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Key to the exindusiate Dryopteris species of Central Africa (D. R. Congo, Rwanda, Burundi) 1 Frond apices with scaly gemmae, stipe scales lacking glands and apex not fimbriate................................................... D. manniana - Fronds without scaly gemmae, stipe scales with glands, or apex irregularly fimbriate, divided into long and uniseriate fissures.................................................................................................................................................................................................................2 2 Lower stipe scales bent downwards, broadly ovate, apex fimbriate, divided into long uniseriate and irregular fissures, lower montane rainforest below 2000 m................................................................................................................ D. schizopaleata sp. nov. - Lower stipe scales bent upwards, ovate–lanceolate, acuminate, margin denticulate, erose, tapering into acuminate apex, upper montane forest to bamboo forest above 2000 m..................................................................................................... D. ruwenzoriensis, Published as part of Fischer, Eberhard & Lobin, Wolfram, 2023, Synoptic Revision of the exindusiate species of Dryopteris (Dryopteridaceae) from Central Africa (Democratic Republic of the Congo, Rwanda, Burundi) with description of Dryopteris schizopaleata sp. nov. from Rwanda, pp. 136-148 in Phytotaxa 587 (2) on page 138, DOI: 10.11646/phytotaxa.587.2.3, http://zenodo.org/record/7731870

Dryopteris ruwenzoriensis C.Chr. ex Fraser-Jenkins (1986: 204) (Figs. 4 C–F, 7–8). Type:—D. R. CONGO. Massif du Ruwenzori, versant Ouest (Congo Belge, Ituri), 2000 m, 22 July 1930, Mission H. Humbert Plantes de l’Afrique Equatoriale 8825ter (holotype BM000605445, digital image!). Specimens seen:—D. R. CONGO. Massif du Ruwenzori, versant Ouest (Congo Belge, Ituri), forêt ombrophile, haute vallée de la Butagu, ca. 2000 m, July 1929, H . Humbert, Plantes de l’Afrique Equatoriale 8849 (ancient 8825ter)(BR0000013468422!, possibly isotype). RWANDA. Northern Province: Volcano National Park, saddle between Karisimbi and Bisoke, 3200 m, 22 September 2021, E . Fischer s.n. (KOBL); Gahinga, bamboo forest, 3000 m, 20 March 2022, E . Fischer s.n. (KOBL). Habitat:—Montane forest to bamboo and Hagenia -Hypericum forest, elev. 2000–3200 m. First recorded for Central Africa:—D.R. Congo (Fraser-Jenkins 1986: 204), Rwanda (Roux 2012: 97). Distribution in Central Africa: —D.R. Congo (Ruwenzori massif), Rwanda (Virunga massif). Distribution in Africa:—D.R. Congo, Rwanda, Uganda (Ruwenzori). Note:— Dryopteris ruwenzoriensis is an Albertine Rift endemic restricted to the Ruwenzori massif and the Virunga massif., Published as part of Fischer, Eberhard & Lobin, Wolfram, 2023, Synoptic Revision of the exindusiate species of Dryopteris (Dryopteridaceae) from Central Africa (Democratic Republic of the Congo, Rwanda, Burundi) with description of Dryopteris schizopaleata sp. nov. from Rwanda, pp. 136-148 in Phytotaxa 587 (2) on page 142, DOI: 10.11646/phytotaxa.587.2.3, http://zenodo.org/record/7731870, {"references":["Fraser-Jenkins, C. R. (1986) A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). Bulletin of the British Museum (Natural History), Botany 14 (3): 183 - 218.","Roux, J. P. (2012) A revision of the fern genus Dryopteris (Dryopteridaceae) in sub-Saharan Africa. Phytotaxa 70: 1 - 118. https: // doi. org / 10.11646 / phytotaxa. 70.1.1"]}

