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 : 342-348

38. Cyathea lastii Baker

( Figs 38 ; 45E ; 52F )

Journal of Botany 29: 3 (1891); Christensen, Dansk Botanisk Arkiv 7: 32, pl. 6 figs 16-18 (1932); Tardieu in Humbert, Flore de Madagsacar et des Comores, IVe famille, Cyathéacées : 26 (1951). — Alsophila lastii (Baker) R.M.Tryon, Contributions from the Gray Herbarium 200: 30 (1970). — Type: North West Madagascar, Bé Kilus Mountains, XII.1890, Last s.n. (lecto-, K! [2 sheets: K000009940, -41], here designated; isolecto-, K! [4 sheets]).

ADDITIONAL MATERIAL EXAMINED. — Madagascar. Maromandia, Ankaramy, 13°59’S, 48°10’30’’E, 11.VI.1923, Decary 2167 (P). — Antsiranana, Manongarivo, Bekolosy, 14°02’S, 48°19’E, 1280 m, 30.III.1996, Gautier et al. 2929 (G). — Idem, 800 m, 10.IX.1997, Gautier et al. 3261 (G, P). — Idem, crête entre les deux bras de l’Ambahatra, 14°00’S, 48°23’30’’E, 850 m, 5.III.1999, Gautier et al. 3474 (G, P). — Idem, 13°59’51’’S, 48°25’43’’E, 1150 m, 24.IX.2004, Janssen et al. 2381 (P, TAN). — Idem, 14°01’22’’S, 48°25’03’’E, 1614 m, 26.IX.2004, Janssen et al.2390 (MO, P, TAN). — Idem, 13°59’51’’S, 48°25’43’’E, 1200 m, 1.X.2004, Janssen et al. 2430 (P, TAN). — Idem, 1147 m, 1.X.2004, Janssen et al. 2431 (MO, P, TAN). — Antsiranana, Manongarivo, E of Ankaramy, Antsatrotro, 14°05’S, 48°23’E, 600-900 m, III.1993, Malcomber et al. 2290 (G, MO). — Idem, 14°05’S, 48°24’E, 1000-1400 m, 11.X.1991, Rakotondrainibe 1385 (P). — Manongarivo, Bekolosy, 14°02’S, 48°19’E, 550 m, 24.I.1992, Rakotondrainibe 1415 (P). — Idem, Mt. d’Antsatrotro, 14°05’S, 48°24’E, 1200 m, 20.V.1992, Rakotondrainibe 1712 (K, MO, P). — Antsiranana, Manongarivo massif, above village of Ambodisakoana, 14°05’S, 48°20’E, 500-1000 m, 20.X.1994, van der Werff et al. 13538 (MO, P). — Without locality, 1860, Barbey s.n. (G).

FIELD OBSERVATIONS. — Trunk: HT up to 4.5(-8) m, DT 11-13 cm excluding, 15-20 cm including the persistent petiole bases that cover the entire trunk or its upper part only, the leaf scars usually visible below; trunk thickened at its base by a mantle of adventitious roots; trunk surface dark brown, muricate.

Petiole: with a row of white to brown aerophores on either side; petiole base more or less sigmoid.

Leaf scars: 2-4 × 3-9 cm, elliptic to ovate, with orifices below the scars; spirally arranged.

Crown: large and umbrella-shaped with arching petioles and rachises.

Trunk apex: densely covered with brown scales, concealed among the more or less spaced petioles; some dead leaves persistent and hanging from the apex.

Lamina: elliptic to ovate; LL 250-300 cm, WL 120- 180 cm, FW (75-)120-145(-195) cm, NP 11-20.

DESCRIPTION

Petiole: 50-80(-130) cm long, 2.5-3 cm in diameter; green, abaxial face violaceous, distantly and sharply muricate; glabrous, covered with a bluish to glaucous waxy layer, that can be easily wiped off, invisible in herbarium specimens; reduced pinnae absent from the lower half of the petiole.

Lamina:bipinnate-pinnatisect to tripinnate, herbaceous to subcoriaceous (not coriaceous), fertile-sterile dimorphism absent; shiny light to dark green above, pale light green below, lamina base shortly attenuate to truncate, basal pinnae patent to slightly reflexed, more or less conduplicate; rachis of the same colour as the petiole, completely green distally.

