Polish Botanical Journal 58(1): 117–125, 2013 DOI: 10.2478/pbj-2013-0013 Two new Fissidens species (FissidenTaceae) From Tropical easTern aFrica and Fissidens bessouensis corb., a remarkable species wiTh FimbriaTe leaves and mulTicellular spores1 Maria alida BruggeMan-nannenga abstract. Fissidens acrophilus Bruggeman-Nannenga, sp. nov. (subgenus Fissidens) from Kenya and Uganda and F. obscuri- frons Bruggeman-Nannenga, sp. nov. (subgenus Fissidens?) from Tanzania are newly described and figured. F. obscurifrons is compared to the little known Madeiran F. nobreganus Luisier & P. de la Varde. A revision of F. bessouensis Corb. (subgenus Octodiceras) with description, diagnosis and figures and a discussion including adaptations to an aquatic habitat is presented. Multicellular spores and fimbriate leaf margins are reported in the Fissidentaceae for the first time. Rectification: F. anguste- limbatus Mitt. is eliminated from the African flora. key words: Taxonomy, Africa, Fissidens acrophilus spec. nov., Fissidens obscurifrons spec. nov., Fissidens nobreganus, Fis- sidens bessouensis, fimbriate leaves and multicellular spores in Fissidens Maria Alida Bruggeman-Nannenga, Griffensteijnseplein 23, 3703 BE Zeist, The Netherlands While preparing a paper on subgenus Fissidens 3960 m. 17 January 1992. Kungu 9222/AD (holoType: in tropical eastern Africa two new species were L; isoType: EGR). discovered. Both are from wet habitats. Fissidens descripTion (perigonial data from paratype). acrophilus Bruggeman-Nannenga, sp. nov. is a lim- Growing in mats; stem with central strand of bate species with smooth cells found on Mt. Elgon ± 10 cells, with 1–3 tiers of inner and 1–3 tiers (Kenya and Uganda) between 3250 and 3960 m. of outer cortical cells; rhizoids basal and frequently Fissidens obscurifrons Bruggeman-Nannenga, sp. axillary, brown, smooth; axillary nodules weakly nov. has smooth cells and partly limbate leaves. It differentiated; perichaetial stems shorter than veg- was collected by Tamás Pócs and collaborators in etative stems; vegetative stems branched or not, the Nguru and S-Pare Mts in Tanzania and grows 8.0–10.0 × 2.0–2.5 mm, pinnate; leaves pale green, between 700 and 1000 m. The last part of this paper older leaves often with dark costa and limbidia, is a revision of the West African F. bessouensis distant to partly overlapping, up to 20 pairs, hardly Corb. (subgenus Octodiceras). This remarkable crispate when dry, broadly lanceolate, acute-acu- species has two features that are to my knowledge minate to acute-mucronate, 1.10–1.30 × 0.40–0.45 hitherto unknown in the Fissidentaceae: fimbriate mm, L/W 3, limbate; limbidium ± reaching the leaf leaf margins and multicellular spores. apex, confluent or not; confluent at the apex of the vaginant lamina, reaching the insertion of the vagi- Fissidens acrophilus Bruggeman-Nannenga, nant lamina; reaching the insertion of the dorsal sp. nov. Fig. 1 lamina, becoming indistinct on the decurrency, in Type: KENYA. Mount Elgon National Park, at the mid of dorsal lamina 24 µm wide, 1–2 stratose; Suam River sources 1 km WSW of Koitoboss summit. on vaginant laminae to 6 cells wide, to 38.5 µm Carex runssorensis-Alchemilla bog, on peaty soil. wide, 1–2 stratose, marginal; vaginant lamina 4/5 the leaf length, slightly open to closed, at the base less wide to about as wide as the stem, slightly 1 With pleasure I dedicate this paper to Professor Tamás Pócs for his 80th birthday. rounded at insertion, not decurrent, unistratose; 118 POLISH BOTANICAL JOURNAL 58(1). 