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 : 273-277

16. Cyathea costularis Bonap.

( Figs 16 ; 44J; 48F )

Notes ptéridologiques 5: 44 (1917); l.c. 9: 51 (1920); Christensen, Dansk Botanisk Arkiv 7: 25 pl. 3 figs 13-15 (1932); Tardieu in Humbert, Flore de Madagascar et des Comores, IVe famille, Cyathéacées : 16 (1951). — Alsophila rolandii R.M.Tryon, Contributions from the Gray Herbarium 200: 31 (1970). — Type: Maroantsetra, 15°26’S, 49°45’E, 300 m, VIII.1912, Perrier de la Bâthie 7980 (lecto-, P! [3 sheets: P00389606-08], here designated; isolecto-, P! [3 sheets]).

ADDITIONAL MATERIAL EXAMINED. — Madagascar. Antsiranana, Nossi-Be, forêt de Loucoubé, 13°17’30’’S, 48°15’30’’E, I.1850, Boivin s.n. (P). — Forêt d’Ambatosoratra, 14°46’S, 48°52’E, 1000 m, 8.I.1949, Cours 3413 (P). — Route d’Ambodihasina à Manandaleha, 30.XII.1950, Cours 3936 (P). — Toamasina, Sandrakatsy, forêt de Bekopila, 16°20’S, 49°37’E, 350 m, 15.XI.1959, Cours 5345 (P). — Forêt de Lokobe, 13°24’30’’S, 48°18’30’’E, 260 m, 4.XII.1989, Deroin et al. 180 (P). — Vallée de la Lokoho, Ambalavoniho, 14°34’S, 49°44’E, 75-300 m, I.1949, Humbert et al. 22967 (K). — Antsiranana, Nossi-Be, 13°17’30’’S, 48°15’30’’E, Humblot s.n. (P). — Toamasina, Maroantsetra, Ambanizana, Ambon’Andraoka, 15°37’53’’S, 49°58’33’’E, 2-471 m, 20.X.2004, Janssen et al. 2478 (MO, P,TAN), 2486 (MO, P,TAN). — Idem, 15°37’51’’S, 49°58’35’’E, 2-503 m, 21.X.2004, Janssen et al. 2489 (MO, P,TAN). — Idem, entre Ambanizana et Andranobe, 14°40’53’’S, 49°57’26’’E, 0-30 m, 25.X.2004, Janssen et al. 2518 (MO, P, TAN). — Idem, Andranobe, entre Andranobe et Bedinta, 15°40’34’’S, 49°58’03’’E, 0-628 m, 26.X.2004, Janssen et al. 2523 (MO, P, TAN). — Baie d’Antongil, 1897, Mocquerys 329 (G). — Toamasina, Nosy Mangabé, 15°29’30’’S, 49°46’E, 1897, Mocquerys 423 (G). — Antsiranana, Nossi-Be, bois Lokobe, 13°17’30’’S, 48°15’30’’E, VIII.1913, Perrier de la Bâthie 7616 (P, paralectotype). — Toamasina, Maroantsetra, Ambanizana, Andranobe, 15°41’S, 49°58’E, 100 m, 18.XII.1993, Rakotondrainibe et al.2052 (P, TAN). — Nosy Mangabe, 15°30’S, 49°46’E, 230 m, 1.XII.1993, Rakotondrainibe et al. 2054 (P, TAN), 2054 bis (P). — Toamasina, Maroantsetra, Ambanizana, Andranobe, 15°40’54’’S, 49°57’26’’E, 18.XII.1993, Rakotondrainibe 2068 (P). — Antsiranana, Andapa, RNI du Marojejy, Manantenina, 14°26’12’’S, 49°46’30’’E, 450 m, 5.X.1996, Rakotondrainibe 3294 (P, TAN). — Idem, 500 m, 6.X.1996, Rakotondrainibe 3301 (K, P, TAN). — Antalaha, Ambohitralanana, Sahafari, 15°16’18’’S, 50°20’35’’E, 430 m, 18.II.2001, Rasolohery 313 (MO, P, TAN). — Maroambihy, Sambava, 14°27’30’’S, 49°42’30’’E, 11.XII.1961, Réserves naturelles 11737RN (P). — Maroantsetra, Nosy Mangabe, 15°29’30’’S, 49°46’E, 0-330 m, 11.X.1987, Schatz 1625 (P). — Antsiranana, Nossi Bé, Loucoubé, 13°24’S, 48°19’E, I.1883, Thiebaut s.n. (K, P). — Nossibé, forêt de Loucoubé, 13°24’S, 48°19’E, 27.III.1882, Thiebaut 60 (P). — Toamasina, Maroantsetra, Ambanizana, 15°38’S, 49°58’E, 25-200 m, 25.X.1992, van der Werff et al. 12759 (P).

