MARATTIACEAE [Draft]
合囊蕨科 he nang jue ke
Zhang Xianchun (张宪春)[1]; Cristina H. Rolleri 2
Angiopteridaceae C. Chr. in F. verdoorn (Ed.), Man. Pterid.: 527. 1938
Christenseniaceae Ching, Bull. Fan Mem. Inst. Biol, Bot. 10: 227-229. 1940
Danaeaceae C. A. Agardh. Aphor.117. 1822
Kaulfussiaceae C. Presl, Suppl. Tent. 17. 1845
Marattiaceae Bercht. & J. S. Presl, Přirozen. Rostlin 1:272. 1820. non Kaulf. Enum. 31. 1824.
Terrestrial or amphibious ferns. Erect or postrate, carnose, amylaceous, polycyclostelic to dictyostelic, naked rhizomes, without sclerenchymatic tissues. Fronds entire or divided, 1-4 pinnate, monomorphic or dimorphic. Stipes flexible or rigid, with basal, amylaceous stipules. Cortical spines, aerophores and pulvinules (nodes) present or absent. Stipes, costae and costulae with sclerenchymatic tissues. Laminae entire or divided (ternate, palmate, 1-4-pinnate), very small (ca. 0.20 cm) to large (ca. 6 m). Veins free, simple or anastomosing. Venuloids present or absent. Indument eglandular (proscales) and glandular (trichomes). Secretory mucilage system. Idioblasts containing opal A in the epidermis and/or the inner tissues. Marattiaceous, cyclocytic or porocyclocytic stomata. Intercellular pectic protuberances (IPP) in the mesophyll cells. Eusporangia free (sori) or partially to completely fused (synangia), with paraphyses. Spores trilete and monolete, with bulliform, muriform or stelliform ornamentation.
A family of five genera: Angiopteris, Archangiopteris, Christensenia, and Danaea. Homogenous cytology of the genera based on an x=40 (39). All genera bear IPP AND share the ability to biomineralize SiO2 and produce opal. According to Ching’s system (1978), this group of primitive ferns is divided into rather small families, including Angiopteridaceae, Christenseniaceae, and Danaeaceae.
Key to genera of the Marattiaceae
1. Fronds entire, ternate or palmate; veins anastomosing, with polygonal areoles, and free veinlets; glandular indument; sinangia circular in outline, without paraphyses; opal epidermal idioblasts present; spores trilete, ornamentation stelliform, sporoderm echinate; plants small to medium, 30-60 cm. Christensenia
1. Fronds entire or 1-4 pinnate, dimorphic or monomorphic; veins free, simple, geminate and/or furcate; venuloid present or absent; eglandular or glandular indument; sinangia (ellipsoidal) or sori; piliform branched paraphyses; opal epidermal idioblasts present or absent; spores trilete or monolete, ornamentation stelliform, muriform and/or bulliform; plants small (0.20 m) to large (6-7 m). 2
2. Fronds dimorphic, entire or 1-pinnate; rhizomes postrate to suberect, never massive; venuloid absent; glandular indument, mucilaginiferous proscales and trichomes; sinangia to 200-240 fussed eusporangia, without paraphyses; opal epidermal idioblasts absent; spores monolete, ornamentation stelliform, sporoderm echinate; plants very small ( 0.20 m) to large (3 m). Danaea
2. Fronds monomorphic, 1-4 pinnate; rhizomes postrate or erect and massive; venuloid present or absent; eglandular indument, only proscales; sinangia or sori; branched multicelular paraphyses always present; opal epidermal idioblasts always present; spores trilete or monolete, ornamentation stelliform, muriform and/or, bulliform. 3
3. Plants large, to 6-7 m; rhizomes massive, to 0,50 m; fronds arising in clusters, approximate, 1-4 pinnate, imparipinnate or paripinnate; spores trilete or monolete. 4
3. Plants smaller, rarely 2-3 m; rhizomes postrate to suberect, not massive; fronds not clustered; 1-pinnate, imparipinnate or paripinnate; pinna escarce, usually a few pairs; venuloid commony present; sori with 40-140 free, sometimes partially fussed eusporangia; spores trilete, ornamentation stelliform or muriform, sporoderm crestate, echinate or other, with heterogeneous micrornamentation. Archangiopteris
4. Fronds 1-pinnate, imparipinnate; venuloid present, short to long and reaching the costa; marginal sori with 6-12(14) free eusporangia; spores trilete, ornamentation bulliform, sporoderm pustulate. Angiopteris
4. Fronds 1-4 pinnate, imparipinnate; venuloid short or absent; intramarginal to medial, ellipsoid sinangia attached to protruding receptacles; spores monolete, ornamentation muriform, bulliform or stelliform (uncommon), sporoderm rugate, rugulate, pustulate or echinate (rare). Marattia
1. MARATTIA Swartz, Prodr. 128. 1788.
合囊蕨属 he nang jue shu
Discostegia C. Presl, Suppl. Tent. : 11(postpr. 281). 1845.
