Exploration of Mount Žeden and discovery of Proterebia phegea (Borkhausen, 1788), a new species for the fauna of North Macedonia.
Brief summary of current knowledge on Proterebia phegea.
Published online: 30.vi.2025.
DOI: 10.5281/zenodo.15723544

Michel Taymans¹ | Sylvain Cuvelier²
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¹ Taymans M., Clos du Moulin Royal 2 bte 02, B-6900 Marche-en-Famenne, Belgium. michel.taymans@hotmail.com
² Diamantstraat 4, B-8900 Ieper, Belgium. sylvain.cuvelier@telenet.be

Abstract
The butterfly species Proterebia phegea (Borkhausen, 1788), previously unrecorded in North Macedonia, was discovered during field surveys on Mount Žeden in April 2025.
This finding extends the known range of the species in the Balkan Peninsula, which includes populations in Croatia, Bosnia and Herzegovina, Albania, and Greece.
Mount Žeden, located approximately 20 km west of Skopje, is characterised by a continental Mediterranean climate and diverse karstic topography, providing a unique habitat for various butterfly species. The discovery of P. phegea on Mount Žeden contributes to the understanding of the species' distribution and highlights the importance of this area for butterfly conservation.

Key words
Proterebia phegea — Papilionoidea — Nymphalidae — Satyrinae — Faunistics — Taxonomy — Conservation — Mount Žeden — North Macedonia — Balkan Peninsula.

Introduction
A preparatory map survey of Mount Žeden revealed two stone roads accessible by car, both leading to the massif's summits from the eastern side. These stone tracks enabled three field explorations (on April 19th, 20th, and 30th, 2025), during which a notable diversity of butterfly species was recorded.
Among the species recorded was Proterebia phegea (Borkhausen, 1788), which had not been previously reported from North Macedonia.
Mount Žeden (Жеден in Cyrillic) is a moderate-elevation mountain massif located in northwestern North Macedonia, approximately 20 kilometers west of the capital, Skopje. It is bounded to the south by the valley between Skopje and Tetovo, through which the main highway connecting the two cities runs, and to the north by the Vardar River and the foothills of the Šar Mountains and Skopska Crna Gora.
Mt. Žeden extends in a southwest–northeast direction. The landscape features predominantly rounded relief, culminating in several peaks, the highest being Vodeni Kamen (1.264 m a.s.l.). The summits can be reached via two stone roads originating near the villages of Kopanica and Boyanë.
As its name suggests (Žeden means "without water" in Macedonian), the mountain is devoid of tall vegetation and surface water, though several springs arise at its base. The most significant of these has a high flow rate and supplies the city of Skopje with potable water. Only a few small, slightly incised valleys host denser vegetation and a richer fauna.
Mt. Žeden experiences a continental Mediterranean climate, characterised by hot, dry summers marked by abundant sunshine and relatively low precipitation, and cold winters, with heavy snowfall. Spring and autumn are transitional seasons with moderate rainfall contributing to the mountain’s diverse vegetation and microclimates.
Mt. Žeden is primarily composed of sedimentary rocks, including limestone and dolomite, which date back to the Mesozoic era. Its geological structure features karst formations, characterised by caves, sinkholes, and rugged rocky outcrops. The mountain’s rounded relief and steep cliffs result from long-term erosion and tectonic uplift.

Biotopes
Three distinct biotopes were identified during the ascents:
A. The lower grassland zone at around 400–500 m a.s.l. (Fig. 1-2)
Immediately after leaving the aforementioned villages, the tracks pass through open grasslands, characterised by sparse vegetation, predominantly juniper (Juniperus spp.) and Christ's thorn (Paliurus spina-christi). Near the villages (around 400 m), the landscape remains green at this time of the year, but above 500 m, vegetation becomes sparse and dry.
The following species were documented in this area:
Iphiclides podalirius (Linnaeus, 1758), Papilio machaon Linnaeus, 1758, Erynnis tages (Linnaeus, 1758), Leptidea sinapis (Linnaeus, 1758), Colias croceus (Geoffroy, 1785), Pieris rapae (Linnaeus, 1758), Anthocharis cardamines (Linnaeus, 1758), Lycaena phlaeas (Linnaeus, 1761), Callophrys rubi (Linnaeus, 1758), Celastrina argiolus (Linnaeus, 1758), Tarucus balkanica (Freyer, [1843]) (1), Aricia agestis ([Denis & Schiffermüller], 1775), Polyommatus icarus (Rottemburg, 1775), Issoria lathonia (Linnaeus, 1758), Vanessa cardui (Linnaeus, 1758), Lasiommata megera (Linnaeus, 1767), Coenonympha pamphilus (Linnaeus, 1758), an unidentified Procridinae, and Proterebia phegea (Borkhausen, 1788).
Only a few scattered and isolated specimens of P. phegea were recorded in this zone. The males, generally worn and scarce, flew long distances without settling, while the females were more numerous, fresher, and frequently observed feeding on flowers.