FIELD OBSERVATIONS. — Trunk: HT up to 6 m, DT 7-9(-12) cm, dead petioles caducous and leaf scars exposed, but sometimes a few spiny petiole bases persistent; trunk surface dark brown to black, densely spiny-tuberculate with sclerified, pyramidal, more or less antrorse scales up to 1.5 cm long, trunk surface finely muricate among these spines.

Petiole: with 1 or 2 rows of distant, light brown aerophores on either side, distributed over the entire abaxial face near the petiole base; petiole bases sometimes rather long sigmoid and appressed to the trunk.

Leaf scars: 2-2.5 × (2.5-) 4-7 cm, rounded to narrowly elliptical, with some orifices on their lower rim, sometimes with a rudiment of the petiole, spirally arranged.

Crown: erect-arched, often with a distinct infundibuliform centre.

Trunk apex: densely scaly, black, somewhat raised and well visible through the distant petioles.

Lamina: elliptical; LL 150-200 cm, WL 60-80 cm, FW 70-85(-100) cm, NP 15-20.

DESCRIPTION

Petiole: 13-28(-45) cm long, 1.8-2.2(-3) cm in diameter; stramineous to green, abaxial face violaceous brown.

Lamina: bipinnate to pinnate-pinnatisect, very coriaceous, pale green below, shiny yellowish to dark green above; lamina base shortly attenuate, basal pinnae more or less strongly reflexed and conduplicate; rachis stramineous to green.

Largest pinnae: 40-60 cm long, distant by 9-10(-14) cm, adjacent pinnae spaced to distinctly overlapping, their apex acute to shortly caudate, pinnatifid; costae and costulae of the same colour as the rachis.

Largest pinnules:5.5-8.5 × 0.7-1(-1.2) cm, spaced by less than to about their width, sessile, at most the 1 or 2 proximal pairs petiolulate, frequently broadly adnate to the costa in the upper half of the pinna; narrowly oblong, margin subentire to shallowly crenate, slightly revolute, apex acute and serrulate, base truncate to subcordate, rarely slightly auriculate; veins twice furcate.

Scales and hairs: scales present from the petiole base upwards to 40 cm on the petiole and rachis, distant to contiguous, but never very dense, persistent, narrowly triangular to deltoid, 1-1.5(-2) × 0.2-0.5 cm, abaxial scales patent, strongly indurated and transformed into more or less pyramidal spines, shiny black to dull brown, their apex antrorse and often breaking away in herbarium specimens, adaxial scales coriaceous, not appressed to the petiole, crispate, usually dull brown, scales further up on the petiole less indurated and more or less deltoid and appressed; sparse short, stiff, contorted, brown hairs mixed with caducous, dull brown, filiform scales on the adaxial face of the rachis; dense multicellular, dark brown, antrorse hairs on the adaxial face of the costae; leaf otherwise glabrous.

Sori: subcostular, contiguous to slightly spaced, 0.15-0.2 cm in diameter, covering entire pinnules; indusia globular, brown, subcoriaceous to coriaceous, at maturity dehiscing in 3 or 4 lobes, but usually not down to their base; receptacle capitate, shorter than to as long as the rim of mature indusia, paraphyses inconspicuous.

DISTRIBUTION

Northern Madagascar:Nosy Be, Marojejy, Masoala; endemic.

ECOLOGY

0-600(-1000) m. Low altitude dense evergreen rainforests and littoral forests. Also on forest margins.

REMARKS

The species is well characterised by its strongly indurated, patent to antrorse petiole scales. It is also distinct from other members of group IIc by its pinnules being frequently adnate in the upper half of the pinnae and never conspicuously petiolulate.