2013 dorsal lamina narrow, slightly rounded at base, lanceolate, acute-acuminate to acute-mucronate reaching the insertion, often slightly decurrent, leaves with per- to excurrent costae and large, dorsal and apical lamina unistratose; costa per- 10.0–19.0 × 7.0–11.5 µm, laminal cells; it has to excurrent; mid dorsal laminal cells hexagonal, long, pinnate stems with basal clusters of short, plane to slightly convex, 10–18 × 7–10 µm, frondiform perichaetial stems and short, axil- smooth; mid vaginant laminal cells 10.0–20.5 × lary perichaetial and perigonial branches; and 5.0–11.0 μm; gemmae not observed. short peristomal filaments. Variability: the type Fertile parts. Cladautoicous, perigonia ter- specimen differs from the other collection in the minal on short, axillary branches, budlike or relatively long vaginant laminae. Fissidens acro- slender, 0.3–0.7 mm long, antheridia 200 µm philus can be confused with large expressions of long; perichaetia terminal on short stems and F. curvatus. This species can be distinguished by short branches; perichaetial stems, 2–3 × 1 mm, its limbidia that are confluent at the apex and its unbranched, frondiform, in clusters at the base of narrower, stiff vegetative leaves with dorsal lam- vegetative stems; leaves crowded, up to 5 pairs, inae that are not decurrent. Fissidens androgynus perichaetial leaves up to 1.7 mm long; arche- can be separated by its terminal antheridia and gonia ± 300 µm long; sporophyte, seta 6 mm archegonia. The type-specimen was earlier re- long, smooth; capsule symmetrical to slightly ported as F. anguste-limbatus Mitt. (Bruggeman- oblique, 0.7–0.9 × 0.5 mm, ± 64 columns of ob- Nannenga 2009); indeed, it resembles this species long exothecial cells; peristome bryoides type, in the shape of the leaves, and the dark brown teeth 45.0–49.5 µm wide at base, relatively short, coloring of cell walls, costae and limbidia in older 250 μm long; operculum 0.45 mm long; calyptra specimens. However, F. anguste-limbatus has not seen; spores subglobose, 16.5–19.5 µm, no central strand, lacks axillary nodules and has coarsely papillose, green. archegonial plants that are similar to vegetative stems. Moreover, it is typically found at lower eTyMology. The name acrophilus is de- elevations, 0 and 1200 m, though once at 3200 rived from the Greek words ακρoν (topmost) and m (Pursell 2007). φίλoσ (loving) indicating that it grows at high The new species belongs in subgenus Fissidens altitudes. as it has limbate leaves, smooth cells and a bry- suBsTraTe. On peaty soil, wet rocks. oides type peristome. haBiTaT. Carex runssorensis-Alchemilla bog; afro-alpine zone unspecified. Fissidens obscurifrons Bruggeman-Nannenga, sp. nov. Fig. 2 alTiTude. 3250–3960 m. Type: TANZANIA. Nguru Mts, Kanga Forest Re- disTriBuTion. So far known only from Mt serve, on the SE slope of Mt Kanga. Lowland rainforest, Elgon (Kenya and Uganda). on irrigated rocks under small waterfall. 700–800 m. paraType: UGANDA, Mt Elgon, wet rocks in afro- 4 December 1987. Pócs, Nsolo & Temu 87226/AL alpine zone, on wet soil. Wesche 9085 (private herbarium (holoType: L; isoType: EGR). Bruggeman-Nannenga). Growing from a dense mat of rhizoids, thalloid This new species is known from high altitudes protonemata (Fig. 2: 11) present or not; central in Kenya and Uganda. It is characterized by its strand weak or lacking, with 1–2 tiers of inner and pale green, limbate, slightly decurrent, broadly 2 tiers of outer cortical cells; rhizoids basal, bright Fig. 1. Fissidens acrophilus Bruggeman-Nannenga, sp. nov. 1 – vegetative stem with small, basal perichaetial stem; 2 – vegeta- tive stem; 3 – sporophytic stem; 4 – short perichaetial stem (detail from 1); 5–8 – leaves; 9 – perichaetial leaf; 10–12 – leaf apices; 13 – mid leaf; 14b & 15 – basal parts of leaf with decurrent dorsal lamina; 16 – axillary nodule. 