Synonymy of Pseudochazara misjai and Pseudochazara tisiphone dibra: taxonomic reassessment of an Albanian butterfly population.
Is the glass half full or half empty? An inconvenient truth.
Published online: 25.v.2025.
DOI: 10.5281/zenodo.15492634

Sylvain Cuvelier¹ | Mohammad AJ Marafi² | Anila Paparisto³
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¹ Diamantstraat 4, B-8900 Ieper, Belgium. sylvain.cuvelier@telenet.be
² Central Region Section, Department of Restoration of Terrestrial and Marine Ecosystems, Public Authority of Agriculture Affairs and Fish Resources (PAAFR), P. O. Box 21422, Safat, 13075, Kuwait City, State of Kuwait. Mohammadam@paaf.gov.kw
³ Faculty of Natural Sciences, Department of Biology, Bulevardi Zogu i Pare, Tirana University, Al-1001 Tiranë, Albania. anila.paparisto@unitir.edu.al

Abstract
A population of Pseudochazara butterflies from Albania was formally described as the subspecies Pseudochazara tisiphone dibra on January 24, 2025. However, in a separate publication on March 1, 2025, the same population was described as a distinct species, Pseudochazara misjai.
This paper reassesses the taxonomic status of the population, critically evaluating the evidence supporting both descriptions. Based on a careful review of morphological, climatic, ecological, and molecular data, we propose that Pseudochazara misjai is a junior synonym of Pseudochazara tisiphone dibra, which should remain the valid name for the population. This decision follows the guidelines of the International Code of Zoological Nomenclature (ICZN), prioritizing the earlier description to maintain nomenclatural stability.
The current study furthermore addresses inaccuracies and oversights in the publication of March 1, 2025, and reviews updates published between 2018 and January 9, 2025 on the dynamic website Fluturat e Shqipërisë, which includes a Monograph on Pseudochazara tisiphone.

Key words
Synonymy — Papilionoidea — Satyrinae — Pseudochazara — genetic diversity — DNA barcoding — morphological variability — phenotypic plasticity — reproductive isolation — ICZN — subspecies — ssp. nov. — Pseudochazara tisiphone dibra Cuvelier & Marafi, 2025 — Pseudochazara misjai Parmentier, 2025 — syn. nov.

Abbreviations
AL: androconium length from basal stalk (bs) to terminal points above apex (tp).
AB: androconium breadth across widest part of lamina (l).
Avg.: average.
bs: basal stalk.
°C: degrees Celsius.
d: days.
ESU: Evolutionary Significant Unit.
ICZN: International Code of Zoological Nomenclature.
l: lamina.
MORB: Mid-Ocean Ridge Basalt.
SSZ: Supra-Subduction Zone.
Tmax: mean daily maximum.
Tmin: mean daily minimum.
tp: terminal points at apex.

Introduction

The genus Pseudochazara de Lesse, 1951, includes a diverse group of butterflies with several species present in Europe, primarily in the Balkans, and one species in southern Spain.
The taxonomic classification of certain populations within the genus has often been debated, especially when new populations are discovered or existing ones are re-evaluated.
Brown [1981] proposed that the Pseudochazara populations on Mount Smolikas (Greece) represented an undescribed subspecies of Pseudochazara cingovskii (Gross, 1973), which he classified as ssp. tisiphone (url).
However, based on external morphology, Hesselbarth, van Oorschot, and Wagener (1995) reclassified P. cingovskii tisiphone as Pseudochazara mniszechii tisiphone (Brown, 1980) a change that was later confirmed by Wakeham-Dawson's (1997) discriminant analysis based on androconia within the genus Pseudochazara. Since then, this taxon has often been referred to as the subspecies tisiphone of the Turkish Pseudochazara mniszechii (Herrich-Schäffer, [1851]).
Takáts & Mølgaard (2016) and Verovnik & Wiemers (2016) employed DNA barcoding to address the taxonomic status of the described Pseudochazara species and to provide insights into the phylogeny of the genus. Takáts & Mølgaard (2016) maintained the status of P. mniszechii tisiphone due to the lack of samples of Turkish P. mniszechii. Based on their findings, Verovnik & Wiemers (2016) proposed a recognition of Pseudochazara tisiphone Brown, [1981] as a distinct species.
Multiple sources have reported the presence of P. tisiphone in the Korçë district (South East Albania) since the 1980s (Misja & Kurrizi 1984; Misja 1993; Misja 2005). Since these early publications the species was confirmed from many localities in the Korçë district and was recently also discovered in Gjirokastër county (Cuvelier & Paparisto 2025).
On 13.vii.2017, the first observations of a potentially disjunct Pseudochazara population were made near Bulqizë by S. Cuvelier (Cuvelier, Parmentier et al. 2018 and Suppl. Mat. 2), where the butterfly was found to be common across four visited localities.
On 18.vii.2017, L. Parmentier independently confirmed the occurrence of the population in the Bulqizë area and also observed it at a few kilometres near Krastë (Cuvelier, Parmentier et al. 2018 and Suppl. Mat. 2).
X. Qirinxhi, accompanied by R. Fero, reported their discovery of a population in the Lurë-Dejë National Park, which was made on 23-24 July 2022. The National Park is located more than 30 km north of Bulqizë and Krastë (Cuvelier et al. 2023).
Pseudochazara samples from Dibër County are included in the neighbor-joining tree presented in Dincă et al., Supplementary Data 14 (2021), and in Dapporto et al., Appendix S1 (2022). Samples are present in the analysis of the Atlas of mitochondrial diversity of Western Palearctic butterflies. Cuvelier (2023) provides a summary of the current status of this haplotype, concluding with the suggestion for an Evolutionary Significant Unit (ESU).
On January 24, 2025, a population of Pseudochazara from Bulqizë (Dibër district, Albania) was described in AWPL 2025(1) as the subspecies Pseudochazara tisiphone dibra Cuvelier & Marafi, 2025. However, in a subsequent article, an initial abstract was published in the Figshare repository (Figshare v1) on January 27, 2025, and in Phegea on March 1, 2025 (url). The full article was later deposited on March 2, 2025 (Figshare v2), with a second version likely appearing in Phegea on the same day or shortly thereafter. Across these four versions, the same population was described as a distinct species, Pseudochazara misjai Parmentier, 2025.
The primary objective of this study is to resolve the taxonomic ambiguity surrounding this Pseudochazara population.
Additionally, the goal is to clarify any potential confusions and inaccuracies regarding the second version (Figshare v2) from March 2, 2025, deposited in the repository, and to provide an overview of the evolution of updates (from 2018 to the present) in the P. tisiphone Monograph on the Fluturat e Shqipërisë website.

Materials and methods

The study population of Pseudochazara tisiphone dibra Cuvelier & Marafi, 2025 was sampled from Bulqizë, the type locality in Dibër County (Albania) and described in Cuvelier & Marafi (2025). Meanwhile, the population described as Pseudochazara misjai Parmentier, 2025 in Parmentier & Qirinxhi (2025) was sampled from Kräste, the type locality of P. misjai, as well as from Bulqizë and the Lurë-Dejë National Park, all located in Dibër County (Albania).
Both morphological and molecular data were used to assess the variability or distinctiveness of the population in comparison to P. tisiphone populations from Korçë County and Greece.
Specimens were identified based on standard taxonomic features, including external and genitalia morphology.
In addition, mtDNA sequencing (COI) was performed to examine potential genetic differentiation between P. tisiphone and the population described as P. tisiphone dibra and the population classified as P. misjai (Dincă et al., Supplementary Data 14 2021; Dapporto et al., Appendix S1 2022; Cuvelier 2023; Cuvelier & Marafi 2025 and Parmentier & Qirinxhi 2025).
Separately, a new DNA barcode dataset consisting of Pseudochazara cytochrome c oxidase subunit I (COI) sequences without ambiguous base calls published by Dincă et al., Supplementary Data 14 (2021) (78 sequences), and by Parmentier & Qirinxhi (2025) (9 sequences) was compiled. Chazara briseis (Linnaeus, 1764) and Chazara prieuri  (Pierret, 1837) were added as outgroup (one sequence each). This resulted in an alignment of 89 COI sequences that was used to build a neighbor-joining (NJ) tree (Supplementary material S4) in MEGA version X (Kumar et al. 2018), using pairwise deletion and 1000 bootstrap replicates.
A haplotype network was inferred for taxa belonging to the clade formed by Pseudochazara williamsi (Romei, 1927), Pseudochazara euxina (Kuznetsov, 1909), Pseudochazara mercurius (Staudinger, 1887), P. cingovskii, P. orestes, P. tisiphone and P. tisiphone dibra. Only COI sequences of 658 base pairs (bp) and having zero ambiguous base calls were retained, resulting in a dataset of 45 DNA barcodes, which were collapsed to unique haplotypes with TCS version 1.21 (Clement et al. 2000). The same program was used to construct a maximum parsimony haplotype network that had a 95% connection limit (Supplementary material S5). Supplementary material S6 includes a haplotype summary and list.