Dryopteris schizopaleata Eb.Fisch. & Lobin sp. nov. (Figs. 1 E–F, 2–3, 4 A–B). Type:— RWANDA. Western Province: Nyungwe National Park, Cyamudongo Forest, forest floor along stream, 1941 m, 18 September 2021, 2°32`26.89`` S, 28°59`20.46``E, E.Fischer s.n. (holotype BR0000015253521 V!; isotype B200220601!, B200220602!, B200220603!). Diagnosis: — Dryopteris schizopaleata is exindusiate, and shares this character in Africa only with D. manniana, D. filipaleata and D. ruwenzoriensis. From D. manniana it is easily separated by the lack of gemmae.A striking character to recognise the new species are the unique scales, which are broadly and irregularly fimbriate at the apex, differing from D. filipaleata and D. ruwenzoriensis (Fig. 4). Dryopteris filipaleata has a scale apex that is irregularly denticulate, terminating in a short uniseriate series of cells. In Dryopteris ruwenzoriensis the scales apex terminates in a short uniseriate series of oblong cells. Description: —Terrestrial fern. Rhizome erect, scaly, scales broadly–ovate, dark to light brown, up to 4 × 2.3 mm, at apex deeply and irregularly fimbriate, divided into long uniseriate and irregular fissures. Fronds tufted. Stipes (22.8–)27–41.5(–57,1) × 0.3–0.5 cm, at the base thickened to 5–6 cm, densely covered with scales that are bent downward and similar to those from rhizome. Lamina 57–87 × 31–62 cm, broadly–ovate, fresh green, bipinnate-pinnatifid. Rachis middle to dark brown, scaly, winged, secondary rachis and costules sparsely set with trichomes 0.25–0.3 mm, cells 0.3 × 0.05 mm. Pinnae in 14–20 pairs, alternate, arranged in an angle of 45°, (15.2–)21–26(–36.8) × (7–)9.9–12(–14.8) cm, basal pinnae only slightly reduced. Pinnules in 15–17 pairs 3–5.4(–7.6) × (0.8–)1–1.7(–2.2) cm, opposite to subopposite; pinnule lobes in (8–)11–13 pairs, 0.5–1.2 × 0.3–0.5 cm, with rachis winged, veins branched, only in basal pinnules few dichotomosly branched. Sori up to 5–10 per pinnule, almost round, exindusiate, 0.78–0.96 × 0.63–0.82 mm. Spores with perispore folded to form narrow or compressed reticulate ridges, 32.5–34 × 25.7–27 µm, ridges 7–13 × 2–2.5 µm. Additional specimens seen (Paratype):— RWANDA. Western Province: Nyungwe National Park, Cyamudongo Forest, forest floor along stream, 1955 m, 18 April 2022, 2°32`26.89`` S, 28°59`20.46`` E, E . Fischer s.n. (KOBL). Distribution:— Rwanda, Nyungwe National Park, Cyamudongo Forest, only known from the type locality. Habitat:—Montane forest floor along stream, partly extending into buffer zone with pines, 1940–1960 m Etymology:—The name refers to the main character, the scales from the rhizome and base of stipes, with which the new species is easily recognized. Notes:— Dryopteris schizopaleata sp. nov. was collected in the Nyungwe National Park, a very species-rich area of montane rainforest, with 1468 species of ferns and seed plants recorded, of which 259 are endemic (224 Albertine Rift Endemics, 35 Local Endemics, Fischer unpubl.). The Cyamudongo Forest is only 300 ha in size, an isolated part of the Nyungwe National Park by 8 km of unprotected land. Despite this, Cyamudongo is exceptional in its beta-diversity. Here 674 species of vascular plants are recorded, including 88 endemics; 42 of these endemic species are exclusively found in Cyamudongo. Concerning the ferns and fern allies, 39.8% (57 out of 143 species) recorded from Nyungwe National Park are found in Cyamudongo Forest, and 11 species are considered endemic (species numbers after Fischer unpubl.). Two other new fern species, Asplenium uschiae Eb.Fisch. & Lobin (2023a) and Diplazium cyamudongense Eb.Fisch. & Lobin (2023b) have also been discovered in Cyamudongo. Conservation status: —With only a single collecting site, it is not possible to calculate an Extent of Occurrence (EOO) according to the IUCN criteria (IUCN 2012, 2022). The area of occupancy (AOO = 9 km 2) falls within the limits for the Critically Endangered category. However, as the species occurs within a well-protected area, and therefore to experience no threat or decline, it cannot be assessed in any threat category; therefore, it should be assessed as Least Concern (LC).