1–4, 7–10 and 12–14 from holotype Kungu 9222/AD (holotype: L); 5–6, 11, 15, 16 from Wesche 9085 (paratype: private herbarium Bruggeman- Nannenga). M. A. BRUGGEMAN-NANNENGA: TWO NEW SPECIES OF FissiDENs 119 120 POLISH BOTANICAL JOURNAL 58(1). 2013 brown, smooth, up to 33 μm wide; axillary cells This new species is only known by its vegeta- weakly differentiated; stems unbranched, 1.5–3.5 tive state. It is characterized by typically narrow, × 0.8–1.7 mm, pinnate; leaves distant to close, obscure, elliptical, elliptico-lanceolate to oblanceo- up to 13 pairs, hardly altered when dry, obscure, late leaves; limbate vaginant laminae with limbidia elliptical, elliptico-lanceolate and oblanceolate, extending up to 150 μm onto the apical laminae acute, 0.5–1.0 × 0.1–0.6 mm, L/W 3–9, margin limbate; dorsal laminae frequently elimbate. Both subentire, partly limbate, poorly developed plants collections grow on a mat of up to 33 μm wide sometimes elimbate throughout; limbidium mostly rhizoids mixed with diatoms and branched cyano- restricted to the vaginant lamina and basal part phytes. In its leaf shape and obscure leaves the new (up to 150 μm) of the apical lamina, on dorsal species resembles the little known Madeiran en- laminae lacking or represented by a few cells in demic F. nobreganus Luisier & P. de la Varde, but the mid part; on mid of dorsal lamina 0–16 µm in this species vegetative stem leaves are elimbate. wide, unistratose; on the vaginant lamina 0–3 Moreover, F. nobreganus is not hydrophilous. The cells wide, 16 μm wide, unistratose, marginal or new species further could be taken for a miniature intramarginal; vaginant lamina 1/2–3/5 the leaf F. leucocinctus. This species can be distinguished length, closed to slightly open, at the insertion less by its larger, typically wider (L/W 3–4), limbate wide than the stem, slightly concave, not decur- leaves, stout 3–5 stratose limbidia and by larger rent, unistratose; dorsal lamina narrow, hardly size (stems 5–30 × 2–3 mm). From the also partly rounded at the insertion, reaching the insertion, limbate F. bogosicus Müll. Hal. it differs by the not decurrent, dorsal and apical lamina unistra- obscure, typically narrower leaves. The new spe- tose with bistratose areas, alongside the costa and cies probably belongs in subgenus Fissidens, but near the insertion uni- to bistratose; costa ending subgenus Aloma cannot positively be excluded 5–7 cells below the leaf apex; mid dorsal laminal without sporophyte. cells hexagonal, 5.0–11.5 × 5–8 µm, smooth; mid speciMen exaMined for coMparison: Fissidens vaginant laminal cells 6–13 × 3–8 μm. Fertile nobreganus Luisier & P. de la Varde: Madère, sur le parts unknown. tronc de Ocotea foetens. Nord de l’Isle. 1 X 1952. Nó- eTyMology. The name refers to the obscure brega s.n. (Type: PC-PV, slide). leaves. From the Latin words obscurus (dark) and frons (frond). Fissidens bessouensis corB. haBiTaT. Near and under small waterfalls; In 2006 Alain Vanderpoorten and Eberhard Fisher riverine and lowland rainforests. collected a number of Fissidens specimens in suBsTraTe. Shady rocks; on irrigated rocks. Gabon. Amongst these was an aquatic species with fimbriate leaf margins. Having never before alTiTude. 