Comparisons are made using data from Cuvelier & Marafi 2025 and Parmentier & Qirinxhi 2025, in addition to previous publications on various aspects of P. tisiphone.
This study considers the following aspects to provide a comprehensive analysis of the two publications regarding the population(s), their traits and potential status:

1/ Do both publications refer to the same Pseudochazara population?

2/ Are there habitat differences between the disjunct populations in the Korçë and Dibër Counties?

3/ Are there climatic differences between the disjunct populations in the Korçë and Dibër Counties?

4/ What methods were used to analyse wing morphology, and what patterns of variability and statistically significant differences were identified?

5/ What data are available regarding androconial scales, and how do they compare with previous publications?

6/ What is the present status of morphological data pertaining to genital structures?

7/ What is the current extent and quality of the available genetic data?

8/ Addressing potential confusions and inaccuracies in the article deposited in the Figshare repository on March 2, 2025.

9/ Clarification of the publication timeline since Gross's 1973 description of P. sintenisi cingovskii, highlighting subsequent taxonomic revisions, including name changes and synonymies.

10/ Review of the literature on P. tisiphone in Albania, from its initial mention to the present.

11/ Review of the updates (2018 to the present) of the dynamic website Fluturat e Shqipërisë which features a Monograph on P. tisiphone.

Species names were selected according to the systematics outlined in the dynamic checklist of the Western Palearctic butterflies (Taymans & Cuvelier 2025).

Results

1/ Both publications refer to the same population
The DNA barcode data from both publications overlap through three identical specimens, RVcoll14U545, RVcoll14U546, and RVcoll14U547, which appear in each study.
The type locality of the holotype of P. tisiphone dibra is Bulqizë, while for P. misjai, it is Krastë. Both localities are situated in the Dibër district, approximately seven kilometres apart, and share similar ophiolitic habitat characteristics.

2/ Habitat differences between the disjunct populations in Korçë and Dibër Counties
The ophiolite complexes found in Dibër County and Korçë-Gjirokastër Counties are both part of the Tethyan Ophiolite Belt, a globally significant network of ophiolite sequences that stretches across several countries, formed by similar tectonic processes during the closure of the Tethys Ocean. In Albania, this tectonic activity, south of the Scutari-Pec line (Smith 1993), led to the formation of the Mirdita zone and is clearly divided into a western belt, with MORB-type ophiolites and an eastern belt with SSZ ophiolites (Çina 2016).
In the map of Albania from Çina (Fig. 1 2016), the three regions hosting Pseudochazara populations (Korçë- Gjirokastër Counties, the Bulqizë area and the Lurë area) are all located within the eastern belt, represented in darker blue, characterised by SSZ-type ophiolites. In Greece, a clear distinction cannot be made (Barth et al. 2008). The Pindos ophiolite in northwestern Greece includes both MORB-type and SSZ-type sequences (Smith 1993). The habitats of Greek Pseudochazara populations (Pamperis 2025) are characterised by typical ophiolitic rocks and are primarily, though not exclusively, associated with one type of ophiolite.
Bulqizë is particularly known for its rich chromite deposits, and mining activities, which are associated with the peridotitic rocks in the region. Depending on the mineral composition, the terrain exhibits a range of colours, from dark yellow-ochre to brown, with occasional subtle hues of dark green in certain areas.
Similarly, the ophiolites in southeastern Albania, such as those in Boboshtiçë, also contain significant mineral deposits, including chromite, copper and nickel, though their distribution and concentration may vary across the region. These mineral-rich deposits often give rise to distinct dark, metallic colours in the rocks, particularly in areas with high chromite content.

3/ Climatic differences between the disjunct populations in Korçë and Dibër Counties
To compare the macro-climate data with high-quality, historical records, Meteoblue and NOAA's Climate Data Online are excellent resources. Meteoblue is particularly user-friendly for accessing long-term trends while NOAA provides raw historical data for professional use, including climate studies. Meteoblue offers data at a 1x1 km coordinate resolution, whereas NOAA's Climate Data Online (CDO) provides climate data at various spatial resolutions, though it generally does not offer 1x1 km resolution for all datasets.
Therefore, Meteoblue was selected to compare two habitats with healthy Pseudochazara populations at altitudes as closely comparable as possible. The habitat near Bulqizë (41.44N 20.20E) is at 1061 m asl, while in Korçë County, the habitat near Boboshtiçë (40.54N 20.77E) is at 1134 m asl.
The elevation difference of 73 meters can be adjusted for using the lapse rate, which describes how temperature decreases with altitude: (73 m/ 1000) × 6.5°C = 0.4745°C. To compare Tmax at 1134 m to 1061 m, add 0.47°C to the 1134 m value.
While both Tmax and Tmin are affected by altitude, Tmin is generally more sensitive to changes in altitude due to the cooling of the air overnight.
Bulqizë has a continental climate, characterised by cold winters with snowfall and warm summers. Boboshtiçë has a Mediterranean-continental climate, with somewhat milder winters compared to Bulqizë, but still cold enough to see some snow. Summers are warmer in Boboshtiçë. Precipitation in Boboshtiçë is generally lower than in Bulqizë and summers are generally drier in Boboshtiçë compared to Bulqizë although occasional thunderstorms can still occur.
In Table 1, a summary of climate data between the two localities is presented, while the full data including graphics (Supplementary material S1) are accessible through AWPL online.
The definitions of the climate variables can be consulted on the Meteoblue website.

Table 1. Summary of climate data between the habitats in Bulqizë and Boboshtiçë.

Fig. 1. The wind speed analysis table is complemented by two wind rose diagrams from Meteoblue, illustrating wind patterns: Bulqizë (left) and Boboshtiçë (right).

4/ Wing morphology
Based on external characters and androconial scales, extracted from the wing of one specimen from Bulqizë, similar to the previously studied specimens (Cuvelier & Mølgaard 2015) from southeastern Albania, Cuvelier, Parmentier et al. (2018) assign the discovered population in Dibër County to P. tisiphone.
Cuvelier & Marafi (2025) give a qualitative description: “On average, both ♂♂ as ♀♀ of P. tisiphone from Dibër county appear smaller and paler, though without clear outliers, than specimens from Korçë county, and exhibit notable variability in the dimensions of the orange submarginal bands on the upperside of the hindwings."
The pronounced intraspecific variability of P. tisiphone in Korçë County was documented by Cuvelier & Mølgaard (2015) and was also observed in the population from Dibër County.
In Table 1B-C, Parmentier & Qirinxhi (2025) present a quantitative analysis of 9 specimens from unspecified localities in Dibër County and 10 specimens from other unspecified localities. Five wing measurements were selected for males: forewing length, two measurements of the forewing submarginal band width, hindwing submarginal band width, and the thickness on the hindwing at the marginal band. Five wing measurements were also selected for females: forewing length, forewing breadth of the darkened veins in the postdiscal band, two measurements of the forewing submarginal band width, and the forewing surface area of the orange zone at the end of the postdiscal band in cell S1a. For further details, see Fig. 1 and page 26 in Parmentier & Qirinxhi (2025). Univariate statistical tests on metric data of the traits, which show statistically significant differences in Fig. 1B-C, and a non-metric multivariate technique (NMDS), with plots presented in Fig. 6, were both applied.

5/Available data regarding androconial scales
The androconial scales of the Dibër population are illustrated in Cuvelier, Parmentier et al. (2018) and in Parmentier & Qirinxhi (2025).
In the 2018 publication, androconial scales from a single specimen, removed from two areas on the forewing, were photographed without the use of cover glass, using a calibrated 5 megapixel Dino-Lite AD- 7013MZT. These images, presented in Fig. 12c-d of Cuvelier, Parmentier et al. (2018), closely resemble the studied androconial scales of P. tisiphone specimens from southeastern Albania, as illustrated in Cuvelier & Mølgaard (2015). Comparing the androconial scales of this Dibër specimen with the figures of P. tisiphone androconial scales in Wakeham-Dawson (1997), reproduced hereunder in Fig. 2, with the kind permission of A. Wakeham-Dawson, and Fig. 1 in Wakeham-Dawson & Kudrna (2006) show a similar shape matching best with group 4 androconial scales as described in Gross (1978) and Wakeham-Dawson (1997).

Fig. 2. Androconial scales of Pseudochazara, used with the kind permission of A. Wakeham-Dawson (Source: Wakeham-Dawson 1997).

