. Documentation of external characters was carried out using a Visionary Digital photography system (LK Imaging System, Dun. Inc.) equipped with a Canon EOS 5DSR camera, as well as an Olympus E3 digital camera. Genitalia preparations were carried out following standard techniques (e.g., Robinson 1976) and vesicae were everted following the method described by Sihvonen (2001). Before embedding, genitalia characters were photographed in their natural position using a Keyence VHX-5000 digital microscope, following the methods proposed by Wanke & Rajaei (2018) and Wanke et al. (2019, 2021). Genitalia were then embedded in Euparal on permanent slides and photographed with the same Keyence digital microscope.

DNA barcoding and analysis

Analysis of mitochondrial DNA was carried out by submitting one dry leg of each specimen to the Canadian Centre for DNA Barcoding, Guelph, Canada. Extraction and amplification of DNA and sequencing of the barcode fragment (658 base-pairs at the 5' terminus) of the mitochondrial gene Cytochrome-C Oxidase I were performed using standard protocols (e.g., Ivanova et al. 2006). MEGA X (Kumar et al. 2018; Stecher et al. 2020) was used for the maximum likelihood analysis (with 1000 bootstrap replications) and calculation of genetic distances based on the K2P model (Kimura 1980). A list of all specimens used for the analysis is presented in Appendix 1 along with their sampling sites and Process ID numbers. Sequences, photographs and label data are accessible on BOLD, as dataset DS-SABZIA (doi:  https://dx.doi.org/10.5883/DS-SABZIA).

Systematic part

Sabzia Wanke & Rajaei, gen. n. (Figs. 1–5, 10, 11A, 12, 15)

  • Type species

  • Phorodesma graminaria Kollar, 1850: 51, 53.

  • Description

  • Wingspan: 24–32 mm in males, 22–27 mm in females. Antennae dorsally scaled, bipectinate in both male and female, branches shorter in females (longest branch about 1 mm in males, 0.25 mm in females). Frons flat, scaled, reddish-brown. Chaetosemata developed as two separate patches behind the eyes. Length of labial palpi in lateral view approximately equal to diameter of eye. Proboscis absent. Epiphysis almost equal to length of foretibia. Midtibia with one pair of spurs almost equal in size. Hindtibia with one pair of spurs, the inner spur slightly shorter than the outer spur. Thorax grass-green.

  • Wings grass-green, without transverse lines or dis-cal spots. Forewing more densely scaled and slightly darker in colour than hindwing, costa beige. Like in many Geometrinae, the wing colour may vary due to killing and preservation methods and aging. Here, the colour varies from beige to yellow (Figs. 1–5). Fringes slightly lighter than wings. Wings rather long and narrow. Venation of forewing with discocellular vein strongly angled. Vein R1 basally fused with vein Sc, the latter distally not reaching costa. Areole absent. Vein R2 fused with vein R3, veins R2–M1 on a common stalk and veins M3 and CuA1 stalked. Hindwing veins Rs–M1 and M3–CuA1 stalked (Fig. 11A). Frenulum absent in both sexes.

  • In the male genitalia (Fig. 12), the uncus and gnathos are strongly sclerotized and pincer-shaped, both apically pointed. Uncus in lateral view curved and thick. Gnathos ventrally covered with tiny, sclerotized spines; hook-shaped in lateral view, with a serrated inner edge (Fig. 12c). Socii present, membranous and broad. Valva elongated but not exceeding tip of uncus, basal part strongly sclerotized towards centre, apical part membranous. Valva slightly concave ventrally and with a heavily sclerotized central ridge (harpe). Tip of harpe broad, apical part den-tate and curved towards centre of valva. Saccus rounded (Fig. 12a). Aedeagus sclerotized and short, thin at base, broadening towards apex (Fig. 12b). Vesica without cor-nuti. Second tergite without interior processes (apodemes). Posterior margin of sternum A8 concave (Fig. 12d).

  • Female genitalia with broad papillae anales, length of apophyses anteriores approximately 1/2 length of apophyses posteriores. Lamella antevaginalis sclerotized. Ductus bursae broad and tubular, membranous; corpus bursae oval, membranous, without signa (Fig. 15).

