Anthocharis damone shahrdarii Naderi and Back, 2025

Naderi, Alireza, Back, Werner, Scalercio, Stefano & Nazari, Vazrick, 2025, Higher phylogeny of Orange Tips (Anthocharis) with a focus on population structure in A. damone (Lepidoptera, Pieridae), Ecologica Montenegrina 83, pp. 74-91 : 84-91

Anthocharis damone shahrdarii Naderi and Back , ssp. nov. ( Fig. 4 m –p )

https://zoobank.org/ urn:lsid:zoobank.org:act:EB69365D-FC90-4D19-BD22-317E4778C919

Holotype: ♂ Iran: Alborz: Karaj, Azgi Dareh , 1600 m, 25.IV.2008, leg. A. Naderi (coll. MMTT).

Paratypes: Iran: Mazandaran: 1♂ 1♀ Ramsar, Samamus Mt. , 17.VI.2005, leg. A. Naderi (coll. MMTT) ; 2♂ ibid (coll. Naderi) ; 2♀ Veresk, Oshtorinankuh Mt. , 3.VI.2004, leg. A. Karbalaye (coll. Back) ; 1♂ 1♀ Kelardasht, Sirtchal , 8.V.1983, leg. Khial. (coll. HMIM) ; 1♀ Polur, Sarcheshmeh , 25.v.1990, Anonymous leg. (coll. HMIM) . Qazvin: 13♂, 2♀ N of Qazvin , 2500 m, 30.IV.2004, leg. Naderi (coll. Naderi) ; 1♂ ibid (coll. Back) ; 1♂ N. Qazvin, Kafar Meydan , 26.V.2011, leg. A. Naderi (coll. Back) ; 3♂ 1♂ Highlands N. of Qazvin, Aloulak , 17.V.2010, leg. J.D. Bahrami (coll. Zehzad) .

Semnan: 1♂ N. Aradan, Rameh village , 2.V.2018, leg. A. Naderi (coll. Naderi) . Alborz: 1♂ Karaj, Azgi dareh, 1600 m, 25.IV.2008, leg. A. Naderi (coll. Naderi); 7♂, 4♀ ibid (coll. Back); 1♂ ibid, 25.V.2007, leg. A. Naderi (coll. Naderi); 1♂ 15 km N. Karaj, E. Aderan, 4.IV.2018, W. & C. Back (coll. Back); 3♂ Karaj, Sijan (coll. Back); 5♂ Karaj, Pamchal, 2, 11, 18.V.1994, leg. Nazari (coll. HMIM); 1♀, Karaj, Vardeh, 1600-1800m, 19.V.1995, leg. Nazari (coll. HMIM); 2♂ 1♀ Dizin, 2500-2700 m, 18.VI.2009, leg. A. R. Naderi (coll. Zehzad); 1♂ 1♀ Dizin, 2800-3000 m, 12.VI.2007, leg. A. R. Naderi (coll. Zehzad); 21♂, 22♀ Dizin, 2000 m, 3.VI.2001 (coll. Back); 1♀ ibid, 2200-2600m, 22.V.2001, leg. Naderi (coll. Naderi); 1♂ Dizin, 18.VI.2009, leg. A. Naderi (coll. Back); 2♂ Chalus road, Dizin, 18.VI.2004, leg. A. Naderi (coll. Naderi); 1♂ 5 m NE Gatchsar , Dizin rd. , 26.V.2013, leg. W. Back (coll. Back); 31♂, 18♀ ibid, 2000 m, 18.V.2013 (coll. Back) . Tehran: 1♂ N . Tehran, Darband , 2500m, 11.VI.1994, leg. Nazari (coll. HMIM); 1♂ ibid, 1900 m, 23.IV.2017, leg. A. R. Naderi (coll. Zehzad); 1♂ ibid, 1800 m, 15.IV.2021, leg P. Zehzad (coll. Zehzad); 3♂ 1♀ N Tehran, Tochal, 2500 m, leg P. Zehzad (coll. Zehzad); 1♂ Firuzkuh, Chehelcheshmeh , 2000 m, 24.vi.1998, Mofidi-Neyestanak leg. (coll. HMIM); 5♂, 3♀ Damavand, 3000 m, 7.VII.1995 (coll. Back); 1♀ Damavand, Polur-Rineh, 2800- 3200 m, 17.vi.1995, Nazari leg. (coll. HMIM); 1♂ 1♀ Fasham, Rudak [mislabeled as Gheshm Island, Rudak], 20.v.1967 (coll. HMIM) .

