Janssen, Janssen & Rakotondrainibe, 2008

Janssen, Thomas & Rakotondrainibe, France, 2008, A revision of the indusiate scaly tree ferns (Cyatheaceae, Cyathea subgen. Alsophila sect. Alsophila) in Madagascar, the Comoros and the Seychelles, Adansonia (3) 30 (2), pp. 221-376 : 257-260

9. Cyathea ligulata Baker ( Figs 9 ; 45F ; 48B )

Journal of Botany 22: 140 (1884); Christensen, Dansk Botanisk Arkiv 7: 26, pl. 5 figs 11-13; Tardieu in Humbert, Flore de Madagascar et des Comores, IVe famille, Cyathéacées : 14. — Alsophila ligulata (Baker) R.M.Tryon, Contributions from the Gray Herbarium 200: 30 (1970). — Type: Madagascar, Humblot 299 (holo-, K! [K000009938]; iso-, MO!, P! [3 sheets], TAN!, W!). — Madagascar, Prov. d’Anstiranana, Andapa, RNI 12 du Marojejy, au bord d’un affluent de la rivière Manantenina, à 10 km au NW du village de Manantenina, 14°26’S, 49°45’42’’E, 750-800 m, 22.X.1996, Rakotondrainibe 3443 (epi-, P! [3 sheets: P00084923-25], here designated).

ADDITIONAL MATERIAL EXAMINED. — Madagascar. Toamasina, Makira , 15°26’S, 49°23’E, 2004, Antilahimena et al. 2543 ( MO, TAN) . — Anjanaharibe , 15°11’S, 49°38’E, 800 m, 27.XII.1950, Cours 3892 (P). — Massif de l’Anjanaharibe , pentes et sommet Nord , 14°36’S, 49°27’E, 900 m, 1951, Humbert et al. 24583 (G, P). — Toamasina, Maroantsetra , piste menant au sommet de l’Ambohitsitondroinan’ Ambanizana , 15°31’08’’S, 50°00’16’’E, 620 m, 22.X.2004, Janssen et al. 2494 ( MO, P, TAN). — Idem , 15°34’S, 50°00’E, 680 m, 9.XII.1993, Rakotondrainibe et al. 2057 (P). — Antsiranana, Anjanaharibe-Sud, Befingotra , 14°42’30’’S, 49°27’30’’E, 24.X.1994, Rakotondrainibe 2196 (P). — Idem, 14°45’18’’S, 49°30’18’’E, 810 m, 27.X.1994, Rakotondrainibe et al. 2213 (K, MO, P, TAN). — RNI du Marojejy, Manantenina , 14°26’12’’S, 49°46’30’’E, 450m, 5.X.1996, Rakotondrainibe 3272 ( MO, P, TAN). — Idem , 14°26’S, 49°45’42’’E, 780 m, 15.X.1996, Rakotondrainibe 3377 (K, P, TAN) . — Forêt de Betaolana, Ambodiangezoka, 14°32’18’’S, 49°26’18’’E, 870-880 m, 8.X.1999, Rakotondrainibe et al. 4811 (P, TAN) , 4813 (P). — Idem , 800 m, 12.X.1999, Rakotondrainibe et al. 4910 (P) . — Antsiranana, massif d’Anjanaharibe-Sud , forêt d’Analabe, SW de Befingotra , 14°46’S, 49°26’30’E, 1120 m, 29.X.1999, Rakotondrainibe et al. 5110 (P). — PN de Marojejy, Doany, 14°25’36’’S, 49°36’30’’E, 800 m, 19.X.2001, Rakotondrainibe et al. 6301 (K, MO, P, TAN) .

FIELD OBSERVATIONS. — Trunk: HT up to 4 m, DT 5-8 cm, dead petioles caducous and leaf scars exposed, sometimes a rudiment persistent on the scars; trunk surface tuberculate with patent, black, pyramidal, spiny scales, up to 0.5 cm wide at their base, mixed with many smaller squaminate spines, trunk surface brown to black and rather smooth between the spines.

Petiole: with 1 row of brown, much spaced aerophores on either side; petiole bases more or less straight.

Leaf scars: about 2 × 3 cm, elliptical, spirally arranged.

Crown: more or less funnel-shaped, petiole and base of rachis straight, upper half of rachis arched.

Trunk apex: densely spiny-scaly, scales patent, black, apex raised and well visible through the distant petiole bases.

Lamina: elliptical to ovate; LL 120-200 cm, WL 70 cm, FW 79 cm, NP 18-30.

DESCRIPTION

Petiole: 15-80 cm long, about 2 cm in diameter; castaneous, rarely greenish, covered with a caducous indument of minute brown squamules.

Lamina: bipinnate, herbaceous to subcoriaceous (never coriaceous), pale green below, dull dark green above, basal pinnae usually strongly reflexed; rachis of the same colour as the petiole.