Largest pinnae: 60-75 cm long, distant by 16- 20 cm, often distinctly petiolulate with the petiolule 1-4 cm long, adjacent pinnae overlapping; costae and costulae of the same colour as the rachis, the costulae usually darker than the costae when dry.

Largest pinnules: (10.5-)12-14 × (1.5-) 2.5-3 cm, the 2 or 3 proximal pinnule pairs distinctly petiolulate, petiolule up to 1 cm long, adjacent pinnules spaced by less than their width, linear-oblong, their apex conspicuously caudate, divided to the costula into broadly adnate segments, the 1-3 proximal segment pairs distinctly petiolulate to sessile and deeply crenate, segments proximally decurrent, the bases of adjacent segments confluent from about the middle of the pinnule; pinnule segments 0.3-0.5 cm wide, spaced by less than to about their width, more or less falciform, the margin of proximal segments crenate, becoming serrate in distal segments, flat, the apex of proximal segments obtuse, becoming acute in distal segments, rarely rounded; lateral veins in the segments once to rarely twice furcate.

Scales and hairs: scales present from the petiole base upwards to about 10 cm on the petiole, at its base moderately dense and overlapping, rapidly caducous further up, narrowly triangular, 3-4.5 × 0.2-0.4 cm, straight, with a twisted and slightly crispate apex, shiny brown, concolourous or with a very narrow, light brown, erose margin, not appressed to the petiole, not indurated; costulae and upper part of the rachis adaxially with sparse contorted, light brown, multicellular hairs, the lower part of the costa and the rachis glabrous adaxially; leaf otherwise glabrous.

Sori: in a median position between the midvein and the segment margin, distinctly spaced from each other by less than to about their width, about 0.1 cm in diameter, covering the entire segment except its apex; indusia cup-shaped, even when young, subcoriaceous to coriaceous, light brown to stramineous, at maturity persistent as a cup with an entire rim, very rarely with 1 or 2 slits; receptacle elongate, somewhat flattened, slightly to conspicuously longer than the rim of mature indusia, its apex with inconspicuous short, filiform paraphyses.

DISTRIBUTION

Northern Madagascar: Manongarivo massif; endemic.

ECOLOGY

550-1600 m. Dense evergreen rainforests and crest forests.

REMARKS

A very characteristic taxon, which can easily be distinguished from other species of the Western Indian Ocean region by its big, petiolulate pinnae and big, acute pinnule segments as well as by its always cupshaped indusia and its sori being spaced from each other and from the midvein of the segment. In the forest, it is easily recognized by its big leaves and naked green petioles covered with a bluish waxy layer ( Fig. 52 F’). Rakotondrainibe 1415 has 0-3 pairs of reduced pinnae inserted in the lower half and especially near the base of the petiole.

TYPIFICATION AND SYNONYMY

Five sheets of Last s.n. (K!) have been traced comprising two detached fertile middle pinnae, each halved and glued on two separate sheets as well as a smaller fertile pinna with a rachis fragment probably taken

A

B

D

near the apex of the leaf. All sheets are equivalent in carrying the name and collection locality in Baker’s handwriting. The sheets that need to be associated to constitute an entire pinna are obvious from the arrangement of the pinnules and we here designate an entire fertile pinna as a two-sheet lectotype of C. lastii Baker (K000009940, -41). Epitypification is superfluous as all important differential characters are available in the fragmentary original material.

