PETROSAVIIDAE S. W. Graham et W. S. Judd

Graham et Judd in Taxon 56: E25. Aug 2007


[Petrosaviaceae+[Taccales+Pandanales]+[Liliales+[Iridales+Commelinidae]]]


PETROSAVIACEAE Hutch.

Hutchinson, Fam. Fl. Pl. 2: 36. 20 Jul 1934, nom. cons.

Miyoshiaceae Nakai in J. Jap. Bot. 17: 190. Apr 1941; Miyoshiales Nakai in J. Jap. Bot. 17: 189. Apr 1941; Japonoliriaceae Takht. in Bot. Žurn. 81(2): 85. Mai-Jun 1996; Petrosaviales Takhtajan, Divers. Classif. Fl. Pl.: 577. 24 Apr 1997; Petrosavianae Doweld, Tent. Syst. Plant. Vasc.: lx. 23 Dec 2001

Genera/species 2/3–4

Distribution Eastern China, Japan, Taiwan, Southeast Asia, West and Central Malesia.

Fossils Unknown.

Habit Bisexual, perennial herbs. Achlorophyllous holomycotrophic (Petrosavia) or autotrophic (Japonolirion).

Vegetative anatomy Roots fibrous; root stele triarch or tetrach (Petrosavia); root in Petrosavia without medulla; root cortex consisting of four to six layers of parenchyma cells containing fungal hyphae. Phellogen absent. Rhizome with scale-like leaves. Stem vascular bundles forming a cylinder. Vessels present in Japonolirion, usually absent in Petrosavia (roots sometimes containing vessels). Vessel elements with scalariform perforation plates; lateral pits? Imperforate tracheary xylem elements tracheids. Wood rays absent. Axial parenchyma? Sieve tube plastids P2ccp type, with starch, many cuneate protein crystals and one polygonal protein crystal. Nodes? Calciumoxalate as raphides and druses present in Petrosavia.

Trichomes Hairs absent.

Leaves Alternate (spiral), simple, entire, in Petrosavia scale-like, with ? ptyxis. Stipules absent; leaf sheath short? Venation parallelodromous. Stomata anomocytic. Cuticular wax crystalloids? Leaf margin entire.

Inflorescence Terminal (Petrosavia) or axillary (Japonolirion), raceme. Floral prophyll (bracteole) sublateral or absent.

Flowers Actinomorphic, small. Hypogyny (Japonolirion) or almost epigyny (Petrosavia). Tepals 3+3, with imbricate aestivation, petaloid, persistent, free (Japonolirion) or connate at base (Petrosavia); in Japonolirion sepaloid, tepals of outer and inner whorls of slightly different shape. Septal nectaries infralocular. Disc absent.

Androecium Stamens 3+3. Filaments subulate, free from each other and from tepals, in Petrosavia adnate to tepal bases. Anthers basifixed (Japonolirion) or dorsifixed (Petrosavia), non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains monosulcate, shed as monads, bicellular at dispersal. Exine intectate (Japonolirion) or semitectate (Petrosavia), with columellate infratectum, gemmate (Japonolirion) or reticulate (Petrosavia).

Gynoecium Pistil composed of three carpels connate below or at base only (Petrosavia), or free (Japonolirion); carpel plicate and ascidiate, fused by papillae on septal side; carpel fusion usually postgenital (rarely also congenital). Ovary superior (Japonolirion) or semi-inferior (Petrosavia), trilocular below, unilocular above. Stylodia three, short, recurved (Japonolirion). Stigmas decurrent (Japonolirion) or subcapitate (Petrosavia), type? Pistillodium absent.

Ovules: Placentation dorsal. Ovules usually four (Japonolirion) or numerous (Petrosavia) per carpel, anatropous (in Petrosavia possibly also campylotropous), bitegmic, crassinucellar (or semicrassinucellar). Micropyle endostomal. Outer integument two cell layers thick. Inner integument two cell layers thick. Hypostase not formed. Parietal cell formed from archesporial cell. Integumentary obturator present in Petrosavia. Nucellar cap two cell layers thick, functioning as obturator. Megaspore tetrad T-shaped. Megagametophyte in at least Petrosavia monosporous, Polygonum type. Antipodal cells persistent (Japonolirion). Chalazal chamber not formed. Endosperm development ab initio cellular (in Petrosavia possibly also nuclear?). Endosperm haustoria absent. Embryogenesis?

Fruit A septicidal capsule (Japonolirion) or a follicle with often more or less persistent tepals (Petrosavia).

Seeds Aril absent. Seed coat testal-endotegmic. Testa unwinged (Japonolirion) or winged (Petrosavia). Exotestal cells often collapsed. Endotesta well developed. Exotegmen well developed. Endotegmen crushed. Perisperm not developed. Endosperm copious, oily and proteinaceous. Suspensor formed. Embryo small, in Petrosavia undifferentiated, chlorophyll? Cotyledon one. Radicula poorly developed. Germination?

Cytology n = 12, 13 (Japonolirion); n = 15, 30 (Petrosavia, also tetraploid)

DNA

Phytochemistry 3-O-glycosides of 6-C-glycosylquercetin and 6-C-glycosylkaempferol, isoorientin, quercetin-3-O-glycoside (isoquercitrin), quercetin-3-O-arabinoside (avicularin), vicenin-2, orientine, and chlorogenic acid present.

Use Unknown.

Systematics Petrosavia (2–3; eastern China including Hainan, Japan, Taiwan, Burma, Thailand, Vietnam, the Malay Peninsula, northern Sumatra, the Philippines), Japonolirion (1; J. osense; mountains on Hokkaido and Honshu in Japan, on serpentine).

Petrosaviaceae are sister to all monocots except Acorus and Alismatales.

Of the flavonoids investigated, the two flavonol-C-glycosides 6-C-glycosylquercetin-3-O-glycosides and 6-C-glycosylkaempferol-3-O-glycosides are unique to Petrosaviaceae (Iwashina & al. 2005).

The presence of P2ccp type sieve tube plastids in Petrosaviaceae and Tofieldiaceae is probably a plesiomorphy (Behnke 2000; Cameron & al. 2003). Some species of Velloziaceae (Pandanales) possess a superficially similar type of sieve tube plastids, P2cap (Behnke & al. 2000), and Lanaria (Lanariaceae, Iridales) has the type P2ccps (Behnke 2000).


Literature

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