Distribution. Alborz Mountains in Northern Iran.

Etymology: The new taxon is named after Mr. Alireza Shahrdari (Tehran, Iran), a wildlife conservationist and a close friend of the first author.

Description: Male. Fw length 17-20 mm (Holotype 20 mm). Ups gc dull yellow with dull orange apex. The triangular black patch on the tip of orange area usually totally black without any disconnection. The black line of fw in discal area is irregular and is completely stuck to both outer margin and costal area. The discal spot is round. The black line of fw between orange triangular and yellow background is usually absent and sometimes present. Hw mostly yellow without any black spots or dots. Uns ♂ fw with clear yellow apex and, without any regular pattern. Hw with irregular green patches with in yellow background. Female. Fw length: 20-21mm. Ups gc snowy white with totally black apex or sometimes with some white short lines on S3–S5. The discal spot regular without extra lines and borders. Uns as in ♂.

Variation: Size is relatively constant. Males from Samamus are dull pale-yellow while in southern slopes of Alborz the ground colour is deeper yellow. In some individuals the black line separating the orange apex from inner of forewing is absent, while in others it is black and prominent. The black lunulus in the marginal area in both sexes are sometimes well developed and similar to those in ssp. karabalyei .

Differential diagnosis. ♂ size is moderate and similar to ssp. karbalayei and ssp. eunomia but closer to ssp. eunomia . Ups the yellow gc and the orange apex are fair and clear. Most specimens lack the black line separating the orange apex and yellow background. The ups are simply yellow and orange in ♂ while in ssp. karbalayei on the hws a black spot is visible in the in outer margin of S8. In ssp. syra , the black discoidal spots, the apical markings, and the black basal border of the apical orange patch are weakly developed, giving it a lighter look ( Back & Bozano, 2020). Uns simple yellowish with an orange band in fw and sparse green patches in hws, while it is densely patched in ssp. karbalayei ( Back & Bozano, 2020) . ♀ fw gc is snowy white while in ssp. karbalayei it is duller. The black apex has deep white triangular patches, while in the other subspecies these patches are shorter in ssp. karbalayei and ssp. syra . Uns the green shadow-like and irregular patches of hws are less dense than the other subspecies.

Notes. A. damone shahrdarii is an early flier in the steep rocky mountain slopes along the riversides and gardens and roads. The first males emerge while their host plants are still in their early flowering period. Its accompanying species include Tomares callimachus , Anthocharis gruneri , Euchloe ausonia and Euchloe lessei . The host plants of the populations of ssp. eunomia in Alborz are Isatis species (e.g. I. kotschyana Boiss. & Hohen. , I. tinctoria L., and I. cappadocica Desv. ). An albino form of A. damone from Alborz has been reported by ten Hagen (2008).

Discussion

Considering the extent of historical attention paid to Anthocharidini since even before Linnaeus’ time (e.g. see Schenk & Sell 2024), it is surprising that the higher systematics of this group should still be debated. The recent suggestion by Zhang et al. (2021) that the well-established pierid genus Zegris should be sunk a subgenus of Anthocharis goes directly against the status quo of the past 300 years in this group. The phylogeny by Zhang et al. was based on a small sample size of six species ( Paramidea scolymus , Tetracharis cethura , T. midea , Zegris eupheme and Z. pyrothoe ), with three species of Euchloe ( E. charlonia , E. crameri , E. tagis ) used as outgroups. Previously, molecular studies had suggested Euchloe + Zegris as sister to Anthocharis ( Braby et al. 2006; Wahlberg et al. 2014; Braga et al. 2021; Wiemers et al. 2020; but see Marabuto et al. 2020), however none of these studies included members of subgenera Tetracharis or Paramidea . Our well-supported phylogenetic hypothesis based on a larger sampling and several nuclear genes does not support the notion put forth by Zhang et al. Instead, based on the paraphyly of the proposed subgenera within Anthocharidini (sensu Zhang et al. 2020), we propose that Tetracharis and Paramidea should be recognized as separate genera in their own right (stat. nov.), and not as subgenera of Anthocharis .