  • Diagnosis

  • The new genus is compared to the closest genera, Xenochlorodes and Hissarica. Viidalepp & Kostjuk (2021) recently restored Hissarica from synonymy with Xenochlorodes, where it had been placed as a valid subgenus (Hausmann 1996). Sabzia graminaria comb. n. had been previously assigned to the genus Xenochlorodes (PROUT 1938). However, S. graminaria comb. n. differs from Xenochlorodes species and the monotypic genus Hissarica by the narrow and elongated wings without transverse lines (Figs. 1–10). In the forewing venation, Sabzia gen. n. differs from Xenochlorodes and Hissarica in the absence of an areole, the fusion of vein R2 with vein R3, and in having veins M3 and CuA1 stalked (areole present, vein R1 fused with veins R2+R3, and veins M3 and CuA1 con-nate in Xenochlorodes; areole present, vein R1 fused with veins R2+R3, and veins M3 and CuA1 arising separately from cell in Hissarica). In the hindwing of Sabzia gen. n., veins M3 and CuA1 are stalked (arising from cell in Xenochlorodes; arising separately from cell in Hissarica) (Fig. 11). In the male genitalia of Sabzia gen. n. the gnathos is laterally hook-shaped with a serrated inner edge, the valva has a sclerotized harpe with a broad tip, and the saccus is rounded (gnathos laterally curved, harpe absent, saccus forked in Xenochlorodes; gnathos laterally curved, harpe thin, its tip tapered, and saccus rounded in Hissarica). In Sabzia gen. n. the aedeagus is short, basally thin, broadening towards the apex (aedeagus long and slender in Xenochlorodes; aedeagus long and thick in Hissarica). The second tergite in males of Sabzia gen. n. lacks interior processes (apodemes) and the posterior margin of sternum A8 is concave (second tergite with paired, club-shaped apodemes and sternum A8 bilobed in Xenochlorodes; second tergite without any processes and sternum A8 concave with tip in the centre in Hissarica) (Figs. 12–14).

  • In the female genitalia of Sabzia gen. n. the ductus bursae is tubular and membranous and the corpus bursae is oval (ductus bursae narrowing proximally and strongly sclerotized, corpus bursae oval in Xenochlorodes; ductus bursae sclerotized, corpus bursae elongated in Hissarica) (Figs. 15–17).

  • In addition to morphological characters, we used DNA barcode data from the BOLD database to check whether or not Sabzia graminaria comb. n. clusters with the genus Xenochlorodes. The results of this analysis support our hypothesis to combine S. graminaria with a new genus, as it diverges by 11% from the type species of Xenochlorodes, X. olympiaria (Herrich-Schäffer, [1852]) (Fig. 18). Furthermore, the genetic distance from all other Palearctic Geometrinae is approximately 10%. We emphasize here that these results from COI barcoding alone should not be regarded as sound data for drawing any phylogenetic conclusions.

  • Tribal assignment

  • While placed within the genus Xenochlorodes, Sabzia graminaria was previously assigned to the tribe Hemistolini. According to the results of Murillo-Ramos et al. (2019), who subordinated Hemistolini under Hemitheini, we tentatively assign Sabzia gen. n. to the tribe Hemitheini. However, a comprehensive molecular phylogenetic study including Sabzia gen. n. is recommended for a reliable tribal assignment of the new genus.

  • Etymology

  • The name of this genus derives from the Persian word “Sabz”, meaning green, and refers to the wing colour of type species Sabzia graminaria comb. n.

  • Figs. 1–10.