700–1000 m. seen a Fissidens with fimbriate leaves the first disTriBuTion. Tanzania, S-Pare Mts and thought was that this represented an undescribed Nguru Mts. species close to F. bessouensis Corb. However, an examination of several specimens of this species, paraType: TANZANIA, S-Pare Mts, on the E slope including the type, showed that fimbriae were often of Mt Kwizu, WNW of Kisiwani village. Riverine forest, present. Further examination of the specimens on shady rocks near small waterfalls, 1000 m, 27 June 1972, Pócs 6716/J mixed with F. robynsianus and showed that spores of F. bessouensis are multi- F. leucocinctus (EGR, L). cellular (type specimen and in Overlaet s.n.). Apart Fig. 2. Fissidens obscurifrons Bruggeman-Nannenga, sp. nov. 1 – vegetative plant; 2–5 – leaves; 6 – leaf apex; 7 – mid leaf with elimbate dorsal lamina; 8 – mid leaf with weakly limbate dorsal lamina; 9 – detail of 8; 10 – basal part of leaf; 11 – rhizoids with thalloid protonemata – 1, 5–7 from Pócs, Nsolo & Temu 87226/AL (holotype L), 2–4, 8–11 from Pócs 6716/J (paratype: L). M. A. BRUGGEMAN-NANNENGA: TWO NEW SPECIES OF FissiDENs 121 122 POLISH BOTANICAL JOURNAL 58(1). 2013 from these two unique features, F. bessouensis is wide, to 17.5 µm wide, marginal; vaginant lamina also remarkable for its rostrate perichaetial leaves 2/5–2/3 the leaf length, basally narrower than the and subsessile thecae which made Pursell (1987) stem, narrow and straight towards the insertion, observe that F. bessouensis is truly a most remark- then frequently suddenly widened at the inser- able species of Fissidens. tion, slightly decurrent or not, unistratose, nearly or completely open (infrequently slightly open), Fissidens bessouensis Corb. Fig. 3 smaller lamella distally ± rounded; dorsal lamina tapering towards the insertion, narrow, straight Bull. Mus. Natl. Hist. Nat. 18: 109. 1912 – Type: Cen- to slightly rounded at the insertion, reaching the tral African Republic [Haut-Oubangui], Bessou, le long insertion or not, not decurrent; dorsal and apical des rives du fleuve, entre Bangui et le confluent de la lamina unistratose with scattered bistratose areas Kémo. Pendant des branches des arbres à la hauteur in mid region, juxta-costal 1–2 (–3) stratose; costa atteinte par les cruz. 2 Sept 1902. Chevalier s.n. (ho- poorly defined distally, ending 6–31 cells below loType: PC; isoType: PC-PV). Pursell (1987) cited the specimen in Potier de la Varde’s herbarium (PC-PV) as the leaf apex, bryoides-type, but frequently irreg- holotype. Since the species was described by Corbière ular, lateral cells few and large, lateral epidermis the specimen in this author’s herbarium in PC must be not to poorly differentiated, with 1 row of 2 large the holotype. cells or irregular above the vaginant laminae; mid Fissidens jordanii J. Taylor & P. de la Varde, Bull. Misc. dorsal laminal cells smooth, plane, hexagonal, Inform. Kew 1954: 505, f. A-E. 1954 – Type: Sierra thin to firm-walled, 6.5–22.5 × 6.5–14.5 µm; mid Leone, Mange, Bure. Epiphytic on small shrub (Croton vaginant laminal cells 8.0–29.0 × 6.5–14.5 μm; scarceisii Sc. Elliot) abundant on rocks now exposed basal vaginant laminal cells oblong, to 38.5 µm in river. 5. I. 1953. H. D. Jordan 851A (lecToType long, becoming linear-oblong toward the margin; designated by Bruggeman-Nannenga & Pursell 1995: gemmae frequent, manifold; hyaline to brown, PC-PV; isolecToTypes: BM). filamentous, uni-to multicellular, branched or un- Vegetative parts. stems firmly attached to branched fimbriae on marginal cells of all laminae the substrate, floating, greatly branched, 50–100 (Fig. 3: 5); epiphyllous hyaline, green or brown, × 1.5–5.0 mm (with branches), pinnate; central uni- to multicellular, uniseriate, simple or branched strand lacking, 2–6 tiers of inner and 1–8 tiers of filaments (Fig. 3: 6) as well as occasional fascicles small to large outer cortical cells; rhizoids plen- of rhizoid-like filaments originating from laminal tiful, basal and along the stem, axillary or not, cells near leaf apices (Fig. 3: 2); axillary, branched and apically on leaves, brown, smooth; axillary or