In Cuvelier, Parmentier et al. (2018), the androconium length (AL) measured from the basal stalk to the terminal points above the apex, was recorded for three androconial scales: 0.289, 0.292 from one area, and 0.328 mm from the second area. The androconium breadth across the widest part of the lamina (AB) for two androconial scales from the two areas was 0.029 mm and 0.032 mm, respectively. After considering the significant variation in the scales removed from the two areas of the forewing, the AL/AB ratio for a single specimen seemed less meaningful for analysis.
In Parmentier & Qirinxhi (2025), androconial scales from 9 specimens of the Dibër population were photographed under cover-slips, using a calibrated Dino-Lite Edge Microscope, model AM7915MZT, and compared to scales from 10 specimens of “known P. tisiphone populations”.
Regarding the group of androconial scales as described in Gross (1978) the Dibër population is classified as follow: “Classified according to the list of Gross (1978), the ratio can be attributed to class 1–2.”
The AL ranges from 0,291 to 0,312 mm, with an average of 0,3 mm. The AB ranges from 0,020 to 0,024 mm, with an average 0.022 mm. The given mean calculated ratio AL/AB: 13,82 (Supplementary material S1 in Parmentier & Qirinxhi 2025). Univariate statistics are summarised in Tables 1A. The data for the individual specimens are provided in Supplementary material S3 of Parmentier & Qirinxhi (2025).

6/ Present status of morphological data pertaining to genital structures
In Cuvelier & Marafi (2025), it was noted that major differences in genitalia could suggest potential reproductive isolation. However, the genitalia of closely related Pseudochazara species are remarkably similar, offering limited diagnostic value. The genitalia of the two disjunct P. tisiphone populations were not compared, as the likelihood of finding distinct differences is minimal but recommended further research on male and female genitalia.
Parmentier & Qirinxhi (2025) partially addressed this issue through a limited study of the male genitalia of the two populations. The number of specimens studied is unclear, though the illustrations include two specimens from Krastë (Dibër County, Albania), one specimen from Distrato (Greece), and one specimen from Gjergjeviçë (Korçë County, Albania).

7/ Current extent and quality of the available genetic data
Until now, studies on Pseudochazara species have primarily relied on COI barcoding, which, while valuable, does not encompass the full mitochondrial genome. Takáts & Mølgaard (2016), Verovnik & Wiemers (2016), Dincă et al., Supplementary Data 14 (2021), Dapporto et al., Appendix S1 (2022), Parmentier & Qirinxhi (2025) together with the phylogenetic tree presented in Supplementary material S4 and haplotype network shown in Supplementary material S5 of this study.
Furthermore, data from single nuclear genes, as well as larger studies utilizing high-throughput sequencing or full genome sequencing, are lacking.
The DNA barcode sequencing results of three specimens (RVcoll14U545, RVcoll14U546, and RVcoll14U547) are included in both Cuvelier & Marafi (2025) and Parmentier & Qirinxhi (2025). In Parmentier & Qirinxhi (2025) new COI sequences, deposited in GenBank, with accession numbers are mentioned.
A phylogenetic tree of Pseudochazara species based solely on complete COI sequences (658 bp), together with the haplotype network for the Pseudochazara group to which P. tisiphone belongs, is presented in Supplementary material S4 and Supplementary material S5, and further explored in the Discussion chapter.

8/ Addressing potential confusions and inaccuracies in the article deposited in the Figshare repository on March 2, 2025
The publication by Parmentier & Qirinxhi (2025) contains editorial oversights, inconsistencies, and inaccuracies that may lead to misinterpretations of the facts.
These issues are addressed and, where possible, clarified in the discussion section of the present publication.

9/ Publication timeline since Gross's 1973 description of Pseudochazara sintenisi cingovskii, including subsequent taxonomic changes and synonymies
Gross (1973) is the source of the original taxonomic description of Pseudochazara sintenisi cingovskii.

Koutsaftikis (1974) reported Pseudochazara sintenisi Stgr. from Greece for the first time. The figure of a female specimen in this publication clearly depicts an individual that was later associated with tisiphone.

Brown (1976) established P. cingovskii Gross as a new status, with the holotype described from Mt. Smolikas.

Gross (1978) discussed P. cingovskii from Yugoslav and Greek Macedonia. He did not assign the Greek population to any subspecies. The publication includes an illustration of the androconial cell of P. cingovskii from Prilep (present-day North Macedonia).

Brown [1981] described P. cingovskii tisiphone as a new subspecies.

De Prins & van der Poorten (1981) described a new species of Pseudochazara from the Drama district (northeastern Greece): Pseudochazara orestes De Prins & van der Poorten, 1981.

Hesselbarth, van Oorschot & Wagener (1995) reclassified P. cingovskii tisiphone as Pseudochazara mniszechii tisiphone stat. nov., based on a single population near Bursa (Turkey) and from Greece.

Wakeham-Dawson (1997) reclassified P. cingovskiii tisiphone as Pseudochazara mniszechii tisiphone stat. nov., independent of Hesselbarth et al. (1995).

This view is also supported by Lukhtanov (2007) in Nymphalidae: Satyrinae (Global Butterfly Names Project).

This opinion, that P. mniszechii tisiphone is the valid taxon, is also supported in Takáts & Mølgaard (2016) in their study on partial mtCOI-sequences of Balkanic species of Pseudochazara, where COI barcoding was first used to confirm the classification.

Verovnik & Wiemers (2016), based on a deep genetic split between P. mniszechii and P. mniszechii tisiphone, elevated the latter to species level.

Cuvelier, Parmentier, Paparisto & Couckuyt (2018) reported, for the first time, a new disjunct population of P. tisiphone in Dibër County.

Dincă V., Dapporto L., Somervuo P., Vodă R., Cuvelier S., Gascoigne-Pees M., Huemer P., Mutanen M., Hebert P. & Vila R. (2021) include P. tisiphone from Dibër county in the phylogentic tree.

Dapporto, Menchetti, Vodă, Corbella, Cuvelier, Djemadi., Gascoigne-Pees, Hinojosa, Lam, Serracanta, Talavera., Dincă & Vila R. (2022) documented COI gene diversity in Balkan Pseudochazara as part of The Atlas of Mitochondrial Diversity of Western Palearctic Butterflies.

Cuvelier (2023) provided further commentary on the IGV Atlas regarding P. tisiphone and designated the population in Dibër County as a distinct ESU.

Cuvelier & Marafi (2025) assigned the population in Dibër County subspecies status as Pseudochazara tisiphone dibra ssp. nov. This new subspecies was registered in ZooBank.

Shortly after the subspecies status was assigned in Cuvelier & Marafi (2025), Parmentier & Qirinxhi (2025) elevated the population in Dibër County to species level, naming it Pseudochazara misjai sp. nov. This new species was registered in ZooBank.

10/ Review of the literature on P. tisiphone in Albania
Multiple sources have reported the presence of P. tisiphone in the Korçë district (South East Albania) since the 1980s: Misja & Kurrizi (1984), Misja (1993); Misja (2005), Eckweiler (2012), Cuvelier & Mølgaard (2015), Šašić et al. (2015), Bjerregård et al. (2023) and yearly updates in Fluturat e Shqipërisë.
On July 13, 2017, S. Cuvelier made the first observations of a potentially disjunct Pseudochazara population in Dibër County, near Bulqizë. (Cuvelier, Parmentier et al. 2018 and Suppl. Mat. 2) in the company of an entomologist, J. Couckuyt, experiencing Balkan butterflies for the first time.
The butterfly was found to be common across four visited localities in the area.
On July 18, 2017, L. Parmentier independently found the population in the Bulqizë area and also observed it at a few kilometres away near Krastë (Cuvelier, Parmentier et al. 2018 and Suppl. Mat. 2).
On July 23-24, 2022, X. Qirinxhi, accompanied by R. Fero, discovered a P. tisiphone population in Lurë-Dejë National Park, located more than 30 km north of both Bulqizë and Krastë. However, concerns were raised by Parmentier regarding a potential confusion with a collecting locality in Korçë County. Despite these initial doubts about the exact location, Qirinxhi's observation was confirmed by R. Fero and subsequently included in Cuvelier, Parmentier et al. (2023).
P. tisiphone was also recently discovered in Gjirokastër County (Cuvelier & Paparisto 2025).
In Parmentier & Qirinxhi (2025), the Lurë locality is confirmed by the first author. The map does not show a significant extension of the known localities in Dibër County, although the publication itself provides no details regarding the exact locations.