Fig. 1. Habitat of Proterebia phegea. Kopanitsa (around 460 m a.s.l.), City of Skopje administrative area, North Macedonia (© Bénédicte Jonckers)
Fig. 2. Habitat of Proterebia phegea. Boyanë (around 450 m a.s.l.), City of Skopje administrative area, North Macedonia (© Bénédicte Jonckers)

B. Mid-elevation valleys, at approximately 500–900 m a.s.l.
The ascent continues through small, gently sloping valleys dominated by shrubs and low-growing plants. Rocky outcrops and scree are common, especially near the summit zone. In addition to all species recorded in zone A, the following additional species were observed here:
Zerynthia polyxena ([Denis & Schiffermüller], 1775) *, Spialia orbifer (Hübner, [1823]), Spialia phlomidis (Herrich-Schäffer, [1845]) *, Pyrgus malvae (Linnaeus, 1758), Muschampia orientalis (Reverdin, 1913), Leptidea duponcheli (Staudinger, 1871), Gonepteryx rhamni (Linnaeus, 1758), Colias alfacariensis Ribbe, 1905, Pieris mannii (Mayer, 1851), Pieris balcana Lorković, [1969], Anthocharis gruneri Herrich-Schäffer, [1851] *, Pseudophilotes bavius (Eversmann, 1832) *, Pseudophilotes vicrama (Moore, 1865), Scolitantides orion (Pallas, 1771) *, Cupido minimus (Fuessly, 1775), Aricia anteros (Freyer, 1839), Lysandra bellargus (Rottemburg, 1775), Polyommatus thersites (Cantener, 1835), Vanessa atalanta (Linnaeus, 1758), Aglais io (Linnaeus, 1758), Nymphalis polychloros (Linnaeus, 1758), Nymphalis antiopa (Linnaeus, 1758), Melitaea ornata Christoph, 1893, and Melitaea cinxia (Linnaeus, 1758).
* species included in the Red List of North Macedonia (Krpač & Darcemont 2022)
Only a few isolated specimens of P. phegea, both male and female, were observed here, feeding on flowers along the trails. These are likely to have dispersed from the upper slopes and appeared somewhat uncharacteristic for this habitat. These mid-elevation valleys could also serve as corridors or transit areas between more suitable biotopes.

C. Summit zone, at approximately 900–1.200 m a.s.l. (Fig. 3-4)
Above 900 m, the landscape opens into a high plateau characterised by grassy and rocky slopes, with little to no tall vegetation. The summit, reaching around 1,200 m a.s.l., is marked by exposed terrain where a few scattered cattle graze. On April 20th, several males of P. phegea were observed flying low over the south-facing slopes. During the third visit, on April 30th, this species was much more common. Both males and females, were present across all accessible slopes.
Few other species were observed at this elevation: P. machaon, C. croceus, A. cardamines, A. agestis, and V. atalanta.

Fig. 3. Habitat of Proterebia phegea. Summit zone above Boyanë (1000-1100 m a.s.l.), City of Skopje administrative area, North Macedonia (© Bénédicte Jonckers)
Fig. 4. Habitat of Proterebia phegea with in the background, the snow-covered Šar Mountains. Summit zone above Boyanë (1000-1100 m a.s.l.), City of Skopje administrative area, North Macedonia (© Bénédicte Jonckers)

Results
A total of 42 butterfly species were documented on Mount Žeden during three field visits conducted over a ten-day period. The discovery of P. phegea constitutes a new addition to the documented butterfly fauna of North Macedonia and suggests that further exploration of the region may yield additional faunal records of interest.
The localities where P. phegea was found on Mt. Žeden across different biotopes are shown in Fig. 5.
Fig. 6 presents the updated distribution of P. phegea across the Balkan Peninsula, emphasizing the distinct and geographically isolated population discovered on Mt. Žeden (North Macedonia).

Fig. 5. Observation points of P. phegea observations on Mt. Žeden and its potential distribution area (© Michel Taymans)

	Potential distribution area of P. phegea.
Observations across habitat zones: Lower grassland zone, Mid-elevation valleys and Summit zone.