    Wing colouration and pattern of Sabzia Wanke & Rajaei, gen. n., Xenochlorodes Warren and Hissarica Viidalepp spp. (a = upperside; b = underside). – 1–5. S. graminaria (Kollar, 1850), comb. n. [1–3: Iran, Fars, Komehr; 2: g. prep. 11052; 3: g. prep. 11051; 4: Iran, Fars, Miyan Kotal, g. prep. 272/2016 S.F.; 5: Illustration from PROUT (1935)]. 6–7. X. olympiaria (Herrich-Schäffer, [1852]) [6: Krk, Fiumebucht, g. prep. 1087/2021, D. Wanke; 7: Turkey, Mugla, vic. Esen, g. prep. 1088/2021, D. Wanke]. 8–9. H. postalbida Viidalepp, 1988 [all Tadjikistan, Kondara; 8: g. prep. 1091/2021, D. Wanke; 9: g. prep. 1090/2021, D. Wanke]. 10. Adult of S. graminaria comb. n. from Iran (Kerman, Hanna protected area; photo by H. Rajaei). Scale bar: 1 cm.

    img-z3-1_01.jpg

    Fig. 11.

    Wing venation of (A) Sabzia graminaria (Kollar, 1850), comb. n., (B) Hissarica postalbida Viidalepp, 1988 and (C) Xenochlorodes olympiaria (Herrich-Schäffer, [1852]). Areole blue coloured; area between R1 to R3 veins orange coloured. No areole is present in S. graminaria comb. n. (rectangle A1) and one areole (blue coloured) is present in Xenochlorodes (rectangle C1). Remark: according to Viidalepp & Kostjuk (2021), veins R2–M1 in the genus Hissarica (C) are stalked from the cell apex, R2 diverges after M1 and anastomoses with R1 towards the costa, therefore we here name this vein as R1+2. In Sabzia Wanke & Rajaei, gen. n. (B), veins R2–M1 are also stalked from the cell apex, but R2 does not diverge after M1 but reaches the costa along with R3; therefore, we here name this vein as R2+3.

    img-z4-1_01.jpg

    Figs. 12–14.

    Male genitalia of Sabzia Wanke & Rajaei, gen. n., Xenochlorodes Warren and Hissarica Viidalepp spp. – 12. S. graminaria (Kollar, 1850), comb. n. (a, c, d: Iran, Kerman, Jiroft, g. prep. 1092/2021, D. Wanke; b: Iran, Fars, Komehr, g. prep. 11051). 13. X. olympiaria (Herrich-Schäffer, [1852]) (a–d: Krk, Fiumebucht, g. prep. 1087/2021, D. Wanke; e–f: Spain, Girona, Puigventós, g. prep. 1086/2021, D. Wanke). 14. H. postalbida Viidalepp, 1988 (a–d: Tadjikistan, Kondara, g. prep. 1091/2021, D. Wanke). a = genitalia capsule; b = aedeagus; c = uncus, lateral view; d = sternum A8; e = saccus; f = tergite 2. Scale bar: 1 mm; c, d, f are out of scale.

    img-z5-1_01.jpg

    Figs. 15–17.

    Female genitalia of Sabzia Wanke & Rajaei, gen. n., Xenochlorodes Warren and Hissarica Viidalepp spp. – 15. S. graminaria (Kollar, 1850), comb. n. (Iran, Esfahan, vic. Sar Chesmeh, g. prep. 1093/2021, D. Wanke). 16. X. olympiaria (Herrich-Schäffer, [1852]) (Turkey, Mugla, vic. Esen, g. prep. 1088/2021, D. Wanke). 17. H. postalbida Viidalepp, 1988 (Tadjikistan, Kondara; g. prep. 1090/2021, D. Wanke). Scale bar: 1 mm.

    img-z6-1_01.jpg

    Fig. 18.

    Maximum likelihood analysis of Sabzia Wanke & Rajaei, gen. n. and closely related Palearctic species from different genera and tribes, based on COI 5' sequences (built with MEGA X; Kimura 2-parameter model; bootstrap method, 1000 replications). Remark: the COI data used here were only used to check whether or not S. graminaria (Kollar, 1850), comb. n. clustered with the genus Xenochlorodes Warren, and not for phylogenetic purposes; for a sound phylogenetic analysis of Hemitheini, see Murillo-Ramos et al. (2019).

    img-z7-1_01.jpg

    Sabzia graminaria (Kollar, 1850), comb. n.

  • Phorodesma graminaria Kollar, 1850: Kollar 1850: 51, 53. Holotype ♂ (South West Persia: Schiraz). Holotype not traced, possibly lost.