11/ Assessment of recent updates (2018–present) to Fluturat e Shqipërisë and its P. tisiphone Monograph
Thanks to the excellent collaboration with Theo Garrevoet, former president of the Flemish Entomological Society (FES), and the Grupi i Punës Fluturat Universiteti i Tiranës, a preliminary version of the Fluturat e Shqipërisë website was being developed.
Following the publication of Cuvelier, Parmentier et al. (2018) the distribution map in the Atlas and Monograph of P. tisiphone included the localities near Bulqizë and Krastë, as well as additional localities in Korçë County.
In 2023, before the development on the FES server was halted by a decision of two FES committee members, the map in the Atlas and Monograph of P. tisiphone were updated to include the localities in the Lurë-Dejë National Park, provided by X. Qirinxhi. These updates remained unchanged at the official launch of the website on the University of Tirana's server on October 30, 2023.
On January 09, 2025 a major update was implemented on the website for the Atlas, and a note was added to the Monograph about an upcoming publication on the separation of a subspecies of P. tisiphone, named P. tisiphone dibra after the County. Additionally, the legends in the Monograph for the photographs in Figs. 1c, e, f, g, h, and j-k were updated to reflect P. tisiphone dibra.
The recent discovery of a population in Gjirokastër County, marks the first record in the area and is the most significant update to the distribution map of P. tisiphone dates also from January 09, 2025.
On January 27, 2025, the P. tisiphone Monograph and the Sources section in Fluturat e Shqipërisë were updated to include the publication by Cuvelier & Marafi (2025). On March 2, 2025, the Monograph and Sources were further revised to incorporate the work of Parmentier & Qirinxhi (2025).
The current version of the Atlas, dated January 9, 2025, has been uploaded to the Zenodo repository, under the title Fluturat e Shqipërisë. Atlas 09.i.2025. The publication date is March 28, 2025, and future updates will also be made available there.

Discussion

1/ Both publications refer to the same population
This population is regarded as the same Pseudochazara population, supported by the inclusion of three identical specimens for DNA barcode analysis. The DNA barcode sequencing results of three specimens (RVcoll14U545, RVcoll14U546, and RVcoll14U547) are included in both Cuvelier & Marafi (2025) and Parmentier & Qirinxhi (2025).
Similar habitats are present throughout the eastern Mirdita massif, with intermediate localities remaining unexplored due to the difficult access to these areas. As a result, the potential connectivity between the Dibër populations and those in Korcë-Gjirokastër Counties remains undocumented.
These areas, characterised by comparable ecological conditions, offer an ideal environment for Pseudochazara populations, further supporting the potential connectivity between these populations.
The consistent flight period across the disjunct areas also suggests synchrony in the species' life cycle, which may facilitate gene flow and population stability.

2/ Habitat differences between the disjunct populations in Korçë and Dibër counties
The three disjunct Albanian areas with Pseudochazara populations (Korçë- Gjirokastër, Bulqizë and Lurë) are all situated in the eastern belt with SSZ-type ophiolites (Çina 2016). The Greek Pindos ophiolite comprises both MORB-type and SSZ-type sequences (Smith 1993). The habitats of Greek Pseudochazara populations, based on the first author’s observations and as confirmed by L. Pamperis (personal communication), are dominated by typical ophiolitic rocks and are found mainly, though not exclusively, on SSZ ophiolites, as shown in the latest distribution map (url).
The ophiolites in the different areas host significant mineral deposits, including chromite, copper and nickel, though their distribution and concentration may vary. These mineral deposits give rise to a range of metallic brown hues in the rocks, which become progressively darker as the concentrations of chromite, serpentine, and other iron- and magnesium-rich minerals increase and go hand in hand with subtle colour differences in many Satyrinae species, as also documented in Pseudochazara, among others, by Weiss (1980), Hesselbarth, van Oorschot & Wagener (1995) & Gil-T (2017).
The Dibër, Korçë-Gjirokastër, and Pindos ophiolite regions are situated in rugged, mountainous terrain, contributing to their dramatic landscapes. Characterised by steep slopes, deep valleys, and varying elevations, these areas influence the local microclimates, ecosystems, and vegetation. The ophiolitic soils found here are typically nutrient-poor, and the predominantly south-facing slopes, combined with sparse vegetation, create harsh environmental conditions. In response, the local flora has developed various adaptations to thrive in rocky soils, withstand droughts, and endure extreme temperatures.

3/ Climatic differences
The comparison of weather variables in Meteoblue between Bulqizë and Boboshtiçë shows that despite the relatively small difference in elevation, Boboshtiçë tends to experience warmer temperatures, more sunny days, significantly less precipitation (mm) and precipitation days, similar amount of snow days and higher wind speeds than Bulqizë.
Bulqizë, on the other hand, has more frequent precipitation, especially in heavier amounts, and experiences colder nights and more frost days. The climate at Bulqizë is characterised by more frequent rainfall and cooler temperatures, indicating a tendency to wetter and colder weather patterns compared to Boboshtiçë.
The differences in temperature, precipitation, solar radiation, and wind between the two locations persist throughout the year and can therefore significantly influence development during both the early summer stages following egg-laying and the spring period when caterpillars progress through their larval stages to pupation.
This information could be valuable for environmental studies requiring a detailed understanding of local weather conditions at these two locations, particularly those focusing on Satyrinae. Previous studies (Dennis & Shreeve 1989; Gibbs et al. 2010) have demonstrated that climatic variables, such as temperature and precipitation, can drive significant morphological changes in wing size, body shape, as well as wing patterns and colouration. Understanding how these factors influence Pseudochazara morphology in different climates can offer valuable insights into phenotypic plasticity and climate-driven adaptation for this genus.

4/ Wing morphology
It is important that the apparent external morphological differences are substantiated, as done in Parmentier & Qirinxhi (2025). However, it is first noteworthy that the Material and methods chapter does not include a description of how the wing measurements were carried out. Secondly, it should be noted that the sample sizes for the two disjunct Pseudochazara populations, nine and ten individuals respectively, are relatively small. These sample sizes are not as large as would be ideal for capturing the full range of variability and drawing more robust conclusions.
Larger sample sizes would provide a more thorough representation of population variability and enhance the reliability of the findings.
As noted in Cuvelier & Mølgaard (2015), the sample size of 38 males and 19 females used to compare two disjunct populations of Pseudochazara amymone Brown, 1976, is suboptimal for such highly variable butterfly populations. For a study investigating differences in 5 wing variables, one would likely need around 50-60 individuals per population for a t-test (per trait) and around 70-80 individuals per population if using multivariate analysis to account for all wing traits simultaneously.
A larger sample size would certainly provide a more statistically robust description of the variability, although it is expected that the overall impression of slightly smaller individuals with a brighter appearance would not be compromised.
Additionally, the lack of a table indicating the localities of the individual specimens should be addressed in the future.
In the introduction to the genus Pseudochazara in Hesselbarth, van Oorschot & Wagener (1995), it is noted that the very similar appearance of adults across many populations and the significant individual variability within populations, makes distinguishing between species and subspecies extremely challenging. They also raise the question of to what extent homoplasy plays a role in this difficulty.
Several authors have explored the potential causes of the significant phenotypic plasticity observed in Pseudochazara species, suggesting that it may be linked to the butterflies' specific ecological needs (Weiss 1980; Tennent 1996; Hesselbarth, van Oorschot & Wagener 1995; Tolman & Lewington 1997; Verovnik & Wiemers 2016; Gil-T 2017).
Most studies primarily focus on the colouration and wing patterns of the butterflies' upper- and undersides in relation to the substrate they inhabit, particularly establishing how this colouration provides an advantage in camouflage or mate attraction across varying environmental conditions. Based on differences in external morphology among Pseudochazara populations and a very small genetic mitochondrial divergence, authors tend to adopt a range of statuses, from the very cautious ecotype, variant, form, subspecies to the more speculative species (Takáts & Mølgaard (2016); Verovnik & Wiemers 2016; Gil-T 2017).
The colour differences and wing patterns observed in the two disjunct Pseudochazara populations, from NW Greece to SE Albania and from Dibër County, show a modest correlation with the subtle colour variations of the ophiolitic substrates in these areas, which are potentially linked to differences in mineral content, including chromite, copper and other trace elements.
To the best of our knowledge, no studies within the genus Pseudochazara or among Satyrinae of the Balkan Peninsula have investigated a substantiated relationship with climatic data.
However, various studies on Satyrinae in Western Europe have shown that climatic factors can significantly influence morphological traits, a relationship that is supported by significant statistical results, in multi-brooded species such as Pararge aegeria, Coenonympha pamphilus and Lasiommata megera, as well as in the single-brooded Hipparchia semele (Dennis & Shreeve 1989; Gibbs et al. 2010).
The most relevant proxy for Pseudochazara in relation to the preference for early successional habitats with bare ground, short herbs and grasses, along with a univoltine life cycle, is H. semele. Its remarkable ecological adaptability is evidenced by the description of no fewer than six subspecies from the British Isles.
In Dennis & Shreeve (1989), changes in wing morphology of H. semele across Britain and Ireland highlight differences in wingspan, the width of the medial band, colour, brightness, and black suffusion. Significant correlations with factors such as the latitudinal gradient, sunshine hours, Tmax, Tmean, total precipitation, humidity, and wind speed enabled the development of a model to explain the geographical variation. The phenetic differences observed in species living in open habitats were significantly greater than those in species living in closed habitats.
Thermoregulation plays a key role in driving phenotypic variation. Cooler, windier, and cloudier conditions slow down biological processes, which in turn affect reproductive success. Under these conditions, specific phenotypic modifications can provide a distinct advantage. Melanism on the underside in lateral baskers, smaller wing size, along with brighter colouration, may help compensate for reduced activity time by improving predator avoidance and enhancing courtship displays.
This is exactly what can be observed in the Pseudochazara populations from NW Greece and SE Albania and Dibër County, where these phenotypic traits appear to be adapted to the specific climatic conditions (Table 1; Supplementary material S1) in these two regions.
The advantageous morphology observed in the Dibër County population, which is adapted to cooler and less sunny conditions, likely results from natural selection favoring traits that enhance reproductive success in this specific area. This selection may be linked to subtle genetic differences that contribute to the expression of these adaptive traits.