Juvenile plants have an obovate lamina with a cuneate base composed of gradually reduced,strongly reflexed, light green, herbaceous (not coriaceous) pinnae. Th eir pinnae are pinnatifid with narrow more or less sharply serrate segments ( Fig. 16G ). The petiole is very short and scales are not or only slightly indurated.

TYPIFICATION AND SYNONYMY

The protolog cites two syntypes. Of these, Perrier de la Bâthie 7980 comprises six sheets at P. Three sheets, carrying a leaf apex, middle pinna and petiole base respectively, have been marked “Original” by Bonaparte and are here chosen as the lectotype of the species.

17. Cyathea hebes Janssen & Rakotondr. , sp. nov. ( Figs 15 F-J; 45C; 49A)

Filix arborescens affinis Cyatheae viguieri, sed differt paleis petioli sordide brunneae, anguste triangularibus, 2.5-3 cm longis et usque ad 0.15 cm latis, non nisi in parte inferiori petioli, saepe leviter induratis et contortis expansisque.Pinnulae parum longiores, 7-8(9) cm longae sine basi auriculata . Indusia brunnea, membranacea vel subcoriacea.

TYPUS. — Madagascar, Toamasina, Maroantsetra, presqu’île de Masoala, Ambanizana , Ambon’Andraoka, piste menant du village d’Ambanizana à la crête E d’Ambanizana 15°37’53’’S, 49°58’33’’E, 2-471 m, 20.X.2004, Janssen et al. 2480 (holo-, P! [4 sheets: P00589575-78]; iso-, P! [3 sheets], TAN!; one trunk surface mould at P!) .

ADDITIONAL MATERIAL EXAMINED. — Madagascar. Toamasina, Maroantsetra, Ambanizana, 15°37’49’’S, 49°58’37’’E, 220 m, 16.IV.2002, Antilahimena et al. 1008 (P). — Toamasina, Maroantsetra, Ambanizana, Ambon’Andraoka, 15°37’53’’S, 49°58’33’’E, 2-471 m, 20.X.2004, Janssen et al. 2479 (MO, P, TAN). — Idem, 15°37’51’’S, 49°58’35’’E, 2-503 m, 21.X.2004, Janssen et al. 2487 (MO, P, TAN). — Idem, entre Ambanizana et Andranobe, 15°40’53’’S, 49°57’26’’E, 0-30 m, 25.X.2004, Janssen et al. 2519 (MO, P,TAN). — Masoala Peninsula, Andranobe, 15°41’S, 49°58’E, 10-110 m, 25.II.1999, McPherson et al. 17689 (P). — Idem, 15°41’S, 49°58’E, 70 m, 19.XII.1993, Rakotondrainibe et al. 2053 (MO, P). — Antsiranana, Andapa, RS d’Anjanaharibe-Sud, 14°45’18’’S, 49°30’18’’E, 800 m, 20.X.1994, Rakotondrainibe et al. 2117 (P). — Toamasina, Masoala Peninsula, near Ambizana, 15°39’S, 49°58’E, 10 m, 30.X.1992, van der Werff et al. 12798 (MO, P).

FIELD OBSERVATIONS. — Trunk: HT up to 6 m, DT 5.5-7 cm, dead petioles caducous and the leaf scars exposed, rudiments of the base may persist in its apical part; trunk surface brown to black, muricate.

Petiole: with 1 or 2 rows of brown aerophores on either side; petiole bases sigmoid, sometimes forming a fascicle above the trunk apex before being gradually recurved.

Leaf scars: 2.5-3 × 5.5-7 cm, elliptic, slightly raised, several orifices below the scars; spirally arranged.

Trunk apex: densely scaly, dull brown; concealed among the more or less close standing petioles.

Crown: horizontal to umbrella-shaped, in young plants more or less funnel-shaped with straight petioles.

Lamina: elliptic; LL 100-175 cm, WL 80-110 cm, FW 50-100 cm, NP 8-14.

DESCRIPTION

Petiole: (20-) 40-55 cm long, 1.5-2 cm in diameter; stramineous green, abaxial face more or less violaceous brown, light brown when dry.