39. Cyathea melleri (Baker) Domin ( Figs 1E ; 39 ; 45L ; 53A )

Pteridophyta: 264 (1929); Domin, Acta Botanica Bohemica 9: 135 (1930); Christensen in Perrier, Catalogue des Plantes de Madagascar, Ptéridophytes: 21 (1931); Christensen, Dansk Botanisk Arkiv 7: 36, pl. 7 figs 11- 17 (1932). — Hemitelia melleri Baker , Synopsis Filicum: 456 (1874). — Gymnosphaera melleri (Baker) Tardieu in Humbert, Flore de Madagascar et des Comores, IVe famille, Cyathéacées : 38 (1951). — Alsophila melleri (Baker) R.M.Tryon, Contributions from the Gray Herbarium 200: 31 (1970). — Type: Madagascar, from the sea to Antinanarivo (sic), VIII-IX.1862, Meller s.n. (lecto-, K! [K000009937], here designated; isolecto-, K!). — Madagascar, Toamasina, Andasibe, RS d’Analamazaotra, autour du Lac Vert, 18°56’S, 48°26’E, 930-950 m, 11.XI.2004, Janssen et al. 2569 (epi-, P! [7 sheets: P00589623-29], here designated; isoepi-, P! [6 sheets], MO!, TAN!; one trunk surface mould at P!).

Hemitelia glandulosa Kuhn ex Baker, Annals of Botany 5: 188 (1891). — Cyathea glandulosa (Kuhn ex Baker) Domin , Pteridophyta: 264 (1929); Domin, Acta Botanica Bohemica 9: 119 (1930). — Type: Madagascar, Imerina , Andrangoloaka, XI.1880, Hildebrandt 4176 (lecto-, B! [B200129546], here designated; isolecto-, B! [4 sheets], BM!, G!, K!, P! [4 sheets]).

Cyathea melleri (Baker) Domin var. virescens C.Chr., Dansk Botanisk Arkiv 7: 36, pl. 6 figs 24-27 (1932). — Cyathea virescens C.Chr. in Perrier, Catalogue des plantes de Madagascar, Ptéridophytes: 22 (1931), nom. nud. — Gymnosphaera melleri var. virescens (C.Chr.) Tardieu in Humbert, Flore de Madagascar et des Comores, IVe famille, Cyathéacées : 38 (1951). — Type: Madagascar, forêt orientale, Mt. Andriantantely au nord d’Anivorano, VI.1922, Perrier de la Bâthie 14746 (lecto-, P! [3 sheets: P00418729-31], here designated; isolecto-, P! [4 sheets]).

ADDITIONAL MATERIAL EXAMINED. — Madagascar. Ambodiriana, 14.XII.1944, Cours 1916 (P). — Fianarantsoa,