unbranched, filamentous gemmae on rhizoids nodules weakly or not differentiated; leaves green, or directly from cortical cells; axillary fascicles numerous, typically distant, slightly crispate when of long unbranched rhizoids with terminal or inter- dry, linear elliptico-lanceolate, acute, 2.0–4.0 × calary areas of short cells separated by transverse 0.3–0.7 mm, L/W 5–10; margin subentire, fre- cross-walls; stems occasionally, in part, covered quently irregular by protruding cells (fimbriae ini- with long thick walled rhizoids with cross walls tials) or fimbriate on all laminae, entire in proximal partly oblique and partly transverse that produce part of vaginant lamina, elimbate or weakly lim- multicellular, filamentous, branched gemmae. bate (best seen in 5% KOH) on basal 1/4 part of Fertile parts. Perigonia and perichaetia ter- vaginant lamina; limbidium unistratose, 1–5 cells minal on short branches, perigonial and perichaetial Fig. 3. Fissidens bessouensis Corb. 1 – habit; 2 – leaf with apical cluster of gemmae and perichaetial branches; 3 – leaf; 4 – fimbriate leaf apex; 5 – marginal fimbriae; 6 – epiphyllous gemmae; 7 & 8 – perichaetial branches; 9 – sporophyte; 10 – sporo- phytic branch; 11 – peristome; 12 – multicellular spores; 13 – transverse section of stem; 14 – transverse section of leaf. 1 from Vanderpoorten 1647 (private herbarium Bruggeman-Nannenga); 2, 6, 7, 11, 12 from Chevalier s.n. (type of F. bessouensis); 3 & 8 from Jones 1539; 4, 5, 13 from Vanderpoorten 1650 (private herbarium Bruggeman-Nannenga); 9, 10, 14 from Overlaet 573 (BM). M. A. BRUGGEMAN-NANNENGA: TWO NEW SPECIES OF FissiDENs 123 124 POLISH BOTANICAL JOURNAL 58(1). 2013 branches frequently in the same leaf axil, 1–5 per & Allen 1994; Bruggeman-Nannenga 2005). Note- axil, tightly attached to the stem by rhizoids; per- worthy they also lack in a number of terrestrial igonial branches budlike, 0.2– 0.5 mm tall, anth- species, e.g., the acostate subgenus Aloma spe- eridia 140–210 µm long; perichaetial branches cies. Adaptations of F. grandifrons Brid. [subgenus short, 0.5–1.6 (3.4) mm tall, perichaetial leaves Pachyfissidens (Müll. Hal.) Kindb., section Pachy- 0.6–0.85 mm long consisting mainly of a vagi- fissidens] to a rheophilous habitat were discussed nant lamina and a long excurrent costa, apical by Iwatsuki and Suzuki (1982), and Pursell and and dorsal laminae lacking or poorly developed; Allen (1994). Apart from the absence of a central archegonia 170–260 µm long; sporophyte, seta strand such adaptations include stiff, stream-lined, short almost nonexistent, capsule symmetrical, im- imbricate, multiseriate leaves, and, astomatose mersed to emergent, 0.6–1.0 × 0.4–0.5 mm with thecae. Species of subgenus Octodiceras, are based 38–40 columns of quadratic to oblong exothecial on morphological adaptations to sites that for most cells; peristome teeth straight, undivided or deeply of the year are inundated often in sheltered places divided, short, 112–150 μm long, 31–38 µm wide (Pursell & Bruggeman-Nannenga 2004). Such ad- at base, strongly papillose; operculum rostrate, aptations similarly include the absence of a cen- 0.3 mm long; calyptra not seen (illustrated by tral strand and astomatose thecae; long, branched, Pursell, 1987); spores large, multicellular, 34–45 flaccid floating stems; linear-lanceolate, distant, × 29–33.5 µm long, slightly papillose, brown. soft, elimbate or weakly limbate leaves with short costae; and sporophytes with short setae, reduced Fissidens bessouensis is distinguished from peristomes and large spores. Adaptations observed other species in subgenus Octodiceras by its in F. bessouensis but not studied in other species ± open vaginant