Eupodium Hook., Gen. Fil. t. 118. 1842.
Gymnotheca C. Presl, Suppl. Tent.: 12 (postpr. 282). 1845.
Myriotheca Juss., Gen. Pl.: 15. 1789.
Stibasia C. Presl, Suppl. tent. pterid.: 15 (postpr. 285). 1845.
Plants terrestrial, to 3-4 m. Erect, massive, naked, amylaceous, polycyclostelic, rhizomes, to 40 cm high. Fronds arising radially, clustered, up to 4 m long. Stipes basally swollen flanked basally by 2 large, fleshy stipules, with aerophores, cortical spines, and deeply fimbriate, auriculate proscales, polycyclostelic, with a 1–2 small central bundles; pulvinules absent. Lamina 1- to 4-pinnate, ultimate rachises usually winged. Pinnules sessile or short-stalked, ovate-lanceolate to oblong, apexes acuminate, serrate to dentate, margins crenate, lobate or serrate. Veins free, simple or furcated. Venuloids short or absent. Superficial linear or narrowsly lanceolate, entire to fimbriate proscales. Synangia medial to submarginal, ellipsoidal, bivalved, with 6-14 fussed eusporangia; protruding receptacles with multicellular branched paraphyses. Spores monolete, ellipsoid, 21-25 x 13-18 µm, muriform or bulliform (rarely stelliform) macrornamentation. x = 39, 40, 78.
A pantropical genus of some 70 or more described species. Seven species recognized for the New World (Lavalle 2003; 2005; 2007). The Old World species are in need of monographic study, and only 8 were revised (Rolleri et al., 2001; Lavalle & Stampella, 2004). Basic chromosome numbers were suggested to be related to geographical distribution, an x = 39 asigned to paleotropical species, and an x = 40 to neotropical species (Takamiya, 1995). Represented in China by a single species found only on the small Orchid Island in the SE sea of Taiwan.
1. Marattia sylvatica Blume, Enum. Pl. Javae: 256. 1828.
Marattia sambucina Blume, Enum. Pl. Javae: 256. 1828. Marattia pellucida C. Presl, Suppl. Tent. Pterid. 10. 1845. Marattia ternatea de Vriese & Harting, Monogr, Marattiac.:4, tab. 3, fig. 16. 1853. Marattia vestita H.- Christ, Philip. J. Sci. 2: 185. 1907. Marattia brooksii Copel., Philip. J. Sci. 7:59. 1912.
Erect rhizomes, up to 40 cm high. Fronds up to 3 m long; stipes shorter than lamina, ca. 2 cm thick, bearing irregularly toothed narrowly lanceolate proscales, ca. 5 mm long and cortical spines. Laminae oblong in outline, 3-pinnate, pinnae up to 60 cm, pinnules 6–10 × 1–1.5 cm, narrowsly elliptic, base cuneate, apex acuminate, margin sub-entire at base, then serrate, finally deeply and notably serrate apically; veins free, 1.5–2 mm apart; texture herbaceous to chartaceous; small, castaneous, thin proscales scales with bullate base along costa. Medial or supramedial synangia, 1–2.2 mm long, with to 8-9 cellular uniseriate paraphyses, ramified paraphyses rare. Epidermal pattern of pinnules deeply sinuate, irregularly thickened at the angulose curves; large opal idoblasts, irregular in outline.
Under forests on slopes. Taiwan (Orchid Island) [Also Malaysia, Borneo, Java, Philippines, and Celebes, Molucas, New Guinea, Admiralty, Caroline, Vanuatu, and Solomon Islands; in rainforests, montane forests, in rideges and forested canyons, near rivers, up to 2000-2100 m altitude].
[1] Herbarium, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China.
2 , Laboratorio de Estudios de Anatomía Vegetal Evolutiva y Sistemática (LEAVES), Facultad de Ciencias Naturales y Museo de La Plata, 64 entre 120 y diagonal 113, B1904 DZB, La Plata,Argentina.