Andringitra, forêt d’Ambodipaiso, 12.I.1945, Cours 2261 (P). — De Nonokambo à Varaina, 17°45’S, 48°45’E, 1200 m, 17.I.1945, Cours G 2382 (P). — Bemainty à Andranomanitra, 800-850 m, 9.III.1951, Cours 4227 (P). — Brickaville, Ambalarondra, 300 m, 21.IV.1951, Cours 4506 (P). — Analamazaotra, 18°57’20’’S, 48°24’30’’E, XII.1905, D’Alleizette 8 (P). — Antananarivo , 18°55’S, 44°31’E, Gilpin s.n. (K). — Manjarivolo , Andrianomy, 22°17’S, 46°52’E , 1300 m, 4.XI.1970, Guillaumet 3509 (P). — Périnet, Analamazaotra, 18°56’S, 48°26’E, 1.X.1971, Guillaumet 3868 (P). — Helsenberg & Bojer s.n. (BM). — Toamasina , Zahamena, massif de l’Andrangovalo, 17°40’S, 48°45’E, 1200 m, X.1937, Humbert et al. s.n. (P). — Bassin de l’Itomampy, Mt. Papanga près de Befotaka, 23°51’S, 46°57’E , 1000-1600 m, XII.1928, Humbert 6894 (P). — Vallée de la Lokoho, près d’Ambalavoniho, 14°34’S, 49°44’E , 75-300 m, I.1949, Humbert et al. 22822 (P). — Idem, mont Ambatosoratra, 14°32’S, 49°42’E , 1000 m, I.1949, Humbert et al. 22845 (BR, K, P). — Toamasina , RNI Betampona, Rendriendry, 17°55’48’’S, 49°12’E, 310-580 m, 6.XI.2004, Janssen et al. 2536 ( MO, P, TAN). — Fianarantsoa , corridor forestier reliant la RS d’Ivohibe au PN Andringitra, forêt d’Angodongodona, 22°25’07’’S, 46°55’31’’E, 1100 m, 20.IV.2005, Janssen et al. 2789 ( MO, P, TAN). — Toamasina , Analamazaotra, 18°56’S, 48°26’E, Jardin Botanique de Tananarive 3231 (P). — Idem, Didy , forêt de Tsiazomborona, 20.XI.2005, Labat et al. 3574 (P). — Idem, RN de Betampona, 17°55’S, 49°13’E , 450 m, 19.XII.1938, Lam & Meeuse 5989 (K, P). — Antsiranana , RN de Marojejy, trail to the summit of Marojejy Est , 14°26’S, 49°46’E, 850- 1000 m, 11.II.1989, Miller et al. 3981 (P). — Toamasina , Andasibe, Analamazaotra, 18°56’S, 48°26’E, 800 m, VII.1914, Perrier de la Bâthie 7579 (P). — Idem, II.1912, Perrier de la Bâthie 11539 (P). — Forêt d’Ambatovy, 11 km NE Moramanga, 18°49’S, 48°18’E, 1997, Rakotomalaza 1599 (P). — Toamasina , Sandrakatzy, forêt de Verezanantsoro, SE de Varary , 16°26’S, 49°38’E, 500 m, 6.I.1994, Rakotondrainibe et al. 2050 ( MO, P, TAN). — Antsiranana , RS d’Anjanaharibe-Sud, SSW de Befingotra , 14°45’18’’S, 49°30’18’’E , 790 m, 20.X.1994, Rakotondrainibe et al. 2115 (P). — Idem, 14°44’42’’S, 49°27’42’’E, 1280 m, 4.XI.1994, Rakotondrainibe et al. 2340 ( MO, P, TAN). — Fianarantsoa , RNI d’Andringitra, piste menant d’Ambalamenjana à Ambatomboay, 22°13’S, 47°12’E, 720 m, 14.V.1995, Rakotondrainibe 2556 (P, TAN). — Idem , berges d’un affluent de la rivière Sahavatoy, 22°13’22’’S, 46°58’18’’E , 1280 m, V.1995, Rakotondrainibe 2714 (P, TAN). — Toliara , Tolanaro, RNI d’Andohahela, NW d’Eminiminy, 24°35’40’’S, 46°44’30’’E, 800 m, 30.X.1995, Rakotondrainibe 3003 ( MO, P, TAN). — Idem , 1100 m, 8.XI.1995, Rakotondrainibe 3050 ( MO, P, TAN). — Antsiranana , RNI du Marojejy, NW de Manantenina , 14°26’12’’S, 49°46’30’’E , 480 m, 5.X.1996, Rakotondrainibe 3271 ( MO, P). — Idem , 450 m, 5.X.1996, Rakotondrainibe 3293 (P, TAN). — Idem , 490 m, 9.X.1996, Rakotondrainibe 3335 (P). — Antananarivo , forêt d’Andranomay, SE d’Anjozorobe, 18°28’48’’S, 47°57’18’’E, 1300-1450 m, 15.XII.1996, Rakotondrainibe 3718 (P). — Fianarantsoa , RS d’Ivohibe, ENE d’Ivohibe, 22°28’12’’S, 46°57’36’’E, 850-950 m, 7.X.1997, Rakotondrainibe et al. 4055 (P, TAN). — Antananarivo , forêt d’Ankilahila, SE de Tsinjoarivo , 19°42’24’’S, 47°51’E, 1400-1560 m, 15.I.1999, Rakotondrainibe 4743 (P, TAN). — Antsiranana , forêt de Betaolana, NW d’Ambodiangezoka, 14°32’18’’S, 49°26’18’’E, 900 m, 14.X.1999, Rakotondrainibe et al. 4911 (P, TAN). — Fianarantsoa , PN de Ranomafana, forêt de Vatoharanana, 21°17’24’’S, 47°26’E, 1000-1100 m, 3.X.2000, Rakotondrainibe et al. 5866 (K, P, TAN). — Antsiranana , PN de Marojejy, SE de Doany , 14°25’36’’S, 49°36’30’’E , 820 m, 14.X.2001, Rakotondrainibe et al. 6219 (P, TAN). — Forêt de Ranomafana , forêt de Talatakely, 21°18’S, 47°38’30’’E , 950 m, 25.VIII.1991, Ranarijaona et al. 91 (P). — Toamasina , PN de Zahamena, Ankosy, 17°29’S, 48°44’E, 1107m, 27.I.2001, Rasolohery 204 ( MO, P, TAN). — Idem , 17°30’14’’S, 48°43’52’’E , 1100-1330 m, 30.I.2001, Rasolohery 290 ( MO, P, TAN). — Idem , 17°41’S, 48°59’E , 650 m, 15.VI.2001, Rasolohery 546bis ( MO, P, TAN). — Moramanga , 18°56’20’’S, 48°13’40’’E , 900 m, 1.II.1959, Schlieben 8114 (B, BM, BR, G, K) .