  • Xenochlorodes albicostaria Brandt, 1938: Brandt 1938: 55. Syn-types ♂♀ ([Iran]: Comèe [Komehr], Bam-i-Firus). Hereby regarded as a new synonym of Sabzia graminaria, syn. n. based on morphological examination and sympatric occurrence (see taxonomic note, below).

  • Material examined

  • Paratype ♂ of X. albicostaria Brandt, 1938: Iran, Fars, Straße Ardekan-Talochosroe [Tall Khosrow], Comèe [Komehr], 3600 m, 25.vi.1937, coll. Brandt, g. prep. ZSM G No. 1545 (ZSM).

  • 1 ♂, 1 ♀, same data as paratype of X. albicostaria, 2600 m, 30.vi.1937, coll. Brandt, g. preps (♂) 11051, (♀) 11052 (NHRS).

  • 2 ♀, Iran, Prov. Esfahan, NE of Naraq, Kuh-e Goran, 34°05′N 50°54′E, 2500 m, 06.vii.2003, leg. G. Ebert & R. Trusch; 1 ♀, Iran, Esfahan, Esfahan, Daran, Ashan vic., 2490–2500 m, 25.vi.2005, leg. A. Hofmann; 1 ♀, Iran, Boyer Ahmad-va-Kohgiluyeh, Yasuj E, Abnar Region, Kakan-Baba Hassan, 2550–2800 m, 24.vi.2005, leg. A. Hofmann; 1 ♂, Iran, Fars, Straße Ardekan-Talochosroe [Tall Khosrow], Comèe [Komehr], ca. 2600 m, 30.vi.1937, coll. Brandt; 1 ♂, Iran, 4–7.vi.1969, Miyan Kotal, 1900 m, östl. Kazerun, 29°30′N 51°40′E, leg. G. Ebert, g. prep. 272/2016 S.F.; 1 ♀, Iran, Fars, Kaserun, Mian-Kotal, 1900 m, 11.vi.1972, leg. Ebert & Falkner, g. prep. 273/2016 S.F. (all SMNK).

  • 1 ♀, Iran, Boyer Ahmad-va-Kohgiluyeh, Yasuj E, Abnar Region, Kakan–Baba Hassan, 2550–2800 m, 24.vi.2005, leg. A. Hofmann; 1 ♀, Iran, Esfahan, Miyandasht NW Afous, Chebleh-Kuh, Sar Chesmeh vic., 2700–2900 m, 12./13.vii.2003, leg. A. Hofmann, J.-U. Meineke, G. Tremewan, g. prep. 1093/2021 D. Wanke; 1 ♂, Iran, Kerman, Jiroft W, Shingera, 2800 m, 26./27.v. 2004, leg. A. Hofmann, J.-U. Meineke, G. Tremewan, g. prep. 1092/2021 D. Wanke (all SMNS).

  • 1 ♀, Iran, Markazi, Tafresh 10 km SSE (Pass), 2400 m, 31.v.2005, leg. A. Hofmann, J.-U. Meineke; 1 ♂, 2 ♀, Iran, Boyeramabad va Kuhgiluye, Kuh-e Dinar, Paß E Sisahkt, 2800–3200 m, 14.vii.2000, leg. ten Hagen, g. preps (♂) 101, (♀) 100/2016 S.F.; 1 ♀, Iran, Boyer Ahmad-va-Kohgiluyeh, Gardaneh, Meymand, 2450–2800 m, 14./15.vi.2001, leg. A. Hofmann, J.-U. Meineke, G. Tremewan, g. prep. 162/2016 S.F.; 1 ♂, Iran, Esfahan, N Fereydun Shahr, vic. Shoohroud, 2700–3200 m, 5./6.vii.2002, leg. ten Hagen, g. prep. 102/2016 S.F.; 1 ♂, 1 ♀, Iran, Kerman, Kuh-e Bochrasman, S Darbmazar, 2800–3000 m, 1.vi.2004, leg. W. ten Hagen, g. prep. (♂) 106/2016 S.F.; 1 ♂, 1 ♀, Iran, Kerman, Jiroft NW, Gardaneh, Sarbishan, Shingara vic., 2700–2900 m, 3./4.vi.2002, leg., J.-U. Meineke, A. Hofmann, A. Kallies et al., g. prep. (♂) 103/2016 S.F.; 1 ♀, Iran, Kerman, Bam SW, Deh Bakri, 2000–2200 m, 1.-3.vi.2002, leg. J.-U. Meineke, A. Hofmann, A. Kallies et al., g. prep. 105/2016 S.F.; 1 ♂, 1 ♀, Iran, Kerman, Jiroft W, Shingera, 2800 m, 26./27.v. 2004, leg. A. Hofmann, J.-U. Meineke, G. Tremewan, g. prep. (♂) 104/2016 S.F. (all PCJM).

