The classification is given according to The Euro+Med PlantBase (source: Christenhusz & Raab-Straube, 2013).
Determined according to the botanical keys by Heywood in Tutin et. al (1964), Heywood revised by Praser-Jenkins in Tutin et al. (1993), Micevski (1985), Fraser-Jenkins (2007), Szczęśniak, Tlałka, and Rostański in Szczęśniak and Gola (2009) and Woziwoda in Szczęśniak and Gola (2009).
Somewhat morphologically similar to Dryopteris filix-mas (L.) Schott, which also occurs in the Osogovo region. D. borreri differs in having somewhat darker and more reddish/brownish, denser petiole scales, and longer and denser rachis scales. Also differs in the shape of the pinnules, which are more oblong, with truncate apices, the lower with rectangular lobes and sometimes auricle. Indusium thin, in ripe sori shriveling and lifting upwards, obtaining cone-like shape (inverted cone or funnel). The specimens from Osogovo (gorge of “Kochanska” river, Ponikva – “Venechka” River region) usually have the proximal basioscopic pinnula of the basal pinnae partly adnate (c. 1/4) to the secondary rachis, while the proximal acroscopic pinnula is stipitate.
Lifeform: Perrenial; Hemicryptophyte according to the Raunkiær system.
Distribution: Native from Europe to Caucasus according to the distribution information provided by the Plants of the World Online portal (Royal Botanic Garden, Kew) and The Euro+Med Plantbase (Christenhusz & Raab-Straube, 2013+).
Habitat: Found in somewhat shaded positions in oak and European beech forest, river gorges, from the lowland up to c. 1600 m altitude.
Locality: Jamishko Osoe region, above the springs of “Venechka” River.
Dryopteris borreri – habit. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – habit. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaves. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae of middle part of leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae of middle part of leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae of middle part of leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae of middle part of leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae of middle part of leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae of middle part of leaf. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnules of middle pinna. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and blackish point of insertion of secondary rachis. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and blackish point of insertion of secondary rachis. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – adaxial side of pinna with sori. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of upper pinna with sori. Indusium more or less thin, not splitting, in mature sori shriveling and lifting to a cone-like shape. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of upper pinna with sori. Indusium more or less thin, not splitting, in mature sori shriveling and lifting to a cone-like shape. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of pinna with sori. Indusium more or less thin, not splitting, in mature sori shriveling and lifting to a cone-like shape. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of pinnules with sori, secondary rachis with scales, blackish point of incision of secondary rachis and rachis with scales. Indusium more or less thin, not splitting, in mature sori shriveling and lifting to a cone-like shape. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of pinnules with sori and secondary rachis with scales. Indusium more or less thin, not splitting, in mature sori lifting to a cone-like shape. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of pinnules with sori and secondary rachis with scales. Indusium more or less thin, not splitting, in mature sori shriveling and lifting to a cone-like shape. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of pinnules with sori. Indusium more or less thin, not splitting, in mature sori shriveling and lifting to a cone-like shape. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – basal pinnae. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – basal pinnae. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – basal pinnae. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – basal pinnae. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – acroscopic and basioscopic pinnules of basal pinnae. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – acroscopic and basioscopic pinnules of basal pinnae. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – acroscopic and basioscopic pinnules of basal pinna. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis scales. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis scales. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petioles. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petioles. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petioles. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petiole scales. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petiole scales. Osogovo – Ponikva, Jamishko Osoe region, 19.10.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaf. Specimen from Osogovo – Ponikva, Jamishko Osoe region. Scale in cm. Photo by Trajche Dimitrovski.Dryopteris borreri – leaf. Specimen from Osogovo – Ponikva, Jamishko Osoe region. Scale in cm. Photo by Trajche Dimitrovski.Dryopteris borreri – leaf. Specimen from Osogovo – Ponikva, Jamishko Osoe region. Scale in cm. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of pinnules with sori. Specimen from Osogovo – Ponikva, Jamishko Osoe region. Scale in mm. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis scales. Specimen from Osogovo – Ponikva, Jamishko Osoe region. Scale in mm. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis scales. Specimen from Osogovo – Ponikva, Jamishko Osoe region. Scale in mm. Photo by Trajche Dimitrovski.Dryopteris borreri – petiole scales. Specimen from Osogovo – Ponikva, Jamishko Osoe region. Scale in mm. Photo by Trajche Dimitrovski.
Locality: Gorge of “Kochanska” River.
Dryopteris borreri – habit. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – habit. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – habit. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaves. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaves. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and pinnae with pinnules. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and pinnae with pinnules. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and pinnae with pinnules. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and pinnae with pinnules. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and pinnae with pinnules. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and pinnae with pinnules. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of pinnae. The indusium is embracing the sporangium when young. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis with scales and abaxial side of pinnules with sori. The indusium is embracing the sporangium when young. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis with scales and abaxial side of pinnules with sori. The indusium embracing the sporangium when young. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and abaxial side of pinnules with sori. The indusium in mature sori lifts upwards and shrinks, becoming and inverted cone of funnel shaped. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – abaxial side of pinnules; sori with lifted indusium. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis with scales; sori with lifted indusium. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis with scales; sori with lifted indusium. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis with scales; sori with lifted indusium. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis with scales. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis with scales. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – lower part of leaves. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petioles of leaves. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petioles with scales. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petiole with scales. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – petioles with scales. Gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – leaves. Scale in cm. Specimen from gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – pinnae. Scale in cm. Specimen from gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis and adaxial side of pinnules with sori. Scale in mm. Specimen from gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – lowest pinna with lowest basioscopic and acroscopic pinnules. Scale in mm. Specimen from gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – rachis scales. Scale in mm. Specimen from gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.Dryopteris borreri – sori under magnification – indusium (upper row) and sporangia (lower row). Scale in mm. The indusium is thin , shriveling and lifting in a cone-like shape. Specimen from gorge of “Kochanska” River, 20.08.2025. Photo by Trajche Dimitrovski.
References:
Dryopteris Adanson. Heywood, V.H. in Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Valentine, D.H., Walters, S.M., & Webb, D.A. (1964). Flora Europaea. Volume 1. Lycopodiaceae to Platanaceae. Cambridge: Cambridge University Press, p. 20 – 22.
Dryopteris Adanson. Heywood, V.H. (edition 1) revised by Praser-Jenkins, C.R. in Tutin, T.G., Burges, N.A. Chater, A. O., Edmondson, J. R., Heywood, V. H., Moore, D.M., Valentine, D.H., Walters, S.M. & Webb, D.A., assisted by Akeroyd, J.R. and Newton, M.E. (1993). Flora Europaea. Volume 1. Psilotaceae to Platanaceae, second edition. Cambridge: Cambridge University Press, pp. 27 – 30.
Dryopteris Adanson. Micevski, K. (1985). The Flora of SR Macedonia Vol. 1, Book 1. Macedonian Academy of Sciences and Arts, pp. 67 – 72.