FIELD OBSERVATIONS. — Trunk: HT up to 8 m, DT 9.5-15(-20) cm, dead petiole bases persistent in the upper quarter only, caducous below and the leaf scars exposed; trunk surface coarsely and bluntly muricate, dark brown to black, some scales persist occasionally; base of the trunk usually considerably thickened by a mantle of adventitious roots.

Petiole: with a subcontinuous row of white to light brown aerophores, up to 2 cm long, on either side; petiole base long sigmoid.

Leaf scars: 3-4 × 5-9 cm, elliptic, about 5-7 very big and shallow orifices on their lower rim; spirally arranged.

Crown: more or less horizontal with stiff petioles and rachises to umbrella-shaped with arching petioles and rachises.

Trunk apex: densely covered with black scales, usually concealed by the more or less spaced petioles; some dead leaves persistent and hanging from the apex.

Lamina: elliptic; LL (130-)185-250(-300) cm, WL 110- 160 cm, FW 70-95 cm, NP 10-20.

DESCRIPTION

Petiole: (20-)40-80(-120) cm long, 3.5-4 cm in diameter; completely blackish to violaceous brown, adaxially sometimes green, distantly and bluntly muricate; covered with a thin caducous tomentum of light brown squamules; always 1 or 2 pairs of aphlebioid pinnae with a much reduced lamina, 15- 25 cm long, inserted at 5-30 cm from the petiole base, distinctly petiolate, usually positioned just above the trunk apex.

Lamina: bipinnate-pinnatifid to tripinnate, herbaceous to subcoriaceous (not coriaceous), fertilesterile dimorphism present; shiny green to dark green above, pale green below, lamina base shortly attenuate to truncate, basal pinnae patent to slightly reflexed, conduplicate; rachis of the same colour as the petiole, gradually becoming green distally.

Largest pinnae: 50-80cm long, distant by 10-20cm, adjacent pinnae contiguous to overlapping; costae and costulae of the same colour as the rachis.

Largest pinnules: 7.5-11 × 1.2-2.2 cm (fertile), 10.5-13.5 × 2-3 cm (sterile), adjacent pinnules contiguous to spaced by less than their width, linear-oblong to rarely triangular, their apex caudate, divided to the costula into broadly adnate segments, the 0-2 proximal segment pairs sessile and often deeply crenate; fertile pinnule segments 0.2-0.3 cm wide, spaced by less than to more than their width, slightly falciform to straight, their margin crenate to serrate to rarely subentire, flat or slightly revolute, their apex rounded, the bases of adjacent segments widened and confluent from the lower third of the pinnule; sterile pinnule segments 0.3-0.4 cm wide, spaced by less than their width, with a subentire to serrulate margin and a rounded to obtuse apex, the bases of adjacent segments usually confluent from the lower half of the pinnule or from its base; lateral veins in the segments once furcate, rarely simple.

Scales and hairs: scales present from the petiole base upwards to 15-30 cm on the petiole, at its base distant to contiguous, at most slightly overlapping, persistent, narrowly triangular, 1-1.5 × 0.1-0.2 cm, straight, slightly contorted further up on the petiole, their apex at most slightly crispate, not twisted, shiny dark brown to black, with a very narrow, light brown, erose margin, at the petiole base paralleling the petiole surface from a very conspicuously indurated and antrorsely curved base, completely appressed further up on the petiole, coriaceous; adaxial face of the rachis, costae and costulae more or less densely tomentose with rather soft, crispate, brown, multicellular hairs; sparse and caducous, ciliate, soft, crispate, light brown scales on the abaxial face of the costulae and segment midveins in young leaves; leaf otherwise glabrous.

