Sphenostemonaceae P. van Royen et Airy Shaw in Kew Bull. 27: 325. 23 Oct 1972; Paracryphiales Takht. ex Reveal in Novon 2: 239. 13 Oct 1992; Quintiniaceae Doweld in Byull. Mosk. Obshch. Ispyt. Prir., Biol. 105(5): 60. 9 Oct 2000; Quintiniales Doweld, Tent. Syst. Plant. Vasc.: lii. 23 Dec 2001; Sphenostemonales Doweld, Tent. Syst. Plant. Vasc.: li. 23 Dec 2001
Genera/species 3/28–31
Distribution Central and East Malesia to eastern Australia, New Caledonia and New Zealand.
Fossils Fossil pollen grains of Quintinia have been reported from the Oligocene to the Pliocene of Australia, and fossil leaf fragments of Paracryphia from the Early Miocene of New Zealand (Pole 2010).
Habit Usually bisexual (Paracryphia and Quintinia sometimes unisexual, andromonoecious), evergreen trees or shrubs.
Vegetative anatomy Phellogen ab initio superficial. Primary medullary rays narrow (Paracryphia). Primary vascular tissue a cylinder, without separate vascular bundles. Vessel elements with scalariform perforation plates (often with much more than 40 transverse ribs); lateral pits scalariform to opposite, simple (Sphenostemon) or bordered (Paracryphia, Quintinia) pits. Vestured pits present (Quintinia). Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits, non-septate. Wood rays uniseriate and multiseriate, heterocellular (in Paracryphia secondarily very primitive). Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty. Secondary phloem in Paracryphia of very primitive structure (secondarily), in Sphenostemon stratified. Sieve tube plastids S type (in Paracryphia very similar to those in Hibbertia in Dilleniaceae). Nodes usually 3:3, trilacunar with three leaf traces (in one species of Quintinia 5:5?, pentalacunar with five? traces). Parenchyma with calciumoxalate as crystal sand, styloids and/or elongate and acicular crystals present in Paracryphia and often in Sphenostemon; druses present in Quintinia.
Trichomes Hairs unicellular, simple (in Paracryphia ferrugineous); Quintinia with multicellular glandular hairs with multiseriate stalk and multicellular peltate head. Peltate scales abundant (in Quintinia very frequent on vegetative parts).
Leaves Alternate (spiral), simple, entire, coriaceous, with ? ptyxis. Stipules and leaf sheath absent or, in Sphenostemon, minute, cauline. Petiole vascular bundle transection in Paracryphia flattened-annular, petiole with medullary bundles; in Sphenostemon three bundles, with transection annular and petiole with wing bundles, or transection arcuate. Venation pinnate, in Quintinia semicraspedodromous to brochidodromous. Stomata anomocytic. Cuticular wax crystalloids? Mesophyll in Paracryphia with sclerenchymatous idioblasts (thin-walled brachysclereids) containing calciumoxalate styloids; abaxial side of lamina in Sphenostemon sometimes with styloids. Leaf margin usually serrate (in Quintinia and Sphenostemon occasionally entire), in Paracryphia with single vein running into tooth apex.
Inflorescence Terminal, cymose or racemose; in Paracryphia spikes in branched panicle, densely beset with ferrugineous hairs; in Sphenostemon raceme; in Quintinia raceme or panicle.
Flowers Zygomorphic, small. Hypogyny (Paracryphia, Sphenostemon) or epigyny to half epigyny (Quintinia). Sepals? (floral prophylls?, bracts?) in Paracryphia usually four (rarely five), with decussate-cochlear aestivation, caducous, free, outer larger and enclosing remaining three; sepals in Quintinia four or five, with decussate or imbricate aestivation, free, persistent; sepals in Sphenostemon four, with decussate aestivation, caducous, free. Petals probably absent in Paracryphia; in Quintinia four or five, with decussate or imbricate aestivation, free, caducous; in Sphenostemon four to six, decussate, fleshy, caducous, free (absent in some species). Nectary and disc absent in Paracryphia; disc absent in Sphenostemon (nectary?); nectariferous disc present on ovary apex in Quintinia.