Largest pinnae: 30-35 cm long, distant by about 6 cm; broadest in or above their middle; adjacent pinnae slightly spaced to contiguous, their apex shortly caudate, pinnatifid; costae and costulae of the same colour as the petiole.

Largest pinnules: 2.3-4(-4.5) × 0.5-0.7(-0.8) cm, spaced by less than to almost their width, sessile, linear-oblong, conspicuously biauriculate, adjacent auricles overlapping, margin crenate to subentire, apex rounded to obtuse, rarely acute, distal pinnules usually with a sharper apex than proximal pinnules; veins once to thrice furcate.

Scales and hairs: scales present from the petiole base upwards to 30(-50) cm on the petiole, spaced, i.e. not overlapping, persistent, conical to pyramidal, 0.5-0.8 × 0.2-0.4 cm, straight, patent to very slightly antrorse, shiny dark brown to black, strongly indurated and spiny; dense triangular to deltoid, brown scales, up to 0.2 cm long, on the abaxial face of the costae and costulae, mixed with whitish acaroid to amorphous squamules; brown, antrorse, contorted multicellular hairs on the adaxial face of the costae and in the apical part of the costae also on the abaxial face; scattered patent, hyaline hairs present on the abaxial face of the costulae; sparse and short contorted hairs and scattered filiform brown scales on the adaxial face of the rachis.

Sori:median to subcostular, contiguous to slightly distant, about 0.1 cm in diameter, covering the lower three quarters of the pinnules or entire pinnules; indusia globular, light brown, membranous, at maturity dehiscing in irregular lobes or persistent as a cup with a lacerate rim; receptacle capitate, about as long as the rim of mature indusia, with inconspicuous filiform paraphyses shorter than the sporangia.

DISTRIBUTION

Northern Madagascar:Marojejy and baie d’Antongil; endemic.

ECOLOGY

(450-) 700-1100 m. Dense evergreen rainforests, frequently in wet places next to small streams.

REMARKS

We here publish the first complete description of this taxon including characters of the petiole base. Cyathea tsaratananensis C.Chr. has similar, but usually smaller, petiole scales and is easily distinguished from the present taxon by its larger pinnules and glabrous adaxial face of costae and costulae.

TYPIFICATION AND SYNONYMY

The original material consists of single pinnae and pairs of pinnae with rachis fragments and lacks petiole scales. We hence designate an epitype to unambiguously define this taxon.

10. Cyathea lisiae Janssen & Rakotondr. , sp. nov. ( Figs 10 ; 45H )

Filix arborescens Cyatheae decrescenti affinis sed differt paleis petioli distantibus, breviter deltoideis, 0.2-0.3 cm longis et 0.1-0.15 cm latis, nitide atris, adpressis vel parum arcuatis, ascendentibus ad primum par pinnarum, transientibus in paleis bicoloribus in parte superiore petiolo (palei bicolores basi straminei membranacei, apice brunnei vel atri et coriacei). Pagina abaxalis rhachidis, costarum et costularum dense obtecta pilis multicellularibus hyalinis interdum apice in palea anguste lanceolata transientibus. Pinnulae 1.5-1.7 cm longae et 0.3 cm latae, costae adnatae, margine integra, apice acuta vel obtusa et crenulata.

TYPUS. — Madagascar, Province de Fianarantsoa, PN Ranomafana, Marotreho à 6 km au S du village de Ranomafana, 21°18’14’’S, 47°27’42’’E, 900 m, 10.XII.2000, Rabarimanarivo 204 (holo-, P! [P00477696]) .

FIELD OBSERVATIONS. — Trunk: HT 4 m, DT 5 cm, dead petioles caducous and leaf scars exposed; surface dark brown, smooth to finely muricate, covered with distant, deltoid, appressed, black scales.

Leaf scars: with 4 or 5 distinct orifices on their lower rim.

Lamina: LL 100 cm, NP 30.

DESCRIPTION

Petiole: 45 cm long, about 1 cm in diameter; light reddish brown to stramineous; aphlebia absent.

Lamina: pinnate-pinnatisect, subcoriaceous, dull pale green below, dark green above (when dry), lamina base acute to truncate, i.e. basal pinnae slowly and gradually reduced in size and not attaining the petiole base; rachis of the same colour as the petiole.

Largest pinnae: 23 cm long, distant by 3-3.5 cm, adjacent pinnae contiguous, their apex caudate, pinnatifid; costae and costulae of the same colour as the petiole.

Largest pinnules:1.5-1.7 × 0.3 cm, spaced by much less than their width, broadly adnate to the costa, oblong, slightly falciform, margin entire, the acute to obtuse apex crenulate; veins once furcate.