Sori: very close to the midvein, contiguous, about 0.1 cm in diameter, covering the entire segment; indusia hemitelioid, i.e. scale-like and inserted at the costular face of the receptacle, with an entire to dentate rim, light brown, membranous to subcoriaceous, often obscured by the sporangia; receptacle columnar, conspicuously elongate, longer than the rim of mature indusia, with inconspicuous filiform paraphyses.

DISTRIBUTION

Madagascar, in the eastern rainforests from the north (Marojejy massif) to the south (Andohahela massif); endemic.

ECOLOGY

(50-)400-1300(-1400) m. Dense evergreen rainforests, also frequently on forest margins, in clearings or on roadsides.

REMARKS

The species is easily distinguished by its short, contiguous to distant, dark petiole scales, aphlebioid pinnae inserted near the petiole base and hemitelioid indusia. Detached pinna have been confounded with C. boivinii , although that species has a shorter receptacle and cup-shaped to asymmetric, but never truly hemitelioid indusia. Indusia of C. boivinii are always prominent and never completely concealed by the sporangia.

TYPIFICATION AND SYNONYMY

None of the sheets of Meller s.n. at K, one carrying a sterile apex and partly fertile middle pinnae and a second with two detached aphlebioid pinnae from the petiole base, carries the name of the species. As only the former sheet contains the manuscript remark “junghuhniana?” referenced in a remark following the protolog and as the protolog does not mention the aphlebioid basal pinnae, we choose the sheet carrying the sterile apex and partly fertile middle pinnae (K000009937) as the lectotype of Hemitelia melleri Baker. Indusium shape being variable in many tripinnate taxa and C. melleri having been occasionally misidentified, we consider it useful to designate an epitype including characters of the petiole base, i.e. scales and aphlebioid pinnae, as well as complete sterile and fertile pinnae to illustrate the leaf dimorphism.

Hildebrandt 4176 is widely distributed and most sheets carry Hildebrandt’s original label with the name “ Hemitelia glandulosa Kuhn n. sp. ”, most likely in Kuhn’s writing. Th ey are furthermore equivalent in carrying fertile middle pinnae. One sheet of the material seen at B, B200129546, is here designated as the lectotype of Hemitelia glandulosa Kuhn ex Baker. Although no petiole base is available for Hildebrandt’s collection, the material is identical to C. melleri with respect to lamina cutting and indusium shape.

All sheets of Perrier de la Bâthie 14746 seem to be located at P and are equivalent in carrying the determination “ Cyathea virescens C.Chr. sp. nov. ” in Christensen’s handwriting. We here designate a 3-sheet lectotype for C. melleri var. virescens C.Chr. including a leaf apex, middle pinnae and the petiole with an aphlebioid pinna, P00418729- 31. Th is taxon cannot be maintained as a distinct variety as it has been described from a young leaf with incompletely developed sori and a grass-green herbaceous (not coriaceous) lamina, characters that have been observed in young leaves of typical C. melleri during fieldwork. Considering all material seen of C. melleri , we cannot find a single morphological character to distinguish Perrier de la Bâthie 14746.

Figures

FIG. 38. — Cyathea lastii Baker: A, pinnules adaxially with a fragment of the costa; B, pinnule segments abaxially with a fragment of the costula; C, part of a pinnule segment abaxially with sori, dorso-lateral view, note the cup-shaped indusium and the distance between adjacent sori; D, habit; E, leaf scars and trunk surface; F, scale from the base of the petiole; G, basal part of the leaf (from the petiole base up to the first pinna pair), lateral view, one half of the petiole pruned away by longitudinal section. A-C, F, G, Janssen et al. 2430 (P); D, Janssen et al. 2431 (P); E, Janssen et al. 2381 (P). Scale bars: A, E, 1 cm; B, 0.5 cm; C, F, 0.1 cm; D, 1 m; G, 5 cm.