lamina 2/5–2/3 the leaf length of the subgenus include multicellular spores, stems with a smaller lamella that is distally more or less firmly attached to the substrate by masses of rhiz- rounded (Fig. 3: 2, 3); large multicellular spores oids; axillary perigonial, perichaetial and infertile (Fig. 3: 12); immersed to emergent sporophytes branches firmly attached by rhizoids. Furthermore (Fig. 3: 10); subsessile setae (Fig. 3: 9); small per- the formation of many structures that probably ichaetial (Fig. 3: 7, 8) and perigonial branches; and function as gemmae (for examples see description perichaetial leaves consisting of a wide vaginant and Fig. 3: 2, 5, 6). lamina plus a rostrum formed by the costa and reduced dorsal and apical laminae. Fimbriate leaf disTriBuTion. Sierra Leone, Gabon, Republic margins (Fig. 3: 5) are frequently found. The stem of the Congo. cortex typically is thin (1 tier) and composed of suBsTraTes. Attached to rocks, twigs of Cro- large thin walled cells, but can be up to 8 tiers of ton, and on roots. small thick walled cells thick. These differences are probably caused by environmental differences. haBiTaT. Rivers, cascades and rapids; sub- merged occasionally exposed. Fissidens bessouensis usually has the lax look so characteristic of subgenus Octodiceras (Brid.) speciMens exaMined. Sierra Leone, Moa River at Broth. (Pursell 1987; Pursell & Bruggeman-Nan- Jawo. Jones 1539 (BM); Mange, Bure. H. D. Jordan nenga 2004), but occasionally it can have the firm 851A (holotype of F. jordanii PC-PV, isotype of F. jor- appearance of subgenus Fissidens (e.g., the type- danii: BM). Gabon, prov. Ogooué-Ivindo, Chutes de Kongou sur l’Ivindo, 0°17′24″N, 12°34′18″E. Van- specimen of F. jordanii). These two expressions derpoorten GAB1647 (LGHF, private herbarium are not sharply distinct and, moreover, share the Bruggeman-Nannenga); rapides de Mbawaka sur le diagnostic features of F. bessouensis. fleuve de l’Ivindo, Vanderpoorten GAB1750 (LGHF, Most Fissidentaceae have stems with a central private herbarium Bruggeman-Nannenga). Democratic strand. Central strands are lacking in subgenus Republic of the Congo, riverside of Lulua [Lua], near Octodiceras and in rheophilous species of several Kalinga. Overlaet 573 (BM); riv. Lulua. Overlaet s.n. other subgenera (Iwatsuki & Suzuki 1982; Pursell (BR and PC-PV), sine locality. Overlaet s.n. (BR, cum M. A. BRUGGEMAN-NANNENGA: TWO NEW SPECIES OF FissiDENs 125 sporophytes);Wambali. Vanderijst 1004 (BR), Vanderijst BruggeMan-nannenga M. a. & pursell r. a. 1995. Notes s.n. (BR); Stanleyville, rapides des Wagenia. Leonard on Fissidens V. Lindbergia 20: 49–55. 1820 (BR). iwaTsuki, Z. & suZuki, T. 1982. A taxonomic revision of the Japanese species of Fissidens (Musci). J. Hattori Bot. illusTraTions. Pursell, 1987: fig. 23–30; Taylor Lab. 51: 329–508. & Potier de la Varde, 1954: 507, fig. A-E (as F. jor- danii). pursell R. A. 1987. A taxonomic revision of Fissidens sub- genus Octodiceras (Fissidentaceae). Mem. New York Bot. Gard. 45: 639–660. acknowledgMenTs. I am obliged to Dr. Ron Pursell for his useful comments to an earlier version of this pursell R. A. 2007. Fissidentaceae. Flora Neotrop. Monogr. paper, and to the anonymous reviewer for valuable 101: 1–278. remarks on the manuscript. pursell r. a. & allen B. H. 1994. A re-evaluation of Fis- sidens subgenus Pachyfissidens, with a detailed discussion of Fissidens grandifrons and F. geijskesii. J. Hattori Bot. references Lab. 75: 15–22. pursell r. a. & BruggeMan-nannenga M. A. 2004. A re- BruggeMan-nannenga M. A. 2005. Two new species of Fis- vision of the infrageneric taxa of Fissidens. 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