5/Androconial scales
A major publication, investigating the structure of androconia in 23 Pseudochazara populations, spanning taxa at both the species and subspecies levels, Wakeham-Dawson (1997), has been overseen in recent publications reviewing the taxonomy of this genus (Takáts & Mølgaard 2016; Verovnik & Wiemers 2016).
There were significant differences in AL and AB between the taxa summarised in Table 1, reproduced hereunder in Fig. 3, with the kind permission of A. Wakeham-Dawson.
This publication is particularly relevant for P. cingovskii, P. orestes and P. tisiphone, as it provides valuable insights into the similarities in the structure of their androconial scales. A comparison (Table 2 in Wakeham-Dawson 1997) between the actual group memberships and the predicted species affiliations revealed that androconial traits could not reliably distinguish ten of the twenty-three species, including cingovskii (for which none of the specimens matched the predicted classification), tisiphone, and others.
It is somewhat surprising that this important publication is not referenced in Parmentier & Qirinxhi (2025) in their discussion of P. tisiphone populations, as well as those of closely related Pseudochazara species in the Balkan Peninsula. Given the relevance of the study to the structural variations in androconial scales, one would expect its findings to be referenced in relation to the new population, especially considering the critical role androconia may play in understanding the evolutionary and taxonomic relationships within the genus. The omission of such a key reference may limit a more comprehensive analysis of the morphological traits that define these populations and their interrelations with other Pseudochazara species in the area.

Fig. 3. AL and AB for 23 Pseudochazara taxa, used with the kind permission of A. Wakeham-Dawson (Wakeham-Dawson 1997).

At first glance, the data for the Dibër population presented in Cuvelier, Parmentier et al. (2018) and Parmentier & Qirinxhi (2025) suggest minimal variation in AL. However, there may be some indication of differences in AB, as well as in the classification based on Gross (1978), as suggested in Parmentier & Qirinxhi (2025).

Did previous publications also document differences in the measurements of AL, AB, the AL/AB ratio and in the Gross classification?
Comparing measurements (Supplementary material S3) for P. cingovskii, P. orestes, and P. mniszechii in Gross (1978), De Prins & van der Poorten (1981), Wakeham-Dawson (1997) and Parmentier & Qirinxhi (2025) reveals significant differences in AL, AB, which consequently result in notable variations in the AL/AB ratio. For P. cingovskii the maximal differences are: AL (4,8%), AB (30,4%), and AL/AB ratio (24,5%). For P. orestes, the maximal differences are: AL (7%), AB (45,8%), and AL/AB ratio (36,3%). Finally for P. mniszechii the maximal differences are: AL (12, 1%), AB (30,4%), and AL/AB ratio (16,4%). The most pronounced differences are observed in AB (see further discussion below) and are considerable.
In several publications of Wakeham-Dawson (1997), Wakeham-Dawson & Kudrna (2000), Wakeham-Dawson (2006) and Wakeham-Dawson et al. (2007), it is clearly marked that measurements include the basal stalk (bs). However, in Parmentier & Qirinxhi (2025), the bs is not visible, and based on the markings in their figures and their Supplementary Material S1, it appears that the bs was not included in the measurement of AL. Although it may be a minor discrepancy, it complicates the comparison with the standard measurements.
Comparing measurements (Supplementary material S3) for P. tisiphone in Wakeham-Dawson (1997), Wakeham-Dawson (2006), and Parmentier & Qirinxhi (2025) also reveals significant differences in AL (9,4%), AB (8%), which consequently result in notable variations in the AL/AB ratio (18,9%). In Parmentier & Qirinxhi (2025), as expected, a shorter AL (likely excluding the bs) is observed, but surprisingly, a broader AB is reported. This finding is puzzling when compared to the data from Wakeham-Dawson (1997) and Wakeham-Dawson (2006), a leading expert on the androconia of Satyrinae and Pseudochazara.
Direct evidence for a correlation between wing size and androconial size for Pseudochazara is not known to us. Based on the low sample size, the impression from Parmentier & Qirinxhi (2025) is that the forewing size in males from the Dibër population may be significantly smaller compared to those from more southern populations.
Although borderline and not statistically significant, the AL of the androconia from the two populations appears to correlate with wing size.
It can be inferred through a proxy, which may provide some insight into this matter, as discussed in Wakeham-Dawson & Dennis (2004): “The apparent direct correlations between overall H. semele adult size and androconial size probably discounts androconia as a useful taxonomic character in British H. semele populations.” This should be confirmed by a study with an adequate sample size for Pseudochazara populations both within species and across different species.
Comparing the Gross classification between De Prins & van der Poorten (1981) and Wakeham-Dawson (1997) for the same species yields the following Gross types: P. cingovskii is classified as type 1-2 in De Prins & van der Poorten (1981) while in Wakeham-Dawson (1997), it is classified as type 4. For P. orestes and P. mniszechii, the classification is type 5 and type 4, respectively. This is also evident when comparing the classification of P. tisiphone in Wakeham-Dawson (2006), where it is classified as type 4, with that in Parmentier & Qirinxhi (2025), where it is classified as type 5.

Could such differences be due to varying interpretations of the classification by Gross (1978)?
Could this variation be attributed to the removal of androconial scales from different areas of the forewings, or is it due to the presence of different types of androconial cells on the wings?
Since the androconial scales shown in Fig. 12c and 12d of Cuvelier, Parmentier et al. (2018) were removed by the first author from different zones of the forewing, it is not entirely surprising to observe size variations across different wing compartments, as has been documented for other types of wing scales in butterflies (Kusaba & Otaki 2009; Dhungel & Otaki 2014), where scale size decreases regularly from the postbasal to the distal areas of the forewing.
Table 1, in Wakeham-Dawson (1997), reproduced here in Fig. 3, provides details regarding the base of the androconial cells in 23 Pseudochazara taxa, including P. tisiphone. For P. tisiphone, the bases are predominantly flat, though occasionally rounded, with noticeable variability both within specimens and across populations. In this publication, polymorphic androconia were observed within specimens and populations of ten species, including P. tisiphone. Due to this variability, androconia could not be reliably used for species identification, with a probability of 0.44 for P. tisiphone.

Did the use or absence of cover-slips impact the image quality and, consequently, the accuracy of the measurements?
Could the differences be due to variations in resolution or illumination between the two digital microscopes?
The resolution of Fig. 12c-d in Cuvelier, Parmentier et al. (2018) and Fig. 4, as well as in Supplementary material S1 in Parmentier & Qirinxhi (2025), is noticeably different.
In Cuvelier, Parmentier et al. (2018) the bs is clearly visible in various androconial cells, with a sharply delined base. The lamina is well contrasted, gradually becoming clearer towards the visible apex (the thin length of androconium between the thicker part of the lamina and the terminal points). In the figures of Parmentier & Qirinxhi (2025), the base is vague, the bs is not visible, and it seems excluded in the measurement of the AL as well as in the Supplementary material S1. Additionally, the apex is hardly visible, and the borders of the androconial scales are poorly contrasted with the background.
The measurement lines for AL in Parmentier & Qirinxhi (2025) suggest that the measurements begin at the base of the androconial cell, excluding the bs.
The difference in the AL/AB ratio for P. tisiphone between Wakeham-Dawson (2006) (15.34 ± 2.46) and Parmentier & Qirinxhi (2025) (12.26 ± 0.013) is striking.

Could this suggest an underestimation of the measured AB in Parmentier & Qirinxhi (2025) due to the image quality?
The limited resolution could indeed cause a negative measurement deviation, having a more significant impact on the smaller structures and measurements, such as AB (see also the discussion above).