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

FIG. 52. — Photographic documentation of field characters in Cyathea: A-D, F, habit; A’-D’, F’, trunk apex; A”-D”, F”, trunk surface at breast height; A-A”, Cyathea boivinii Mett. ex Kuhn var. boivinii; B-B”, C. boivinii var.hildebrandtii (Kuhn) Janssen & Rakotondr.; C-C”, C. boivinii var. andringitrensis Janssen & Rakotondr.; D, D’, C. kirkii Hook.; F-F”, C. lastii Baker; A”, the petiole bases are sometimes completely decayed on older trunks (left), but persistent in most trunks observed (right); E, C. boivinii var. andringitrensis Janssen & Rakotondr., dense tomentum of intricate squamules on the petiole; A, A’, A” (left), C, C’, uncollected; A” (right), Janssen et al. 2604; B, Janssen et al. 2433; B’, Janssen et al. 2442; B”, Janssen et al. 2464; C”, Janssen et al. 2594; D, D’, Labat et al. 3208; E, Janssen et al. 2591; F-F”, Janssen et al. 2431. Vouchers at P. Photos A’, A” (left), B’, B”, C-C”, E by G. Rouhan (MNHN); D, D’ by J.-N. Labat (MNHN); all other photos by T. Janssen. Scale bars: A-D, F, 1 m; A’-D’, F’, A”-C”, F”, 10 cm; E, 1 cm.

FIG. 1. — Spores of Cyathea sect. Alsophila in Madagascar and on the Comoros: A-C, spores of taxa belonging to the bipinnate clade (Janssen et al. 2008) have a perispore formed by dense slender rodlets; D-F, spores of taxa belonging to the tripinnate clade (Janssen et al. 2008) have a perispore with prominent crests; A, C. approximata Bonap.; B, C. kirkii Hook.; C, C. serratifolia Baker; D, C. boivinii Mett.ex Kuhn var.boivinii; E, C. melleri (Baker) Domin;F, C. similis C.Chr.,a young spore with a distantly verrucose perispore (Janssen & Rakotondrainibe 2006; Gastony & Tryon 1976). A, Janssen et al. 2449 (P); B, Rakotondrainibe et al. 6751 (P); C, Rakotondrainibe et al. 6230 (P); D, Janssen et al. 2481 (P); E, Janssen et al. 2536 (P); F, Janssen et al. 2576 (P). Scale bars: 10 μm.

FIG. 39. — Cyathea melleri (Baker) Domin:A, fertile pinnules abaxially with a fragment of the costa, sori only partly indicated; B, sterile pinnules abaxially with a fragment of the costa; C, pinnule segments abaxially with a fragment of the costula (at the junction with the costa), sporangia partly removed to reveal the hemitelioid indusia; D, habit; E, leaf scars and trunk surface; F, basal part of the leaf (from the petiole base up to the first pinna pair), basal part in lateral view, folded middle part in adaxial and upper part in dorsal view, note the presence of a pair of reduced pinnae at 5-10 cm above the base of the petiole; G, pinnule segments adaxially with a fragment of the costula (at the junction with the costa), form with narrow segments; H, scales from the base of the petiole (left: lateral view; right: dorsal view). A-C, E, F, H, Janssen et al. 2569 (P); D, uncollected, photograph at P; G, Rakotondrainibe et al. 5866 (P). Scale bars: A, B, 1 cm; C, H, 0.1 cm; D, 1 m; E, F, 5 cm; G, 0.5 cm.

FIG. 53. — Photographic documentation of field characters in Cyathea: A-D, habit; A’-D’, trunk apex; A”-D”, trunk surface at breast height; A-A”, Cyathea melleri (Baker) Domin; B-B”, C. sechellarum Mett.; C-C”, C. similis C.Chr.; D-D”, C. valdesquamata Janssen & Rakotondr. A, A’, B’, B”, uncollected; A”, Janssen et al. 2569; B, Janssen et al. 3247; C, Janssen et al. 2402; C’, Janssen et al. 2576; C”, Janssen et al. 2393; D, D”, Janssen et al. 2907; D’, Janssen et al. 2904. Vouchers at P. Photos A’-A”, C’ by G. Rouhan (MNHN); all other photos by T. Janssen. Scale bars: A-D, 1 m; A’-D’, A”-D”, 10 cm.