  • Taxonomic note

  • Kollar (1850) described Phorodesma graminaria based on one damaged male, the holotype, from southern Iran. According to Prout (1935), the holotype was left without a type label and, as already mentioned by Kollar (1850), was in damaged condition (five legs and the abdomen missing). Unfortunately, we could not find the holotype in any of the examined collections. Nevertheless, the original description and the additional notes of Prout, as well as his illustration (Prout 1935: 18, row h), served for identification of this species in the present study. Additionally, our revision of all known Iranian Geometrinae species has shown that S. graminaria is the only species with entirely green wings without markings, and that it therefore cannot be confused with other Geometrinae in this region. Considering all of the above, a neotype designation is not considered necessary, as the taxonomy of this species is stable and future retrieval of the holotype cannot be ruled out.

  • Surprisingly, X. albicostaria, described by Brandt (1938) from the southern Iranian province Fars, shows a high similarity with S. graminaria. It matches in all external characters, such as the long and narrow wing shape, the green colouration of the wings, the lack of transverse lines on the wings, the two terminal spurs on the hindleg, the absence of a frenulum, the strongly angled discocellular vein, vein R2 situated near the cell apex and vein M1 arising from the common stalk of veins R2–5 (Figs. 1–5). We therefore synonymize X. albicostaria with S. graminaria, syn. n.

  • Description and diagnosis

  • See generic part.

  • Phenology and habitat

  • Flying from May to July at altitudes from 1900 to 3600 m.

  • Biology

  • Unknown

  • Distribution

  • Species endemic to Iran, occurring from western to southern Iran along the Zagros Mountains.

  • Acknowledgements

    We wish to express our deepest gratitude to Robert Trusch and Michael Falkenberg (both Staatliches Museum für Naturkunde Karlsruhe, Germany), Tobias Malm and Johannes Bergsten (both Naturhistoriska Riksmuseet, Stockholm, Sweden) and Jörg-Uwe Meineke (Kippenheim, Germany) for the loan of valuable specimens from their collections. Many thanks to Norbert Pöll (Bad Ischl, Austria) for sharing the barcode data of S. graminaria with us. We thank Hamid Valipoor (Quchan, Iran) for accompanying the last author on several expeditions in Iran. Thanks to James E. O'Hara (Ottawa, Canada) for providing an indispensable translation of the Latin in the original description of S. graminaria. We are grateful to the editor-in-chief of Integrative Systematics Daniel Whitmore and to two anonymous reviewers for their critical review of the submitted version of the paper and constructive comments.

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    Appendices

    Appendix 1.

    List of specimens used for the maximum likelihood analysis, with species, sampling site and Process ID. Data taken from BOLD, generated by Axel Hausmann except GWORU553-10, generated by Norbert Pöll.

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    Continued

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    © Staatliches Museum für Naturkunde Stuttgart
    Dominic Wanke, Shamsi Feizpour, Axel Hausmann, Jaan Viidalepp, and Hossein Rajaei "Taxonomy and systematics of the enigmatic emerald moth Xenochlorodes graminaria (Kollar, 1850) (Lepidoptera: Geometridae), and its assignment to a new genus," Integrative Systematics: Stuttgart Contributions to Natural History 5(1), 61-71, (18 March 2022). https://doi.org/10.18476/2022.857803
    Received: 8 October 2021; Accepted: 18 February 2022; Published: 18 March 2022
    KEYWORDS
    Iran
    Middle East
    new combination
    new synonymy
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