Dryopteris manniana (Hook.) Christensen (1905: 276) (Figs. 5–6). Aspidium mannianum Hook. (Hooker 1863: 253) Type:— EQUATORIAL GUINEA. Bioko: “Fernando Po, on the peak, 2000 ft, 1860”, G. Mann 351 (holotype K000675774, digital image!). Specimens seen:—D. R. CONGO. Ituri Distr, Mongbwalu Mine Site, Crack Site Pt 082 to pt 083, 25 October 2010, Q . Luke 14598 (BR0000017859837). RWANDA. Western Province: Nyungwe National Park, Territoire de Shangugu (= Rusizi), Forêt de Montagne, 14 February 1985, G . Michel 5133 (BR0000017859936); Nyungwe National Park, Cyamudongo Forest, forest floor along stream, 1941 m, 2°32`26.89`` S, 28°59`20.46`` E, December 2021, E . Fischer & B. Dumbo s.n. (KOBL). Southern Province: Nyungwe National Park, Préfecture de Gikongoro, Commune Kivu, Savane de Rubyiro-Mubuga, dans la vallée de la Tsava, 17 August 1999, C. E. N . Ewango 2200 (BR0000017859943). BURUNDI. Bururi Province: Nella foresta riparia umida della vallata del torrente Siguvyaye vicino al ponte sul torrente, 1700 m, 18 December 1966, R. E. G . Pichi Sermolli 6974 (FI067885 digital image). First recorded for Central Africa:— Pichi Sermolli (1985: 163). Distribution in Central Africa:—D.R. Congo (Ituri Distr.), Rwanda (Nyungwe NP), Burundi (Bururi). Distribution in Africa:—Disjunct distribution, occurring in Guinea, Liberia, Sierra Leone, Côte d’Ivoire, Nigeria, Cameroon, Bioko, S„o Tomé, Equatorial Guinea, D.R. Congo, Burundi, Rwanda, Uganda, Kenya, Tanzania, Malawi, Mozambique, Zimbabwe, Madagascar. Roux (2012) does not list the species for D.R. Congo. As all specimens listed by Mangambu Mokoso et al. (2013) are wrongly identified (see note below), the specimen Luke 14598 is probably the first record for this country. Habitat:—Lower montane rainforest, elev. 1190–2140 m. Note:— Dryopteris manniana is easily recognized by the scaly proliferous buds borne adaxially on the rachises, mostly near the lamina apices, the denticulate scales, and the exindusiate sori (Roux 2012). The following specimens from D.R. Congo at BR were originally identified as Dryopteris manniana, but they have well developed indusia and lack gemmae: Kisantu, Kisungu, Callens 2385, (BR0000017859820!); Parc National de Kahuzi Biéga, Mangambu Mokoso 3149, (BR0000017859912!); ibid. Mangambu Mokoso 3021 (BR0000017859851!, BR0000017859868!); ibid. Mangambu Mokoso 3025 (BR0000017859875!); ibid. Mangambu Mokoso 2756 (BR0000005279920!); ibid. Mangambu Mokoso 3124 (BR0000017859899!, BR0000017859882!); ibid. Mangambu Mokoso 2754 (BR0000005277506!); ibid. Mangambu Mokoso 3149 (BR0000017859905!); ibid. Mangambu Mokoso 2456 (BR0000017859844!); ibid. Mangambu Mokoso 2749 (BR0000005277308!). They all represent taxa of the Dryopteris pentheri -group sensu Roux (2009) and will be dealt with in a separate paper. Therefore, the new record of D. manniana for the D.R. Congo (Mangambu Mokoso et al. 2013) is most probably erroneous. A specimen lacking indusia, but also gemmae, has been collected in Haut-Katanga:PN Kundelungu, Nkonko-chutes Kaloba (pied), 1e halte: rivière Kibila, 19 May 1985, Bodenghien, Goetghebeur. & Malaisse 982 (BR0000017859929!). It cannot be identified due to the scarcity of the material., Published as part of Fischer, Eberhard & Lobin, Wolfram, 2023, Synoptic Revision of the exindusiate species of Dryopteris (Dryopteridaceae) from Central Africa (Democratic Republic of the Congo, Rwanda, Burundi) with description of Dryopteris schizopaleata sp. nov. from Rwanda, pp. 136-148 in Phytotaxa 587 (2) on page 142, DOI: 10.11646/phytotaxa.587.2.3, http://zenodo.org/record/7731870, {"references":["Christensen, C. (1905) Index Filicum sive Enumeratio omnium generum specierumque Filicum et Hydropteridum ab anno 1753 ad finem annum 1905 descriptorum. Copenhagen, Hagerup, I - LIX, 745 pp. https: // doi. org / 10.5962 / bhl. title. 402","Hooker, W. J. (1863) Species filicum 4 (15 / 16). Pamplin, London, pp. 161 - 292, pls 250 - 280.","Pichi Sermolli, R. E. G. (1985) A contribution to the knowledge of the pteridophyta of Rwanda, Burundi, and Kivu (Zaire) II. Bulletin du Jardin botanique National de Belgique / Bulletin van de Nationale Plantentuin van Belgie 55 (1 / 2): 123 - 206. https: // doi. org / 10.2307 / 3668014","Roux, J. P. (2012) A revision of the fern genus Dryopteris (Dryopteridaceae) in sub-Saharan Africa. Phytotaxa 70: 1 - 118. https: // doi. org / 10.11646 / phytotaxa. 70.1.1","Mangambu Mokoso, J. de D., Diggelen, R. v., Mwanga Mwanga, J. C., Ntahobavuka, H. & Robbrecht, E. (2013) Especes nouvellement signalees pour la flore pteridologique de la Republique Democratique du Congo. International Journal of Biological and Chemical Sciences 7 (1): 107 - 124. https: // doi. org / 10.4314 / ijbcs. v 7 i 1.10","Roux, J. P. (2009) Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23: 296 pp."]}

Dryopteris sect. Diclisodon is a small section of ferns with about 12 species mainly distributed in East Asia. Here, we carried out morphological and phylogenetic analyses of this section. A new species from southwest China, D. gaoligongensis, is described and illustrated. Dryopteris gaoligongensis resembles D. indonesiana and D. sparsa, but differs by having a creeping rhizome and large 4-pinnate fronds. We also show that D. glabrior Ching & Z.Y. Liu is a distinct species; however, because it is a later homonym of D. glabrior Copeland, it should be renamed D. renchangiana. We conclude that a species previously known as D. nitidula, also an illegitimate homonym, should be recognized with a new name, D. sinonepalensis. We resolve the phylogenetic position of D. yoroii as sister to other sampled species of D. sect. Diclisodon. Our phylogenetic analyses confirm the distinctiveness of D. gaoligongensis, D. renchangiana, and D. sinonepalensis. A key to species of D. sect. Diclisodon in China is provided.