Androecium Stamens in Paracryphia eight (to eleven), in one whorl,triplostemonous; in Quintinia four or five, haplostemonous; in Sphenostemon four (to 13), in one whorl, haplostemonous, antesepalous, fleshy and often petaloid. Filaments in Paracryphia stout, free from each other and from tepals, in bisexual flowers filiform, in male flowers broad and flat; in Sphenostemon stout, often very short, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, with thecae embedded in thick connective, latrorse to introrse, longicidal (dehiscing by longitudinal slits). Tapetum at least in Quintinia secretory. Staminodia usually absent (present in female flowers in some species of Quintinia).
Pollen grains Microsporogenesis simultaneous (Quintinia). Pollen grains tricolporate (Paracryphia), tetracolporate to hexacolporate (Quintinia) or tri- or tetraporate (Sphenostemon) (rarely inaperturate), shed as monads, bicellular (Quintinia) at dispersal. Exine tectate-perforate to semitectate, with columellate infratectum, in Paracryphia reticulate to almost rugulate, in Sphenostemon reticulate to scabrate or finely verrucate, in Quintinia usually striate-rugulate to radiate-rugulate (sometimes perforate or smooth).
Gynoecium Pistil in Paracryphia composed of eight to 15 laterally conduplicate (ventrally adnate to central ovary tissue), in Quintinia three to five, in Sphenostemon two connate carpels, in Paracryphia attached to central columella. Ovary superior (Paracryphia, Sphenostemon) or inferior to semi-inferior (Quintinia), in Paracryphia octalocular to 15-locular, in Quintinia entirely or partially trilocular to quinquelocular, in Sphenostemon bilocular. Style in Quintinia single, simple, long; absent in Paracryphia and Sphenostemon. Stigmas in Paracryphia central, separate, conduplicate folded, type?; stigma in Quintinia capitate, trilobate to quinquelobate, Wet type; stigma in Sphenostemon large, capitate, type? Pistillodium usually absent (present in male flowers in some species of Quintinia).
Ovules Placentation inParacryphia axile, in Quintinia parietal or axile, in Sphenostemon apical. Ovules four (Paracryphia), numerous (Quintinia) or one (or two) pendulous (Sphenostemon) per carpel, anatropous, unitegmic (Paracryphia, Sphenostemon) or bitegmic (Quintinia), crassinucellar (Paracryphia, Sphenostemon) or tenuinucellar (Quintinia). Micropyle in Quintinia endostomal. Integument three or four cell layers thick (Paracryphia). Funicular obturator present in Sphenostemon. Megagametophyte monosporous, Polygonum type. Endosperm development cellular. Endosperm haustorium possibly micropylar (Quintinia). Embryogenesis?
Fruit In Paracryphia a capsule-like schizocarp with eight to 15 septicidal follicle-like mericarps; carpels contracting acropetally and dehiscing adaxially, ripe carpels separating from persistent central columella except at apex, to which they are adnate through two lignified ventral carpellary vascular bundles; in Quintinia a septicidal capsule with persistent calyx; in Sphenostemon an asymmetrical berry-like drupe with one or two pyrenes.
Seeds Aril absent. Testa in Paracryphia and almost all species in Quintinia winged. Exotestal cells? in Paracryphia with sinuous anticlinal walls and inner walls lignified. Endotestal cells in Sphenostemon with dark-staining content. Perisperm not developed. Endosperm copious (in Sphenostemon often ruminate). Embryo straight, well differentiated (Paracryphia), chlorophyll? Radicula in Paracryphia longer than cotyledons. Cotyledons two. Germination?
Cytology n = 22? (Quintinia), 23?
DNA
Phytochemistry Very insufficiently known. Route I secoiridoids (Quintinia), ellagic acid (Quintinia), hydrolyzable tannins (some species of Quintinia), and saponins present. Flavonoids? Alkaloids? Aluminium accumulated in Quintinia.
Use Timber (Quintinia).
Systematics Paracryphiaceae may be sister to Dipsacales (Winkworth & al. 2008), although they sometimes appear as sister-group to the clade [Apiales+[Bruniales+Dipsacales]].
A possible topology is [Sphenostemon+[Paracryphia+Quintinia]] (Lundberg 2001).