Even more notable is the difference in classification according to the Gross’s criteria (1978) between Cuvelier, Parmentier et al. (2018) and Parmentier & Qirinxhi (2025). In Cuvelier, Parmentier et al. (2018) the base of the androconial cells is straight and the edges of the androconial cell in the first third of the lamina are parallel, rather than gradually tapering as they do from the base, ending in a short apex. These cells most closely correspond to types 4-5 as described in Gross (1978, p. 99) and type 4 in Wakeham-Dawson (1997). In the same publication (Fig. 4) the androconia of P. tisiphone are clustered together with those of the closely related P. cingovskii and P. orestes.
In Parmentier & Qirinxhi (2025), the overall structure of the lamina appears similar, it does not seem to gradually taper from the base to the apex, although the lower resolution may limit the clarity of this observation. The less defined base and apex give a limited impression that the base is neither rounded nor bulbous (as is typical of class 7 scales), and the apex appears shorter than the lamina. These characteristics, do not support classifying the androconial scales as class 1-2. The lamina is also not spindle-shaped, a typical characteristic of class 3 and 6 androconial scales, which is absent in this case.

For a comprehensive comparative overview of the data from previous publications alongside the current study's findings on the androconial scales of the Dibër population, please refer to Supplementary material S3.
Even without delving further into the discordant results, the central question that remains unresolved since it was first raised by Wakeham-Dawson & Kudrna (2000), is the relationship between androconial shape associated with different pheromone production, pheromone chemical structure, and breeding relationships within the genus Pseudochazara and how these factors may be linked to reproductive isolation between taxa. As a result, androconia cannot yet be reliably used as indicators of biological relationships at the species level, and this lack of clarity provides no conclusive information regarding a potential prezygotic barrier.

6/Genitalia
Parmentier & Qirinxhi (2025) confirm that the study of the male genitalia, could not serve for differential detailed morphometrics, which is in contrast to the emphasis placed on more pronounced features at the tip of the valva and the ventral structure of the uncus.
This suggests that genitalia do not act as a mechanical prezygotic barrier to reproductive isolation.

7/ Genetic data
Currently, only the mitochondrial COI gene, has been studied in various publications (Takáts & Mølgaard 2016; Verovnik & Wiemers 2016; Dincă et al., Supplementary Data 14 2021; Dapporto et al., Appendix S1 2022; Parmentier & Qirinxhi 2025). DNA barcodes have been successfully used in Lepidoptera taxonomy, but their limitations, as stated in Verovnik & Wiemers (2016), must be considered when interpreting gene trees. When the COI gene is the sole available data, it may not be sufficient to conclusively determine monophyly.
The COI gene, being mitochondrial and maternally inherited, cannot resolve postzygotic reproductive isolation, which is essential for understanding speciation. Since it only reflects the maternal lineage, it provides no insight into male genetic contributions, limiting its ability to detect barriers like hybrid sterility or inviability.
Given the minimal genetic distance in DNA barcodes among Pseudochazara populations on the Balkan Peninsula, Takáts & Mølgaard (2016) questioned where the boundary lies between species and subspecies in the studied Pseudochazara.
Verovnik & Wiemers (2016) confirm the low level of genetic differentiation between P. tisiphone, P. orestes, and P. cingovskii indicating a relatively recent split of the populations. Indeed, for a population with one generation per year, approximately one mutation is expected every one hundred thousand years.
Verovnik & Wiemers (2016) tend to support the separate species status based on constant differences in wing patterns/colouration. However, they also state in their introduction, “The main reason for the extensive variation in phenotype can be linked with the specific ecological requirements of these butterflies.” yet they do not further elaborate on this point.
As a second supporting element they also refer to their ecological specialization. The different substrates used by these three closely related taxa may be important. However, different substrates (limestone or ophiolite) do not account for the separation of all currently accepted Pseudochazara species. In the Balkan Peninsula, this is evident in the case of Pseudochazara graeca (Staudinger, 1870) and Pseudochazara amalthea (Frivaldszky, 1845).
For P. cingovskii and P. orestes, the distinction between the two might be influenced by their occurrence in limestone areas situated in disjunct massifs with different geological origins. The Pletvar pass, home to P. cingovskii, is located in the Pelagonian zone, while the Oros Phalakro area and nearby massifs, where P. orestes is found, belong to the Serbo-Macedonian Massif. Both regions have distinct geological histories shaped by separate orogenic events and tectonic processes over millions of years. The recent genetic differentiation in the COI gene of both limestone populations contrasts with their distinct geological histories spanning millions of years. However, when compared to the genetically similar P. tisiphone, which exclusively inhabits ophiolite environments, this appears to be a plausible hypothesis.
The discovery of a second genetically distinct Pseudochazara population associated with ophiolite habitats (Dincă et al., Supplementary Data 14 2021; Dapporto et al., Appendix S1 2022; Cuvelier 2023; Cuvelier & Marafi 2025; Parmentier & Qirinxhi 2025), located in separate massifs within the same Tethyan Ophiolite Belt, which was formed by similar tectonic processes during the closure of the Tethys Ocean, calls for a thorough reevaluation of this argument.
The first DNA barcodes of the Pseudochazara population from Dibër County were already presented in Dincă et al., Supplementary Data 14 (2021), as part of the COI-based neigbor-joining tree included in the study.
In contrast, Parmentier & Qirinxhi (2025) present a phylogenetic tree based on “a selection of samples (males and females)”, comprising 62 sequences retrieved from GenBank and 9 newly generated from personal material. The sequences vary in length (618 to 658 bp). The inclusion of shorter COI barcodes may reduce accuracy due to fewer informative sites and potential alignment issues. Although the relative placement of the different Pseudochazara populations remains unchanged, it is nonetheless unexpected that publicly available full-length sequences, were not incorporated into their analysis.
Supplementary material S4 presents a new phylogenetic tree, aligning 87 sequences of Pseudochazara species incorporating, among others, two distinctive P. cingovskii haplotypes (EULEP1567-15 P. cingovskii RVcoll-14-G750 and HBOK048-08), which differ by five mutations from other P. cingovskii specimens from the same locality. Supplementary material S5 presents the haplotype network, based solely on complete COI sequences (658 bp), comprising 45 DNA barcodes from the Pseudochazara group, including P. tisiphone. The analysis reveals limited resolution and an ambiguously defined genetic structure.
Mitonuclear discordance, where mitochondrial DNA and nuclear DNA provide conflicting phylogenetic signals, has been documented in multiple Palearctic Lepidoptera families (Desprès 2019; Dincă et al. 2019; Hinojosa et al. 2019; Ebdon et al. 2020; Joshi et al. 2024). Given the low resolution of DNA barcoding and the poorly resolved genetic structure in this Pseudochazara group, this is even more significant.
A study including larger genomic data (such as high-throughput or full genome sequencing), which allows the detection of gene flow across multiple loci, could help determine whether the observed mutations in the COI gene are isolated or part of a broader pattern of divergence across the genome. This approach would provide the potential to document gene flow, providing deeper insights into whether the observed phenotype is driven by genetic changes, such as speciation, or by environmental factors influencing phenotypic plasticity.

8/ Confusions and inaccuracies in Parmentier & Qirinxhi (2025), deposited in the Figshare repository on March 2, 2025
This chapter addresses the discrepancies and inaccuracies in Parmentier & Qirinxhi (2025), deposited in the Figshare repository on March 2, 2025.
Several issues have been noted, stemming from minor oversights by the author(s), chief editor, and three anonymous reviewers, particularly in the handling of information and names. Notably, there are a few location errors that need to be corrected, as well as misreported dates that could lead to confusion regarding the population’s discovery and historical context. A reference is cited multiple times with conclusions that do not align with the content of the article. A map from a reference appears to have been misinterpreted in relation to the Albanian ophiolitic formations. Additionally, one of the references cited in the article appears to include part of another citation, likely related to a chief editor’s work on African butterflies, even though the main content of the cited article concerns only Western Palearctic butterflies. This misattribution may lead to confusion about the geographic relevance of the research.
These discrepancies suggest an opportunity to enhance the internal review process within FES, building on the suggestion made at the FES annual members' meeting on January 27, 2024 (available on request), to revise the review procedure. Involving independent content reviewers, as proposed, could help ensure greater accuracy and consistency in future assessments. This chapter identifies and provides suggestions for rectifying these issues to contribute to an updated and more reliable version of the paper.