Dryopteris lijianxiuii X. J. Li, a new species of Dryopteris Adans. from Shandong, China, is described and illustrated. It is closer to D. lacera (Thunb.) O. Kuntze for its morphological and palynological characters of LM and SEM studies but differs by a number of characters, such as, the leaves rectangular roundlanceolate, apex acuminate, and without narrow sharply; sori ornamented in the upper or middle part of 1-3 pairs of pinnaes at the base of the leaf, and densely covered with the upper and middle pinnaes back; spore perispore with tuberculiform-rugulate protrusions, and surfaces with melting ice and snow ornamentation. The sp. nov. is described with LM and SEM characters of spore and fronds, compared with that of D. lacera (Thunb.) O. Kuntze and provided with photographs. Bangladesh J. Bot. 50(3): 747-753, 2021 (September) Special

Dryopteris pentheri (Krasser) C.Chr. — Habit: Fern. Habitat: LMWF, LMDF; up to 2 700 m. Distribution: II. Voucher: Castle Forest Station, Alt. 2050– 2100 m, 30 Nov. 1971, Faden et al. 71/898 (EA). Reference: Bussmann (1993)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 20, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Bussmann, R. W. (1993) The forest of Mt. Kenya (Kenya): A phytosociological approach with special reference to ecological problems. Ph. D. Thesis, Eberhard Karls Universitat Tubingen, Tubingen, 119 pp."]}

Dryopteris manniana (Hook.) C.Chr. — Habit: Fern. Habitat: LMWF; 1 400–2 200 m. Distribution: II. Voucher: Irangi Forest Office area, Alt. 2000 m, 9 Aug. 1978, Zogg et al. 276/10 (EA). References: Bussmann (1993, 1994), Bussmann & Beck (1995a, 1998, 1999), Agnew (2013)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 20, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Bussmann, R. W. (1993) The forest of Mt. Kenya (Kenya): A phytosociological approach with special reference to ecological problems. Ph. D. Thesis, Eberhard Karls Universitat Tubingen, Tubingen, 119 pp.","Bussmann, R. W. (1994) The forest of Mt. Kenya (Kenya): Vegetation, ecology, destruction and management of a tropical mountain forest ecosystem. Ph. D. dissertation, Universitat Bayreuth Startseite, Bayreuth, 252 pp.","Bussmann, R. W. & Beck, E. (1995 a) The forests of Mt. Kenya (Kenya), a phytosociological synopsis. Phytocoenologia 25 (4): 467 - 560. https: // doi. org / 10.1127 / phyto / 25 / 1995 / 467","Bussmann, R. W. & Beck, E. (1998) The plant communities of the forests of Mount Kenya - First phytosociological and ecological data. In: Ojany, F. F. (Ed.) African mountains and highlands - planning for sustainable use of mountain resources. United Nations University, Tokyo, pp. 96 - 107.","Bussmann, R. W. & Beck, E. (1999) Vegetation units of Mt. Kenya Forest Reserve: Comment. In: Bitok (Ed.) Vegetations-okologosche, ethnobotanische und faunistische Beitrage uber Aquatorial-Afrika. Bayreuther Forum Okologie 64. Bayreuther Forum Okologie, Bayreuth, pp. 17 - 28.","Agnew, A. D. Q. (2013) Upland Kenya wild flowers and ferns, 3 rd edn. Nature Kenya Publications, Nairobi, 733 pp."]}

Dryopteris antarctica (Baker) C.Chr. — Habit: Fern. Habitat: BZ, UMF, HZ; 2 500–3 200 m. Distribution: II. Voucher: Northeast Mount Kenya, Kirui Volcanic Cone, on the slopes of Ithanguni, Alt. 2 530 m, 28 Feb. 1970, Faden & Evans 70/105 (EA). References: Schelpe (1951), Bussmann (1993, 1994), Roux et al. (2007), Agnew (2013)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 19, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Schelpe, E. A. C. L. E. (1951) The Pteridophyta of Mount Kenya. American Fern Journal 41 (3): 65 - 74. https: // doi. org / 10.2307 / 1545044","Bussmann, R. W. (1993) The forest of Mt. Kenya (Kenya): A phytosociological approach with special reference to ecological problems. Ph. D. Thesis, Eberhard Karls Universitat Tubingen, Tubingen, 119 pp.","Bussmann, R. W. (1994) The forest of Mt. Kenya (Kenya): Vegetation, ecology, destruction and management of a tropical mountain forest ecosystem. Ph. D. dissertation, Universitat Bayreuth Startseite, Bayreuth, 252 pp.","Roux, J. P., Shaffer-Fehre, M. & Verdcourt, B. (2007) Dryopteridaceae. In: Beentje, H. J. & Ghazanfar, S. A. (Eds.) Flora of Tropical East Africa. Royal Botanic Gardens, Kew, London, 54 pp.","Agnew, A. D. Q. (2013) Upland Kenya wild flowers and ferns, 3 rd edn. Nature Kenya Publications, Nairobi, 733 pp."]}

Dryopteris schimperiana (Hochst.) C.Chr. — Habit: Fern. Habitat:LMDF, BZ, UMF; 1 400–3 000 m.Distribution: II. Voucher: N/A. Reference: Fries et al. (1924b)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 20, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Fries, R. E., Fries, T. C. E. & Christensen, C. (1924 b) Beitrage zur Kenntnis der Flora des Kenia, Mt. Aberdare und Mt. Elgon. VI. Notizblatt des Koniglichen Botanischen Gartens und Museums zu Berlin 9 (83): 173 - 189. https: // doi. org / 10.2307 / 3994334"]}