Lamina: bipinnate, very coriaceous, pale green below, shiny green to dark green above; lamina base shortly attenuate to truncate, basal pinnae conduplicate, patent to reflexed; rachis of the same colour as the petiole.

Largest pinnae: 40-50 cm long, distant by (9-) 10- 12 cm, adjacent pinnae overlapping, their apex acute, pinnatifid; costae and costulae of the same colour as the petiole.

Largest pinnules: 7-8(-9) × 1-1.3 cm, spaced by less than to about their width, petiolulate, triangularoblong, their base rounded, truncate or cuneate, but never auriculate, their margin subentire and slightly revolute, sharply serrate in the shortly caudate apex of the pinnules; veins 2 to 3 times furcate, 12-16 veins per cm.

Scales and hairs: scales present from the petiole base upwards to 30 cm on the petiole, never reaching the first pinna pair, dense and overlapping, persistent, narrowly triangular to filiform, 2.5-3 × 0.05-0.15 cm, straight and twisted to very much contorted, not appressed to the petiole, dull brown to dull dark brown, shiny in young specimens only, not or slightly indurated at their point of insertion; sparse, contorted, dark brown, antrorse multicellular hairs on the adaxial face of the rachis and costae; leaf otherwise glabrous.

Sori: subcostular, spaced by less than their width, sometimes by more than their width in young specimens, 0.15-0.2 cm in diameter, covering the entire pinnule; indusia globular, light brown, membranous to subcoriaceous, at maturity dehiscing in 2-4 lobes; receptacle capitate to disciform, as long as to shorter than the rim of mature indusia, paraphyses inconspicuous.

DISTRIBUTION

Northern Madagascar: Masoala peninsula and baie d’Antongil; endemic.

ECOLOGY

0-400 m. Dense evergreen rainforests.

REMARKS

Cyathea hebes is easily distinguished from C. costularis by its muricate, not spiny, trunk surface and its much denser and much less indurated petiole scales. It differs from C. viguieri Tardieu by its narrow and dull brown petiole scales ascending only to about halfway on the petiole as well as by its light brown, membranous to subcoriaceous indusia.

Rakotondrainibe 2117 and Janssen et al. 2479 have shiny petiole scales, but the specimens are taken from young plants.

ETYMOLOGY

The epithet hebes refers to the dull petiole scales in adult plants of this species as opposed to shiny scales in the closely related C. viguieri .

Figures

FIG. 16. — Cyathea costularis Bonap.: A, pinnules abaxially with a fragment of the costa, sori omitted from one pinnule; B, strongly indurated scale from the abaxial face of the base of the petiole; C, habit; D, scale from the lateral face of the upper half of the petiole; E, basal part of the leaf (from the petiole base up to the first pinna pair),dorsal view, lateral view in upper half,one pinna pruned;F, leaf scar and trunk surface; G, basal part of the leaf of a juvenile plant, ventral view, note the strongly reflexed decrescent pinnules and soft scales. A, B, D, E, Rakotondrainibe 3301 (P); C, uncollected, photograph at P; F, Janssen et al. 2489 (P); G, Janssen et al. 2486 (P). Scale bars: A, F, G, 1 cm; B, D, 0.5 cm; C, 1 m; E, 5 cm.

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. 15. — A-E, Cyathea conferta Janssen & Rakotondr.; A, pinnules adaxially with a fragment of the costa, note that the auricles overlap the costa; B, sori, note that these are small and very close standing; C, scale from the base of the petiole; D, petiole, note that the filiform scales are restricted to its base; E, pinna abaxially with a fragment of the rachis, sori and veins only partly indicated, note the close standing pinnules; F-J, C. hebes Janssen & Rakotondr.;F, sori, note that these are big and spaced from each other; G, pinnules abaxially with a fragment of the costa; H, basal part of the leaf (from the petiole base up to the first pinna pair), lateral view; I, scale from the base of the petiole; J, pinna abaxially with a fragment of the rachis, sori only partly indicated. A-E, van der Werff et al. 12850 (P); F-J, Janssen et al. 2480 (P). Scale bars: A, C, G, 1 cm; B, F, 0.1 cm; D, E, H, J, 5 cm; I, 0.5 cm. D, E, Drawings by B. Raufeisen.