A. Co-authorship discrepancies and naming issues in the species description
In Parmentier & Qirinxhi (2025), X. Qirinxhi is listed as a co-author but is not mentioned in the description of the species name. This omission appears unusual and may indicate an oversight or miscommunication. Such discrepancies raise concerns about the clarity of authorship and contributions. It seems that Qirinxhi's role in the work was not fully acknowledged, potentially leading to confusion about her involvement.
Upon addressing these inconsistencies, X. Qirinxhi, a member of the Grupi i punës Fluturat Universiteti i Tiranës was contacted to clarify her contribution. She confirmed that her involvement was limited to fieldwork conducted in 2022, and that she did not participate in the analyses, manuscript drafting, revision of subsequent drafts, or the formulation of conclusions.
X. Qirinxhi was informed by the first author, shortly before publication, of her inclusion as a co-author and only received an abstract version of the manuscript. Unfortunately, there was insufficient time to provide suggestions, and no opportunity to review the full manuscript.
Due to the discrepancies and the lack of clarity regarding the contributions, X. Qirinxhi requested the removal of her name as a co-author in a correction. Unfortunately, the first author conveyed that this request could not be accommodated.
To ensure appropriate credit is given to contributors in scientific publications, it is essential to resolve this issue and provide clear acknowledgment of each contributor’s role.

B. Abstract
The timing of the first publication, concerning the discovery of the disjunct Pseudochazara population, is incorrectly stated in the sentence “Initially classified as P. tisiphone Brown, 1981 in the atlas "Fluturat e Shqipërisë, …".
The discovery of the new population in Dibër County was first made public in Cuvelier, Parmentier et al. (2018), which should be cited instead. This update is documented in Results 11, which tracks developments from 2018 to the present in the dynamic website Fluturat e Shqipërisë, where a Monograph on P. tisiphone is featured.
It is not entirely clear whether the distinction in: “Our results show that the central Albanian populations found in the eastern belt of the Mirdita zone in the vast inner Albanides Massif are distinct from known P. tisiphone populations in the western ophiolitic belt, and the surroundings of the Pindos Massif in Greece.” refers specifically to different zones in Albania. Possibly open to multiple interpretations, a more specific clarification of whether the Dibër population, the Korçë-Gjirokastër population, and the Greek Pindos population belong to distinct geological zones would help clarify this quote.
In the map for Albania (Çina 2016 Fig. 1) the three disjunct areas with Pseudochazara populations (Korçë-Gjirokastër, Bulqizë and Lurë) are all situated in the eastern belt with MORB-type ophiolites. The situation in the Greek Pindos ophiolite belt, comprising both MORB-type and SSZ-type sequences, is less clear as a sharp distinction cannot be made (Barth et al. 2008; Smith 1993). The habitats of Greek Pseudochazara populations (Pamperis distribution map 2025) are dominated by typical ophiolitic rocks, and are mainly but not exclusively found on SSZ ophiolites.

C. Introduction
Page 23: The statement “it was split from P. mamurra and raised to species level as P. tisiphone Brown, 1981 (Takáts & Mølgaard (2016).” is inaccurate.
In Takáts & Mølgaard (2016), P. tisiphone is not associated with P. mamurra, as this may be a confusion with Pseudochazara amymone Brown, 1976. Instead, it is associated with P. mniszechii and remains classified as P. mniszechii tisiphone, not raised to species level. This was confirmed by the second author in correspondence from March 2025.
Page 24: There is vagueness regarding the date of discovery: "In July 2017… the first author discovered an unknown population of Pseudochazara sp."
This contradicts Cuvelier, Parmentier et al. (2018 p. 84), which provides a more precise account: "On 13.vii.2017, VVE1 discovered good populations of P. tisiphone around Bulqizë (Dibër County). This is an important extension of its distribution, some 100 km north of Korçë County. P. tisiphone was present in stony habitats on ophiolite substrate, very similar to the known localities nearby Korçë. On 18.vii.2017, not aware of these findings, VVE2 also observed P. tisiphone in the same area and nearby Krastë (S4 map 171)."
Furthermore, the statement " This observation was also confirmed independently by J. Couckuyt and S. Cuvelier during their joint expedition…" should be revised to reflect the accurate dates and context as presented in the original publication.
Page 24: Another error appears in the following statement: “… shortly after, these observations were merged in a new distribution atlas of Albanian Lepidoptera: “Fluturat e Shqipërisë”.
The discovery in Dibër County was first reported in Cuvelier, Parmentier et al. (2018). A more accurate statement would be: "The discovery of the new population was first reported in Cuvelier, Parmentier et al. (2018), followed by its inclusion in the new distribution atlas."
Page 24: The statement “… and a single rough measurement of male androconial scales (Cuvelier et al. 2018).” is also incorrect. In Cuvelier, Parmentier et al. (2018), androconial scales were measured using a calibrated digital microscope, and three distinct androconial cells were analysed (Figs. 12c-d).
Page 24: Incorrect year. The statement “… was quickly linked to the populations studied near Krastë, Bulqizë back in 2018.” is inaccurate.
Both Cuvelier, Parmentier et al. (2018) and higher on page 24 in Parmentier & Qirinxhi (2025) clearly indicate that the populations were studied in 2017, not 2018. This error should be corrected to reflect the correct year.
Fig. 1. Page 25: Incorrect location data and County assignments. The locations listed as “Albania, Dibër prov., env. Bobosthicë, ... G. Albania, Dibër prov., env. Gjergjevicë, …” are incorrect.
Both Boboshtiçë and Gjergjeviçë are actually located in Korçë county, not Dibër. While it is unlikely that similar mistakes occurred in the analysis, it would be beneficial to publish the data for individual specimens and correct these inaccurate location assignments.

D/ Biometric analysis
On Page 26, the localisation of P. orestes is incorrectly stated: “and P. orestes belonging to the same species group but originating from western Greece.”
In fact, P. orestes was discovered in the province of Drama in eastern Greece and later found in nearby Bulgarian habitats. It is restricted to specific mountain ranges in this region, not western Greece as stated.

E/ Statistics
On Page 27, the information provided about male and female traits is incorrect. The publication states, "4 traits for males and females", but "External wing traits" (p. 26) lists 5 traits selected for both males and females. This should be amended to reflect the accurate number.

F/ Results
Fig. 3. Page 28 states “… found in the eastern belt (A-D) and western belt (E-F) of the ophiolitic Mirdita zone …” However, this classification does not align with Fig. 1 in Çina (2016), which clearly marks the Lurë massif, the Bulqizë massif, the Morava massif near Korçë, the Voskopojë–Gjergjeviçë area, and a region in Gjirokastër County in darker blue as "HOT-type ultramafic massif (harzburgite with interbedded dunites, Eastern belt)."
On Page 28, incomplete information is given regarding Pseudochazara amalthea (Frivaldsky, 1845). The statement “… unpublished sites for Pseudochazara amalthea (Frivaldsky, 1845), were found near Skënderbej and Qarrishtë, …” fails to account for the updated Atlas (2025), which marks a dot near Skënderbej representing an ophiolite habitat.
P. amalthea is primarily found in limestone slopes and has a broader distribution, extending into Bosnia & Herzegovina and Croatia, far to the north of the Skënderbej and Qarrishtë locations. Therefore, extrapolating the distribution of other Pseudochazara species based on P. amalthea may not be appropriate.

G/ Character of anroconial (sic) scales
A typo is present on Page 32 regarding the character of androconial scales. We recommend to correct this in the next version of the publication.

H/ Paratypes
On Page 32, there are inconsistencies in the spelling of the National Park near Fushë-Lurë. We recommend, for consistency, to adjust this in the future version of the manuscript.

I/ Taxonomic position compared to sister species
On Page 34, the date mentioned as “taken from the 2018 expeditions” is incorrect. Both Cuvelier, Parmentier et al. (2018) and Parmentier & Qirinxhi (2025) indicate that the expedition occurred in 2017, not 2018. This discrepancy should be addressed.

J/ Distribution pattern of ophiolitic habitats in the western Balkans
On Page 35, the statement “Knowing the generally accepted status of both taxa as bona species (Takáts & Mølgaard (2016); Verovnik & Wiemers 2016).” is inaccurate.
In Takáts & Mølgaard (2016), P. tisiphone is associated with P. mniszechii, and remains classified as P. mniszechii tisiphone, thus not raised to species level. This classification was confirmed by the second author (mail March 2025) and should be corrected in the manuscript.
On Page 35-36, the previously noted inaccuracy regarding the western and eastern belts of the Mirdita ophiolite, specifically in relation to the distribution of Pseudochazara populations, is repeated. This issue, already addressed in sections B (Abstract) and F (Results. Fig. 3. Page 28) in connection with Fig. 3 on page 28, should be reconsidered in light of the discussion provided in both sections.
On Page 36, the phrase “… with the Voúrnios Massif …” appears to contain a typographical or transliteration error. The correct spelling is Vourinos Massif (Greek: Βούρινος), referring to a mountain range located in the eastern Grevena and southern Kozani regional units of Western Macedonia. It may be worth considering a correction from Voúrnios to Vourinos for clarity.