Dryopteris rodolfii J.P.Roux — Habit: Fern. Habitat: LMDF, BZ; 2 700–3 000 m. Distribution: II. Voucher: N/A. References: Roux et al. (2007), Agnew (2013)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 20, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Roux, J. P., Shaffer-Fehre, M. & Verdcourt, B. (2007) Dryopteridaceae. In: Beentje, H. J. & Ghazanfar, S. A. (Eds.) Flora of Tropical East Africa. Royal Botanic Gardens, Kew, London, 54 pp.","Agnew, A. D. Q. (2013) Upland Kenya wild flowers and ferns, 3 rd edn. Nature Kenya Publications, Nairobi, 733 pp."]}

Dryopteris filipaleata J.P.Roux — Habit: Fern. Habitat: LMWF; up to 2 000 m. Distribution: IIIa. Voucher: N/A. References: Roux et al. (2007), Agnew (2013)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 20, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Roux, J. P., Shaffer-Fehre, M. & Verdcourt, B. (2007) Dryopteridaceae. In: Beentje, H. J. & Ghazanfar, S. A. (Eds.) Flora of Tropical East Africa. Royal Botanic Gardens, Kew, London, 54 pp.","Agnew, A. D. Q. (2013) Upland Kenya wild flowers and ferns, 3 rd edn. Nature Kenya Publications, Nairobi, 733 pp."]}

Dryopteris fadenii Pic.Serm. — Habit: Fern. Habitat: LMWF, BZ; 1 700–2 500 m. Distribution: IIIa. Voucher: N/A. References: Bussmann (1994), Roux et al. (2007), Agnew (2013)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 20, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Bussmann, R. W. (1994) The forest of Mt. Kenya (Kenya): Vegetation, ecology, destruction and management of a tropical mountain forest ecosystem. Ph. D. dissertation, Universitat Bayreuth Startseite, Bayreuth, 252 pp.","Roux, J. P., Shaffer-Fehre, M. & Verdcourt, B. (2007) Dryopteridaceae. In: Beentje, H. J. & Ghazanfar, S. A. (Eds.) Flora of Tropical East Africa. Royal Botanic Gardens, Kew, London, 54 pp.","Agnew, A. D. Q. (2013) Upland Kenya wild flowers and ferns, 3 rd edn. Nature Kenya Publications, Nairobi, 733 pp."]}

Dryopteris tricellularis J.P.Roux — Habit: Fern. Habitat: LMWF, LMDF, BZ, UMF, HZ; 1 700–3 600 m. Distribution: II. Voucher: Chogoria Track, Alt. 2 572 m, 22 Jul. 2014, CPG 28574 (HIB, PE). References: Fries et al. (1924b), Roux et al. (2007), Agnew (2013)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 20, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Fries, R. E., Fries, T. C. E. & Christensen, C. (1924 b) Beitrage zur Kenntnis der Flora des Kenia, Mt. Aberdare und Mt. Elgon. VI. Notizblatt des Koniglichen Botanischen Gartens und Museums zu Berlin 9 (83): 173 - 189. https: // doi. org / 10.2307 / 3994334","Roux, J. P., Shaffer-Fehre, M. & Verdcourt, B. (2007) Dryopteridaceae. In: Beentje, H. J. & Ghazanfar, S. A. (Eds.) Flora of Tropical East Africa. Royal Botanic Gardens, Kew, London, 54 pp.","Agnew, A. D. Q. (2013) Upland Kenya wild flowers and ferns, 3 rd edn. Nature Kenya Publications, Nairobi, 733 pp."]}