Our investigation into the fine-scale population structure in A. damone deals with yet another controversial subject in biological studies, namely the concept of the subspecies. Mayr’s definition of subspecies, i.e. a geographical aggregate of local populations that differs genetically and taxonomically from other subdivisions of the species ( Mayr, 1942; O'Brien and Mayr, 1991), is still widely employed as a useful construct in documenting biodiversity. Other proposed definitions also emphasize isolation, allopatry and diagnosability (e.g. Ramsen, 2010; Braby et al. 2012) and highlight the differences between subspecies and clines: While the emergence of subspecies is believed to be associated with historical dynamics that determine species ranges (isolations during glaciations and subsequent spreading from refugia), clines are determined by the current ecological circumstances and the adaptive population response to the impact of environmental factors changing along spatial gradients ( Thorpe, 1987; Thorpe et al. 1991). Accordingly, subspecies should be groups of geographically separated populations that are phenotypically diagnosable ( Vinarski, 2015).

In their descriptions of subspecies for A. damone , most authors (e.g. Verity, 1911, 1947; Seyer, 1980, 1984, 1985 and others) have relied on very subtle phenotypic differences between individuals from different populations. Even though some of these names have been subsequently synonymized (e.g. see Hesselbarth et al. 1995), most contemporary authors find merit in maintaining subspecies-group names for A. damone (e.g. Tshikolovets et al. 2014; Tshikolovets & Nekrutenko, 2013; Tshikolovets & Ben Yehuda, 2020; Back & Bozano, 2020). Existence of a genetic gap between these populations further supports their historical isolation.

The genetic distance between South Calabrian/Sicilian populations of A. damone ( ssp. damone ) with those in northern Calabria was already noted by Scalercio et al. (2020) who posited that this difference does not seem to be due to a recent geographic barrier. Our calibrated BEAST phylogeny suggests that the split in the last common ancestor of these two populations occurred around 4.83 million years ago in Pliocene, after the Zanclean flood refilled the Mediterranean basin and ended the Messinian salinity crisis (5.96 to 5.33 mya) (Supplementary Figure S4 ). As a result of the sea level oscillations during Pliocene, Calabria underwent repeated transformation into an archipelago, splitting the range of the ancestral population of A. damone into a chain of islands. This was likely followed by more recent dispersal of the Sicilian population to the southern tip of Calabria during the Pleistocene cold stage ( Schmitt et al. 2021). These processes have also been instrumental in shaping the evolution of another pierid butterfly, Euchloe tagis , in the Western Mediterranean ( Marabuto et al. 2020).

Even though the distribution of the populations in the Balkans ( Serbia, Macedonia, Albania) ( ssp. hollanderi ) appears to be continuous towards Greece ( Drndić et al. 2017), there is a clear genetic gap between samples from Macedonia / Serbia and those in central ( ssp. arachovae ssp. nov.) and southern Greece ( ssp. morea ssp. nov.), suggesting historical periods of genetic isolation in this region. The populations in E Turkey, Armenia, Georgia and NW Iran ( ssp. eunomia ) show a large degree of variation in phenotype but also in their mtDNA barcodes, consistent with clinal variation within a single evolutionary unit. The split between this group and their closest sister population ( ssp. shahrdarii ssp. nov.), isolated in the Alborz mountains in N Iran, seems to have occurred at around 3.69 mya (Supplementary Figure S4 ).

Additional sampling and genomic work are required to determine the exact distributional boundaries of these taxa and the presence or degree of gene flow between them. Orange Tips are early fliers and are typically among the very first butterflies that appear in the spring after the snow melts. Vast areas of suitable habitats within the Palaearctic range of Anthocharis that have not yet been explored in the appropriate season may potentially hold additional new taxa yet to be discovered and described in future.