The atlas Die Tagfalter von jugoslawisch Mazedonien (Schaider & Jaksic 1989) is relatively old and certainly outdated compared to the current state of knowledge. Nevertheless, it remains the only centralised source of information on the butterflies of North Macedonia. The species listed below have not been documented within UTM square 34T EM 15, corresponding to the surveyed area: P. machaon, Z. polyxena *, E. tages, L. duponcheli, L. sinapis, G. rhamni, C. alfacariensis, P. rapae, P. mannii, P. balcana, A. cardamines, C. rubi, C. argiolus, T. balkanica, P. bavius *, P. vicrama, S. orion *, C. minimus, A. agestis, A. anteros, L. bellargus, P. icarus, P. thersites, V. cardui, N. polychloros, N. antiopa, M. ornata, M. cinxia, L. megera and P. phegea.

Fig. 6. Updated distribution map of P. phegea in the Balkan Peninsula (© Sylvain Cuvelier)
Known localities of P. phegea in the Balkans.
New localities of P. phegea in North Macedonia.

Brief summary of current knowledge on Proterebia phegea (Borkhausen, 1788)

A. Range (Fig. 7)
P. phegea has a highly fragmented distribution, from Central Asia in the east to the Balkans in the west:
Europe: S Russia, the nominal ssp. was described from the steppes of Saratov and occurs further north on the western slopes of the Ural mountains; Croatia, known since Godart (1824), who described the ssp. dalmata in 1824 from the surroundings of Šibenik; Crimea, known since Nordmann (1851) and Sheljuzkho (1929) who described the ssp. krymaea from the surroundings of Yalta; Greece, known since De Louker & Dils (1987), who described the ssp. pyramus from the Greek province of Kozani; Bosnia and Herzegovina, see Koren & Trkov (2011); two disjunct populations in Albania, see Verovnik R. & Verovnik J. (2022) and Prendi et al. (2022); Ukraine, to our knowledge, recent data of confirmation is lacking; and now North Macedonia.
Asia: Turkey; Armenia; Georgia and Azerbaijan, known since Fruhstorfer (1918), who described ssp. zyxuta from Ganja and generally of the Caucasus; Iran, known since Staudinger (1901 ), who described ssp. hyrcana from Hajjiabad; Kazakhstan and S Russia, known since Fruhstorfer (1918), who described ssp. bardines from Altai; Turkmenistan, known since Goltz (1930), who described ssp. transcaspica from Akhal-Teké; Kyrgyzstan, see Tshikolovets (2005) and Uzbekistan, see Korb & Bolshakov (2016).
In Fig. 7.

Fig. 7. Global range of P. phegea, red dots indicate approximate locations on the map (Source: Wikipedia, modifications © Michel Taymans)

B. Habitat
In Europe, P. phegea generally inhabits hot, stony grassland slopes or vast steppe-like terrains with sparse vegetation on limestone substrate, ranging from sea level on the Island of Pag in Croatia (Zakšek 2005; Mihoci & Šašić 2007) to much higher elevations at around 1400 m in Croatia (Koren et al. 2010).
Some populations have also been observed on ophiolite terrain in northern Greece and Albania, where P. phegea occupies more closed habitats such as grassland patches confined between wooded areas. Depending on the extent of these habitats, the species can be common with a high population density.

C. Life cycle and foodplants
Adults: one generation occurs annually from late March to May, depending on altitude. Butterflies feed on available flowers during this period.
Egg: the egg stage is brief.
Note: a female captured on Mt. Žeden for DNA analysis laid 90 eggs in her storage envelope, and after 15 days, the larvae successfully hatched.
Caterpillar: caterpillars aestivate, resume growth in autumn, and overwinter as mature larvae. Their primary food plants include Festuca ovina, ...
Pupa: the pupal stage is short.

D. Nomenclature
The nomenclature of this species is unstable, as three different scientific names have been applied historically to what appears to be the same taxon. These names are considered to be objective or subjective synonyms, depending on the interpretation, and each has been used at various times by different authors. Although each name has valid historical arguments in its favour, all are also subject to nomenclatural dispute. In order of publication, the three specific names are:

Papilio afer Esper, [1783]. Schmetterlinge in Abbildungen nach der Natur 1(2): 161–162, pl. 83, figs. 4–5. (text : url ; plates : url)
(original combination: 'P[apilio]. Nymph[alis]. Gem[matus]. Afer');

Papilio afra Fabricius, 1787. Mantissa Insectorum 2: 41. (url)
(original combination: 'P[apilio]. N[ymphalis]. G[emmatus]. Afra');

Papilio phegea Borkhausen, 1788. Naturgeschichte der europäischen Schmetterlinge 1: 101–102. (url)
(original combination: '[Papilio oreades] Phegea')