Scales and hairs: scales present from the petiole base upwards to about 45 cm on the petiole, reaching the first pinna pair, distant, persistent, deltoid, 0.2-0.3 × 0.1-0.15 cm, shiny black with a narrow brown margin, appressed to slightly arching, very coriaceous, becoming gradually reduced in size upwards on the petiole, where they become bicolourous with a patent to antrorse, stramineous, membranous base and a blackish brown, coriaceous tip; dense, patent, multicellular, hyaline hairs and scale-topped hairs, up to 3 mm long, present on the abaxial face of the lamina axes and veins, these hairs on the rachis and, to a lesser extent, on the costae with an indurated, dark basal cell; dense, more or less straight, patent to antrose, sometimes appressed, multicellular, brown hairs cover the adaxial face of the rachis and costae.

Sori: subcostular, distant, about 0.1 cm in diameter, covering the pinnules for three quarters of their length; mature indusia membranous, light brown, a relatively shallow cup or appressed collar around the base of the receptacle; receptacle capitate, longer than the rim of mature indusia, paraphyses inconspicuous.

DISTRIBUTION

Southern Central Madagascar: Ranomafana region; endemic.

ECOLOGY

900 m. Evergreen rainforest, next to a small stream.

REMARK

The species is clearly distinct from all taxa of group IIb by its distant short deltoid petiole scales grading into bicolourous scales further up on the petiole. It is, with C. auriculata , the only Madagascan taxon with scale-topped hairs on the lamina axes.

ETYMOLOGY

This species is dedicated to Ms Hery Lisy Ranarijaona, botanist at the University of Mahajanga having extensively worked in the Ranomafana area, in acknowledgement of her substantial contributions to fieldwork associated with the present revision.

Figures

FIG. 9. — Cyathea ligulata Baker: A, pinnae adaxially with a fragment of the rachis, note that the auricles of the pinnules overlap the rachis; B, two strongly indurated pyramidal scales from the base of the petiole (left) and one narrowly triangular scale from the abaxial face of the costa (right); C, habit; D, basal part of a pinnule abaxially (at the junction of the costula with the costa) showing indusia as well as the densely scaly costa and costula; E, basal part of the leaf (from the petiole base up to the second pinna pair), dorsal view, basalmost pinnae are strongly reflexed; F, pinnules abaxially with a fragment of the costa, form with long pinnules with a crenate margin and an obtuse apex; G, pinnules abaxially with a fragment of the costa, form with short pinnules with a subentire margin and a rounded apex; H, leaf scars and spiny trunk surface. A, D, G, Rakotondrainibe et al. 5110 (P); B, E, Rakotondrainibe et al. 2057 (P); C, H, Janssen et al. 2494 (P); F, Rakotondrainibe et al. 2057 (P). Scale bars: A, E, 5 cm; B, F, G, 0.5 cm; C, 1 m; D, 0.1 cm; H, 1 cm.

FIG. 45. — Distribution maps for Cyatheaceae (genus Cyathea) of Madagascar and the Comoros. Species are arranged in alphabetical order.

FIG. 48.— Photographic documentation of field characters in Cyathea: A, B, D-F, habit;A’, B’, D’-F’, trunk apex;A”, B”, D”-F”, trunk surface at breast height.A-A”, Cyathea meridionalis Janssen & Rakotondr.; B, B’, C. ligulata Baker; C, trunk apex of C. acutula Tardieu var.acutula; D-D”, C. acutula Tardieu var.deltoidea Janssen & Rakotondr.; E-E”, C. appendiculata Baker; F-F”, C. costularis Bonap. A, A”, E, F, uncollected; A’, Janssen et al. 2599; B, B’, Janssen et al. 2494; C, Janssen et al. 2401; D-D”, Janssen et al. 2788; E’, E”, Janssen et al. 2764; F’, Janssen et al. 2489; F”, Janssen et al. 2518. Vouchers at P. Photos B, C, F, A’, B’, F’, F” by G. Rouhan (MNHN); all other photos by T. Janssen. Scale bars: A, B, D-F, 1 m; C, A’, B’, D’-F’, A”, D”-F”, 10 cm.

FIG. 10. — Cyathea lisiae Janssen & Rakotondr.: A, pinnae abaxially with a fragment of the rachis, sori only partly indicated; B, pinnules abaxially with a fragment of the costa; C, proximal part of a pinnule abaxially with a dense indument of multicellular hairs and scale-topped hairs, note the rudimentary collar-like indusium; D, leaf scar; E, basal part of the leaf (from the petiole base up to the first pinna pair), lateral view; F, bicolorous scale from the upper third of the petiole; G, concolorous scale from the base of the petiole. A-G, Rabarimanarivo 204 (P). Scale bars: A, E, 5 cm; B, 0.5 cm; C, F, G, 0.1 cm; D, 1 cm.