Acknowledgements

We thank Alireza Shahrdari, Ehsan Jannati and Gholamreza Naderi (Tehran, Iran) who accompanied the first author in his field trips, Dr. Farshad Fattahi (Kermanshah, Iran), Payam Zehzad (Tehran, Iran) and Amirhossein Harandi (Esfahan, Iran) for their support and valuable information and materials, Dr. Valiollah Mozaffarian (Tehran, Iran) for botanical help in identification of host plants, and Amirhossein Bakhtiyari (Iranshenasi Publishing, Tehran, Iran) for assistance with an earlier version of the distribution map. Helen Alipanah (Tehran, Iran) kindly supplied data on material stored at the HMIM (Tehran). We also would like to thank Sara La Cava, Giuseppe Rijllo, and Giada Zucco (Rende, Italy) for their help during field observations in Calabria, and Maximilian von Stetten (Freising, Germany) for his assistance with the photographs. Dr. Alexander Kir’yanov (León, Mexico) provided valuable comments on an earlier draft of this manuscript.

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Supplementary Material

Supplementary Table S1

Material examined and GenBank accessions.

Data type: Excel document

Link: https://www.biotaxa.org/em/article/view/86983/81814

Supplementary Figure S2

Larval images. A) A. d. damone , Italy, Sicily, Etna, 1972, W. Back. B) A. d. calabra, Italy, Calabria, 2023, S. Scalercio. C-D) A. d. hollanderi, North Macedonia, Skopje , Treska Schlucht , V.1975, W. Back. E) A. d. arachovae, Greece, Delphi, 3.5.1991, W. Back. F-G) A. d. shahrdarii, Iran, Alborz, Sijan, IV.2017, W. Back. H) A. d. karbalayei, Iran, Esfahan, Fereidunshahr , 2013, W. Back .

Data type: JPG file

Link: https://www.biotaxa.org/em/article/view/86983/81815

Supplementary Figure S3

Pupal images. A) A. d. calabra ssp. nov., Italy, Calabria, 2023, S. Scalercio. B) A. d. hollanderi, North Macedonia, Skopje, Treska Schlucht, V.1975, W. Back. C-D) A. d. shahrdarii ssp. nov., Iran, Alborz, Dizin V. 2013, W. Back. E) A. d. karbalayei, Iran, Esfahan, Fereidunshahr, 2013, W. Back .

Data type: JPG file

Link: https://www.biotaxa.org/em/article/view/86983/81816

Supplementary Figure S4

BEAST tree with median ages.

Data type: PDF document

Link: https://www.biotaxa.org/em/article/view/86983/81817

Supplementary Figure S5

BEAST tree with posterior probabilities.

Data type: PDF document

Link: https://www.biotaxa.org/em/article/view/86983/81818

Supplementary Figure S6.

IQTree with bootstrap values.

Data type: PDF document

Link: https://www.biotaxa.org/em/article/view/86983/81819

Figures

Figure 2. Distribution map of subspecies of A. damone as determined by this study.

Figure 3. TCS haplotype network of COI barcodes for A. damone.

Fig. 4. Holotype ♂ and paratype ♀ of the new subspecies of A. damone described in this study. A–D) A. d. calabra ssp. nov., ♂♀ Italy: Calabria, Balzate, 39.164, 16.3548, 12.V.2015, leg. S. Scalercio (coll. CREA-FL); E–H) A. d. arachovae ssp. nov., ♂ Greece: Arachova 7.IV.1991 (leg. and coll. Back); ♀ Greece: Delphi, e.o. 25.III.1992, Egg 9.IV.1991 (leg. and coll. Back); I–L) A. d. morea ssp. nov., ♂♀ Greece: Taygetos, 23.IV.2001 (coll. Back); M– P) A. d. shahrdarii ssp. nov. ♂ Iran: Alborz, Karaj, Azgi Dareh, 1600 m, 25.IV.2008, leg. A. Naderi (coll. MMTT); ♀ Iran: Alborz, Chalus road, Dizin road, 2200-2600m, 22.V.2001, (leg. and coll. Naderi).