2. Validity of the name ‘afra Fabricius, 1787’
Since 'Papilio afer Esper, [1783]' would be a homonym of 'Papilio afer Drury, 1782', the next oldest available name is 'Papilio afra Fabricius, 1787'.
The issue lies in the Latin grammar: 'afra' (meaning “African” in the feminine form) is the feminine declension of 'afer' (masculine). Therefore, since a species name must in principle agree in gender with the genus, and the genus Papilio is masculine, 'afra' should be modified to 'afer'. As a result, it would then become a homonym of 'afer Esper, [1783]'. However, simplifying somewhat, since the Code is complex (see Article 31.2 of the ICZN Code), such gender agreement is only required if the taxon was originally described as an adjectival epithet, and not if it was used as a noun. Article 31.2.2 of the Code states that if a taxon name can be interpreted as either an adjective or a noun, it should not be modified for gender agreement.
Some authors argue that Fabricius may have been inspired by Esper’s name, which he translated into German as 'Der Mohr', a noun, not an adjective, meaning 'the Moor' (i.e. 'the African'). Moreover, Fabricius wrote 'Afra' with a capital letter, which would indicate that he intended it as a noun, not as an adjective.
Thus, the entire matter hinges on how the taxon name is interpreted, as a noun or as an adjective, and how the relevant grammatical rules of Latin are applied under the Code.
According to the opinion of Nieukerken et al. (2019), enforcing gender agreement can hinder nomenclatural stability, a view we adopted when compiling the checklist (Taymans & Cuvelier 2024; Taymans & Cuvelier 2025). Based on that reasoning, 'afra' could be accepted as valid.
However, another article of the Code also applies in this case. Article 33 states that if the original spelling of a taxon is changed, and both names are cited, such a change may be considered unjustified. Since Fabricius did cite the name 'Papilio afer' Esper in the publication where he introduced 'afra', his name could be seen as an unjustified replacement name.

3. Validity of the name ‘phegea Borkhausen, 1788’
This is the next available and valid name if the two previous ones ('afer' Esper and 'afra' Fabricius) are considered homonyms

Conclusions
Proterebia phegea (Borkhausen, 1788) is recorded here for the first time from North Macedonia, representing a notable range extension and, based on current knowledge, a geographically disjunct population. Further disjunct populations are likely to be revealed through systematic field surveys in ecologically suitable areas, especially during early spring when environmental conditions favour adult emergence and detectability.
Mt. Žeden represents a biodiversity hotspot for butterflies and harbours multiple species classified as threatened on the North Macedonia Red List. Additional surveys conducted throughout the year are essential to obtain a fuller picture of the total butterfly diversity.

Author contribution
Michel Taymans: conceptualization, field work, analysis, visualization, writing - original draft, writing – review and editing.
Sylvain Cuvelier: analysis, validation, visualization, writing – review and editing.

Acknowledgements
We are sincerely grateful to Peter C. Russell for his careful and thorough review of the final draft. His attention to detail and thoughtful feedback have been invaluable in helping us refine and strengthen the work. We are grateful to Lazaros Pamperis for his input on the distribution data featured in the latest distribution map (2025), and Beau De Block for informing us about a locality of P. phegea he recently discovered in eastern Greece. We thank Vladimir Savchuk and Joseph Verhulst for providing helpful literature references. Finally, we would like to express our gratitude to Bénédicte Jonckers for the habitat photography.

Annotated bibliography

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New taxon: Erebia afer hyrcana.

Taymans M. & Cuvelier S. 2024. A dynamic checklist of the Western Palearctic butterflies, hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Lépidoptères - Revue des Lépidoptéristes de France 33(84): 18-21 (url)

Taymans M. & Cuvelier S. 2025. A dynamic checklist of the Western Palearctic butterflies hyperlinked to the original descriptions at species, genus and family level (Lepidoptera, Papilionoidea). — Archives of Western Palearctic Lepidoptera 2025(1): 1-70. doi.org/10.5281/zenodo.14733224

Tshikolovets V. V. 2005. The Butterflies of Kyrgyzstan. — Kyiv-Brno: Tshikolovets (Ed.). 500 p.

Verovnik R. & Verovnik J. 2022. First record of Proterebia phegea (Lepidoptera: Satyrinae) from Albania. — Natura Croatica 31(1): 115-120, fig. (url)
First observation record in Albania (Dibër county)

Zakšek V. 2005. On the presence of Proterebia afra (Lepidoptera: Nymphalidae, Satyrinae) on the island of Pag, Croatia. — Phegea 33(3): 118-120, fig. (url)
Additional observations in Island of Pag, Croatia.