1/1. Paracryphia (1; P. alticola; New Caledonia). – Evergreen tree. Intervascular pits bordered. Hairs unicellular, ferrugineous. Petiole vascular bundle transection flattened-annular; petiole with medullary bundles. Inflorescence panicle with branched spikes. Hypogyny. Tepals with decussate-cochlear aestivation, caducous. Petals probably absent. Disc absent. Stamens eight (to eleven), in one whorl, twice the number of tepals. Filaments stout, thecae more or less embedded in connective. Pollen grains tricolporate. Exine reticulate to almost rugulate. Carpels eight to 15, inserted at central columella. Style absent. Stigmas central, separate, conduplicate folded. Placentation axile. Ovules four per carpel, unitegmic, crassinucellar. Fruit a capsule-like schizocarp with eight to 15 septicidal follicle-like mericarps; carpels contracting acropetally and dehiscing adaxially, ripe carpels separating from persistent central columella except at apex, to which they are adnate through two lignified ventral carpellary vascular bundles. Columella persistent. Testa winged. Exotestal cells? with sinuous anticlinal walls and inner walls lignified.
The wood anatomy of Paracryphia has been considered one of the most primitive among angiosperms, although these features must be interpreted as reversals. The vessel elements are up to 1,76 μm long in the mature wood. The perforation plates are very oblique and scalariform, with up to 203 transverse bars. The lateral walls have scalariform or transitional intervascular pitting.
1/c 20. Quintinia (c 20; Central Malesia from the Philippines to New Guinea, New Caledonia, eastern Queensland, eastern New South Wales, New Zealand). – Evergreen trees or shrubs. Intervascular pits bordered. Vestured pits present. Entire plant more or less covered with peltate scales. Peltate glands present. Leaves alternate (spiral). Stipules absent. Leaf margin serrate. Inflorescence raceme or panicle. Epigyny or half epigyny. Sepals and petals free. Nectariferous disc present on top of ovary. Stamens four or five. Pollen grains 4–6-colporate. Exine usually striate-rugulate to radiate-rugulate (sometimes perforate or smooth). Carpels three to five. Style single, simple, long. Stigma capitate, trilobate to quinquelobate. Placentation parietal or axile. Ovules numerous per carpel, tenuinucellar, probably bitegmic. Micropyle endostomal. Fruit a septicidal capsule with persistent calyx. Testa usually winged. n = 22? Route I secoiridoids, ellagic acid, hydrolyzable tannins, and saponins present. Aluminium accumulated.
The vessel elements are up to 1,00 μm long in the mature wood. The perforation plates are oblique and scalariform, with up to 116 transverse bars. The lateral walls have opposite to scalariform intervascular pitting.
According to analyses of mitochondrial genes (Soltis & al. 2011), Quintinia is closely allied to Polyosma (Escalloniaceae), possibly an example of horizontal gene transfer. The presence of two integuments in the ovules in Quintinia needs to be confirmed.
Friis & al. (2013) compared flowers of Quintinia with the Late Cretaceous Silvianthemum suecicum and Bertilanthus scanicus and found important similarities in, e.g., the secretory hairs on the floral surface and pedicel, the quincuncial free petals, the small microsporangia with quadriaperturate pollen grains, the postgenital fusion of the apocarpous styles, the placentation on incomplete septa, and bottle-shaped epidermal cells with long thin papilla on the carpels.
1/7–10. Sphenostemon (7–10; Sulawesi to New Guinea, eastern Queensland and New Caledonia). – Usually small evergreen trees. Intervascular pits simple. Secondary phloem stratified. Leaves alternate (spiral). Stipules absent. Petiole with three bundles, with transection annular and petiole with wing bundles, or transection arcuate. Leaf margin coarsely serrate. Inflorescence raceme. Flowers relatively small. Hypogyny. Sepals and petals free. Stamens four (to twelve). Filaments stout. Pollen grains tripor(or)ate. Exine reticulate to scabrate. Carpels two. Style absent. Stigma large, capitate. Placentation apical. Ovules one (or two) per carpel, pendulous, unitegmic. Funicular obturator present. Fruit a two-seeded drupe. Testa not winged. Endotestal cells with dark-staining content.
The vessel elements are up to 3,50 μm long in the mature wood. The perforation plates are very oblique and scalariform, with up to 185 transverse bars. The lateral walls usually have opposite (sometimes transitional) intervascular pitting.
Literature
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