L/ Acknowledgements
On page 37, we suggest to change the name 'Regila Fero' to 'Rigela Fero,' as it appears to be a typographical error.

M/ References
On Page 37, there is a misattribution in the reference to Dapporto L., Menchetti M., Voda R., Corbella C., Cuvelier S., DjemPark et al.: The genus Lecithocera in Kenya and Tanzaniaadi I., Gascoigne-Pees M., Hinojosa J. C., Lam N. T., Serracanta M., Talavera G., Dinca V. & Vila R. 2022. The atlas of mitochondrial genetic diversity for Western Palaearctic butterflies. — Global Ecology and Biogeography 31(11): 2184–2190.
The citation includes part of another reference likely relating to the chief editor’s work on African butterflies, while the cited source itself is focused on Western Palaearctic butterflies. This misattribution may lead to confusion regarding the scope and relevance of the cited research. We recommend correcting this in the next version of the manuscript.

In conclusion, addressing the inaccuracies and discrepancies outlined in this chapter, will contribute to a more accurate and reliable version of Parmentier & Qirinxhi (2025). These corrections will improve the clarity and credibility of the publication, ensuring that future research in this field is built on a solid foundation. Implementing these changes will enhance the overall quality and trustworthiness of the manuscript, benefiting both the scientific community and the broader public interested in Lepidoptera research.

9/ Clarification of the publication timeline since Gross's 1973 description of P. sintenisi cingovskii, highlighting subsequent taxonomic revisions, including name changes and synonymies
Due to inaccuracies and confusion, it became essential to clarify the publication timeline since Gross’s (1973) description of Pseudochazara sintenisi cingovskii, along with the subsequent taxonomic changes and synonymies, unveiling a complex and evolving history of classification within the Pseudochazara genus. For a comprehensive overview, please refer to Supplementary material 2.
Early contributions by Gross (1973), Gross (1978), Koutsaftikis (1974), and Brown (1976) laid the foundation for identifying and differentiating taxa across the Balkans. Over time, additional studies introduced new taxa, reclassified populations, and highlighted inconsistencies in earlier identifications.
The application of COI barcoding, led to new attempts to classify and resolve relationships among closely related Pseudochazara populations. Recent developments, including the consideration of the Dibër County population as either Pseudochazara tisiphone dibra or Pseudochazara misjai, highlight the ongoing complexities and uncertainties in the taxonomic understanding of this group.

10/ Review of the literature on P. tisiphone in Albania, from its initial mention to the present
An overview of the publications and discoveries of P. tisiphone in Albania, including P. tisiphone dibra, has been comprehensively presented in Results, to address inaccuracies and clarify the progression of findings throughout the country.
Further updates will be provided as needed in Fluturat e Shqipërisë and, when applicable, in future publications.

11/ Review of the updates (2018 to the present) of the dynamic website Fluturat e Shqipërisë which features a Monograph on P. tisiphone
An overview of the evolution of Fluturat e Shqipërisë is presented in Results, to address confusions and inaccuracies originating from external sources.

Conclusion

It has been demonstrated that the population in Dibër County, first described in Cuvelier & Marafi (2025) and later referenced in Parmentier & Qirinxhi (2025) with a different proposed taxonomic status, refers to the same population. The ecological niche consists of disjunct yet comparable ophiolite habitats.
Differences in macroclimatic factors such as Tmax, Tmin, solar irradiation, precipitation, and wind exposure, combined with variation in mineral composition and concentration across sites, likely drive the phenotypic flexibility that underlies the documented variability within the population. These findings underscore the potential for future research to explore how climatic factors may influence morphological variations within the Pseudochazara genus. Such studies could provide valuable insights into the adaptive strategies of these butterflies in response to changing environmental conditions.
Discordant data regarding androconial scales, with no clear functional link due to the lack of information on associated chemical compounds, adds further confusion to the taxonomic interpretation. This uncertainty echoes the difficulties highlighted by Wakeham-Dawson (1997), who noted the challenge of reliably distinguishing P. tisiphone and P. cingovskii based solely on androconial traits.
As expected and documented in Parmentier & Qirinxhi (2025), male genitalia did not offer useful traits for differential morphometric analysis. Neither androconial structures nor genitalia provide meaningful insights into potential prezygotic barriers to reproductive isolation, further complicating efforts to delineate species boundaries based on morphological criteria alone.
The available COI data shows low divergence, which alone is insufficient to confirm speciation. While COI is useful for species identification, it represents only mitochondrial DNA and does not capture broader genomic changes. Speciation typically involves divergence across the nuclear genome, not just a single gene. The four COI mutations observed may reflect phenotypic plasticity rather than true genetic isolation. Broader genomic data and consideration of environmental factors are needed to draw reliable conclusions.
An analysis of the available data and published material revealed several oversights, inconsistencies, and inaccuracies. To improve clarity and ensure scientific accuracy, it would be beneficial to correct these issues, ideally through a revised version of the work uploaded to the Figshare repository, where it can be properly updated and referenced.
To avoid future confusion, a thorough review was conducted encompassing global literature, Albanian records, and the historical development of Fluturat e Shqipërisë, in relation to the P. tisiphone Monograph.

The question is not whether the glass is half full or half empty; rather the inconvenient truth is that more robust and comprehensive evidence still needs to be developed.
In light of the current lack of substantial evidence, it would be wise to adopt a cautious approach regarding the populations of P. tisiphone, avoiding a predetermined taxonomic perspective (Dufresnes et al. 2023) and treating the newly discovered population in Dibër County as initially done at the subspecies level until concrete evidence emerges.

The species [Pseudochazara misjai Parmentier, 2025], originally described as a distinct species, is here placed in synonymy with the earlier described subspecies [Pseudochazara tisiphone dibra Cuvelier & Marafi, 2025], as both refer to the same population. This synonymization adheres to the principles outlined in the International Code of Zoological Nomenclature (ICZN), particularly Article 23 (Priority), which asserts that the first validly published name takes precedence. Furthermore, Article 50 emphasizes the importance of nomenclatural stability, and this synonymization helps avoid unnecessary taxonomic confusion. The criteria for synonymy are met, as there is no significant morphological, behavioural, or genetic differentiation between the two, as demonstrated by Dincă et al., Supplementary Data 14 (2021), Dapporto et al., Appendix S1 (2022), Cuvelier (2023), Cuvelier & Marafi (2025) and Parmentier & Qirinxhi (2025). The type specimens of the species and subspecies align, as per Article 24.
The publication complies with ICZN Article 8.5, as amended for electronically distributed works. The publication was made available online in AWPL (ISSN 3041-6531) and the corresponding pdf was permanently archived in Zenodo on January 26, 2025.
The ZooBank registration dates are provided both for the publication itself and for the newly described subspecies.
The action adheres to Article 11 (Name Availability), ensuring that both names were validly published.
Pseudochazara tisiphone dibra Cuvelier & Marafi was officially registered in ZooBank on January 26, 2025, under the LSID: urn:lsid:zoobank.org:pub:E8AC9881-7EDE-47A2-B1CF-067152352AAE.
Pseudochazara misjai Parmentier, 2025, was officially registered in ZooBank (accessed April 14, 2025), but the publication date is not specified, under the LSID: urn:lsid:zoobank.org:act:DEAE68B2-B9A7-405F-966D-C6F58A52F9BB.

Author contribution

The three authors contributed to the conceptualization, writing and manuscript preparation. The first author took the lead in the data analysis, literature review, and reference compilation. All authors also participated in editing the manuscript.

Acknowledgement

We would like to express our gratitude to A. Wakeham-Dawson for granting permission to include figures from his 1997 publication, and to L. Pamperis and J. Coutsis for providing valuable background information on P. tisiphone habitats in Greece. Our thanks also go to M. Taymans for his analysis regarding synonyms and ICZN rules, and to M. Mølgaard for confirming their stance on P. mniszechii tisiphone and its taxonomic status. We thank V. Dincă for valuable discussions on DNA barcoding and the supplementary materials of the genus Pseudochazara. We appreciate T. Garrevoet for his support with the preliminary website of Fluturat e Shqipërisë on the FES server. We gratefully acknowledge X. Qirinxhi for the open and ethical attitude regarding her contribution to the article Parmentier & Qirinxhi (2025). Finally, we acknowledge E. Halimi for serving as an external reviewer.

Supplementary material

S1. Long-term monthly climate data from Meteoblue for Bulqizë (41.440°N, 20.200°E) and Lurë (40.540°N, 20.770°E).
S2. Publication timeline since Gross's 1973 description of Pseudochazara sintenisi cingovskii, including subsequent taxonomic changes and synonymies.
S3. Comparative analysis of androconial measurements.
S4. Pseudochazara neighbor-joining tree.
S5. Pseudochazara haplotype network.
S6. Haplotype summary report and list.

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