Dryopteris kilemensis (Kuhn) Kuntze — Habit: Fern. Habitat: LMWF, LMDF, BZ; 1 800–2 700 m. Distribution: II. Voucher: Castle Forest Station, Alt. 2050 m, 20 Oct. 1979, Gilbert 5774 (EA). References: Fries et al. (1924b), Fries & Fries (1948), Bussmann & Beck (1995a, 1995c, 1998, 1999), Bussmann (1993, 1994, 2006), Roux et al. (2007), Agnew (2013)., Published as part of Zhou, Ya-Dong, Mwachala, Geoffrey, Hu, Guang-Wan & Wang, Qing-Feng, 2022, Annotated checklist of the vascular plants of Mount Kenya, East Africa, pp. 1-108 in Phytotaxa 546 (1) on page 20, DOI: 10.11646/phytotaxa.546.1.1, http://zenodo.org/record/6550464, {"references":["Fries, R. E., Fries, T. C. E. & Christensen, C. (1924 b) Beitrage zur Kenntnis der Flora des Kenia, Mt. Aberdare und Mt. Elgon. VI. Notizblatt des Koniglichen Botanischen Gartens und Museums zu Berlin 9 (83): 173 - 189. https: // doi. org / 10.2307 / 3994334","Fries, R. E. & Fries, T. C. E. (1948) Phytogeographical researches on Mt. Kenya and Mt. Aberdare, British East Africa. Kungliga Svenska Vetenskapsakademiens Handlingar, ser. 3, 25 (5): 1 - 83.","Bussmann, R. W. & Beck, E. (1995 a) The forests of Mt. Kenya (Kenya), a phytosociological synopsis. Phytocoenologia 25 (4): 467 - 560. https: // doi. org / 10.1127 / phyto / 25 / 1995 / 467","Bussmann, R. W. & Beck, E. (1995 c) Regeneration and cyclic processes in the Ocotea-Forest (Ocotea usambarensis Engl.) of Mt. Kenya. Verhandlungen der Gesellschaft fur Okologie 24: 35 - 39.","Bussmann, R. W. & Beck, E. (1998) The plant communities of the forests of Mount Kenya - First phytosociological and ecological data. In: Ojany, F. F. (Ed.) African mountains and highlands - planning for sustainable use of mountain resources. United Nations University, Tokyo, pp. 96 - 107.","Bussmann, R. W. & Beck, E. (1999) Vegetation units of Mt. Kenya Forest Reserve: Comment. In: Bitok (Ed.) Vegetations-okologosche, ethnobotanische und faunistische Beitrage uber Aquatorial-Afrika. Bayreuther Forum Okologie 64. Bayreuther Forum Okologie, Bayreuth, pp. 17 - 28.","Bussmann, R. W. (1993) The forest of Mt. Kenya (Kenya): A phytosociological approach with special reference to ecological problems. Ph. D. Thesis, Eberhard Karls Universitat Tubingen, Tubingen, 119 pp.","Bussmann, R. W. (1994) The forest of Mt. Kenya (Kenya): Vegetation, ecology, destruction and management of a tropical mountain forest ecosystem. Ph. D. dissertation, Universitat Bayreuth Startseite, Bayreuth, 252 pp.","Bussmann, R. W. (2006) Vegetation zonation and nomenclature of African Mountains: An overview. Lyonia 11 (1): 41 - 46.","Roux, J. P., Shaffer-Fehre, M. & Verdcourt, B. (2007) Dryopteridaceae. In: Beentje, H. J. & Ghazanfar, S. A. (Eds.) Flora of Tropical East Africa. Royal Botanic Gardens, Kew, London, 54 pp.","Agnew, A. D. Q. (2013) Upland Kenya wild flowers and ferns, 3 rd edn. Nature Kenya Publications, Nairobi, 733 pp."]}

The long history of using plant species as medicines worldwide provides clues for the search for bioactive phytochemicals. One of the important traditional medicinal plant species of Pakistan is Dryopteris ramosa (Dryopteridaceae). The fronds of Dryopteris ramosa are traditionally used for the treatment of gastric ulcer and constipation. The crude methanol extract and aqueous fraction of Dryopteris ramosa exhibited strong free radical scavenging potential. Two xanthonoid compounds, mangiferin and isomangiferin, were isolated from the aqueous fraction of Dryopteris ramosa by chromatographic techniques (CC Sephadex LH20, followed by MPLC and purified by preparative TLC). Their structures were confirmed by spectroscopic analysis MS, NMR (1H, 13C, HSQC and HMBC). This is the first report on isolation of mangiferin and isomangiferin from Dryopteris genus though previously reported in other fern genera, e.g., Cystopteris, Acystopteris, Gymnocarpium and Asplenium. The antioxidant potential of mangiferin (60.34 ± 0.14 μg/mL) and isomangiferin (55.98 ± 0.39 μg/mL) was comparable to ascorbic acid (57.97 ± 0.93 μg/mL), thus representing promising antioxidant agents against oxidative stress. The investigation of the mechanism of free radical scavenging by mangiferin and isomangiferin showed that the catechol moiety was involved in quenching of free radicals. This study justified the traditional use of Dryopteris ramosa confirming the pharmacological properties of isolated compounds. Further phytochemical studies on Dryopteris ramosa are needed for identification of new bioactive phytochemicals in this plant of ethnobotanical relevance.

Dryopteris sukungiana Z. Y. Zuo, sp. nov. (Figs. 1 & 2), in Chinese ���Dzfl����� (y��n n��n l��n m��o ju��). Type:��� CHINA. Yunnan: Maguan, 103��59��� E / 22��50��� N, 1950 m alt., 22 June 2021, Z. Y. Zuo 4660 (holotype KUN-1519949; isotypes HITBC-0061304, KUN-1519950, PE-02354809). Diagnosis:��� The scales of Dryopteris sukungiana are blackish brown and spreading (Fig. 3E), which is different from all other species in the D. sparsa complex (Fig. 3D, 3F). Dryopteris sukungiana is similar to the ���typical��� Sino-Himalayan D. sparsa with erect rhizomes, deltoid or pentagonal tripinnatifid to tripinnate lamina, basal pinnae abruptly broadened toward the base (Fig. 3A, 3B), but differs in having glandular hairs and small fibrillose scales on the abaxial surface of the lamina and the indusia (Fig. 3G, 3H). It resembles the Sino-Japanese D. sparsa in the catadromous pinnules of the second pair of pinnae and glandular hairs on the indusia and the abaxial surface of the lamina (Fig. 3H, 3I); however, the lamina of the Sino-Japanese D. sparsa is ovate-deltoid, and the basal pinnae are gradually broadened (Fig. 3C). Dryopteris sukungiana also looks like D. angustipalea Darnaedi, M. Kato & K. Iwats. (1989a: 308); however, the latter is endemic to Indonesia, and in which the lamina is ovate-deltoid, the basal pinnae are gradually broadened toward the base, rachis and costae are glabrous. Plants 30���60 cm tall. Rhizome erect, up to 8 cm long and 2 cm in diameter, clothed with blackish brown, linear- or ovate-lanceolate, spreading, entire scales. Stipe slightly shorter than lamina, ca. 17���38 cm, covered with glandular hairs when young, brown at base, with blackish brown linear- or ovate-lanceolate, spreading, entire, scales. Lamina papery, glossy, deltoid or pentagonal, ca. 26���53 �� 18���35 cm, tripinnatifid to tripinnate, base not narrowed, apex acuminate, with glandular hairs on abaxial surface. Pinnae 9���17 pairs, deltate-lanceolate, up to 16 �� 6 cm, apex caudate-acuminate; Basal 1���3 pairs opposite and alternate upward, oblique, stalked approximately 1���2 cm. Basal pinnae abruptly broadened toward the base. Pinnules 9���23 pairs, margin serrate to lobed, opposite at base and alternate upward, lanceolate to rounded, base broadly cuneate, asymmetrical, apex obtuse to truncate. Basal basiscopic pinnules of basal pinnae longest, ca. 5���8 �� 2���3 cm, base widest, 2-pinnate. Pinnules of the second pair of pinnae catadromous. Rachis and costa clothed with glandular hairs and small fibrillose scales. Veins pinnate, forked, distinct on both surfaces. Sori close to costa on pinnules, with a wide sterile belt on both sides of costa distal to sori. Indusia orbicularreniform, entire, with glandular hairs. Reproductive mode and ploidy level: sexual tetraploid. Additional specimens examined (paratypes): ��� CHINA. Yunnan: Baoshan, Z. Y. Zuo 1615 & 4937; Cangyuan, Z. Y. Zuo 2862; Kunming, J. M. Lu 179; Longling, X. Cheng et al., FB581; Tengchong, Z. Y. Zuo 1498, 1520 & 4949. Ecology and geographical distribution: ��� Dryopteris sukungiana was found in mountain forests at alt. 1900��� 2450 m in the central, southern, and southwestern parts of Yunnan Province, China, and perhaps in other neighboring areas. Etymology:��� In honor of late Prof. Su-Gong Wu (Su-Kung Wu in Wide-Giles transliteration) (1935-2013), for his contributions to the taxonomic studies on ferns in China, and especially the contributions to collections and studies on the ferns of the Qinghai-Tibet Plateau. He also edited some sections of Dryopteris in Flora Reipublicae Popularis Sinicae and Flora of China., Published as part of Zuo, Zheng-Yu, Lu, Jin-Mei, Wang, Yue-Hua & Li, De-Zhu, 2022, Dryopteris sukungiana (Dryopteridaceae), a new species of the D. sparsa complex from Southwest China, pp. 256-266 in Phytotaxa 533 (5) on pages 260-264, DOI: 10.11646/phytotaxa.533.5.2, http://zenodo.org/record/6144962

Premise Hybridization is a key process in plant speciation. Despite its importance, there is no detailed study of hybridization rates in fern populations. A proper estimate of hybridization rates is needed to understand factors regulating hybridization. Methods We studied hybridization in the European Dryopteris carthusiana group, represented by one diploid and two tetraploid species and their hybrids. We sampled ~100 individuals per population in 40 mixed populations of the D. carthusiana group across Europe. All plants were identified by measuring genome size (DAPI staining) using flow cytometry. To determine the maternal parentage of hybrids, we sequenced the chloroplast region trnL-trnF of all taxa involved. Results We found hybrids in 85% of populations. Triploid D. ×ambroseae occurred in every population that included both parent species and is most abundant when the parent species are equally abundant. By contrast, tetraploid D. ×deweveri was rare (15 individuals total) and triploid D. ×sarvelae was absent. The parentage of hybrid taxa is asymmetric. Despite expectations from previous studies, tetraploid D. dilatata is the predominant male parent of its triploid hybrid. Conclusions This is a thorough investigation of hybridization rates in natural populations of ferns. Hybridization rates differ greatly even among closely related fern taxa. In contrast to angiosperms, our data suggest that hybridization rates are highest in balanced parent populations and support the notion that some ferns possess very weak barriers to hybridization. Our results from sequencing cpDNA challenge established notions about the correlation of ploidy level and mating tendencies.

Here we investigate the morphology of the spores of 12 native taxa of Dryopteridaceae that grow in Malakand division, Pakistan; most of these species not fully described before. This study intends to survey the taxonomic significance of spore morphological features and their variation useful for species identification in 7 species of Polystichum and 5 taxa of Dryopteris. The spores examinations were accomplished utilizing a light microscope (LM), and scanning electron microscope (SEM). The spores are monolete, medium-sized, with variable shapes in both equatorial and polar views. The mean estimation of equatorial diameter ranging from 28.3 to 58.3 μm and the polar diameter varied from 27.6 to 45.8 μm. The exospore thickness ranges between 1.6 to 3.8 μm, and the thickness of perispore is 0.8-6.7 μm. The perispore is perforated, ornamentation elements occur in different ways in the studied taxa of Polystichum. In investigated Dryopteris taxa, rugate perispore type is found. Results of multivariate statistical analysis (UPGMA, and PCA) established on quantitative and qualitative morphological traits of spores demonstrate that the species form distinct groups, not reflecting any phylogenetic relationships. A key to the spores types is provided for species identification, which dependent on the diagnostic characters of spores. The results of this study indicate that variation of spore morphological traits; in particular, the ornamentation and perispore characteristics, which is surface texture, have diagnostic value at the species level. Our data will help in the identification of spores of the family Dryopteridaceae in palynological and taxonomic studies.

Dryopteris wulingshanensis, a new species growing on limestone in the Wulingshan Mountains, Hunan, China, is described and illustrated. This species is most similar to D. jishouensis and D. gymnophylla on general morphological traits, such as the form of scales, rhizome and sori, but differs by the number of vascular bundles at the base of the petiole, length to width ratio of lamina, stalk length of basal pinnae, division of the lamina, apex form of the pinnule and habitat. Moreover, molecular phylogenetic analysis using the chloroplast rbcL gene suggested that D. wulingshanensis, as the sister group of D. jishouensis, is a monophyletic clade. According to its restricted geographic range, small populations and few individuals, D. wulingshanensis should be considered endangered, according to the IUCN Red List criteria.

The complete mitochondrial genome of Dryopteris crassirhizoma was sequenced for the first time. The mitochondrial genome length was 313,346 bp, with 48.58% GC contents. There were 94 genes annotated, including 27 known protein-coding genes, 49 tRNAs, and 18 rRNAs. The maximum likelihood method was used to establish the phylogenetic tree of six species. The phylogenetic results showed that D. crassirhizoma was sister to Ophioglossum californicum. It reveals the genetic relationship between different species and provides a theoretical basis for the establishment of a classification system.

The spore morphology and wall ultrastructure of Dryopteris filix-mas, D. patula and D. wallichiana from Argentina were studied using light microscope, scanning and transmission electron microscope. The study was carried out with herbarium material from Argentine institutions. Equatorial diameters, polar diameters and laesura length were measured. The spores are monoletes with a rugate ornamentation. The folds are short to long, inflate, irregular in shape and size, and varying from subglobose to elongate. The perispore surface is rugulate. The exospore of all the species analyzed is two-layered in section. Simple and branched channels are also present. The perispore is composed of two layers, the inner one forms the ornamentation and the outer covers all the outer and inner surfaces. Some abnormalities, such as globose, triangular or twisted spores were observed. The morphology and ultrastructure of the species are very similar. The differences observed are related to the length and thickness of the perispore folds. The characteristics of these spores would not provide relevant information to differentiate species or sections within the genus, but can provide information for phylogenetic studies as well as for alterations in the biological cycles., Facultad de Ciencias Naturales y Museo

The presence of 267 vascular plants species from 190 genera, 63 families, 26 orders, 4 classes (Equisetopsida, Polypodiopsida, Pinopsida, Angiospermae) and 3 divisions (Equisetophyta, Polypodiophyta, Spermatophyta) was established for Kuznetsov Mountain. The absolute majority of species belong to Spermatophyta (258 species, 96,7%), Angiospermae (257 species, 96,3%), Dicotyledones (228 species, 85,4%). Vascular spore plants are represented by 9 species belonging to 6 genera ( Equisetum , Matteuccia , Athyrium , Cystopteris , Dryopteris , Pteridium ), 5 families (Equisetaceae, Onocleaceae, Woodsiaceae, Dryopteridaceae, Dennstaedtiaceae), 2 classes (Equisetopsida, Polypodiopsida) and 2 divisions (Equisetophyta, Polypodiophyta). The leading families according the species diversity are Compositae (44 species, 16,5%), Leguminosae (22 species, 8,2%) and Rosaceae (18 species, 6,7%). The leading families according the number of genera are Compositae (29 genera, 15,3%), Rosaceae (14 genera, 7,4%) and Labiatae (13 genera, 6,8%). The largest number of species is concentrated in the genera Trifolium (7 species), Viola , Campanula , Galium and Carex (4 species each). According to the climamorphic system of K. Raunkiaer, hemicryptophytes predominate in the flora (148 species, 55,4%). According to the biomorph system I.G. Serebryakov and T.I. Serebryakova - perennial polycarpic short-stemmed, long-rooted, and rod-root grasses (47,0%) predominate. 76,8% of the species are vegetatively still and immobile.

Dryopteris blanfordii (C.Hope) C.Chr. is a member of the Dryopteridaceae, growing in high altitude Picea or Abies forests (2900–3500 m) in China and India. Phylogenetic relationships between D. blanfordii subsp. nigrosquamosa and closely related species of Dryopteris were investigated using a combined analysis of multiple molecular data sets (the protein-coding region of rbcL and matK genes and intergenic spacers psbA-trnH, trnP-petG, rps4-trnS, trnL-trnF and rbcL-accD). An assumption about the position of D. blanfordii subsp. nigrosquamosa within Dryopteris was made by using the Maximum Likelihood and Bayesian Inference approach and chloroplast marker sequences of Dryopteris species from GenBank. The results demonstrated that Asian taxa D. blanfordii subsp. nigrosquamosa and D. laeta as well as two American species D. arguta and D. marginalis belong to the same clade, all four of them being part of Dryopteris section Dryopteris.