Basal Angiosperms
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The Flowering Plants Handbook A practical guide to families and genera of the world James W. Byng CONTENTS Chapter 1: Introduction....................................................................... 1 Notes on how to use this book.................................................. 2 Useful diagnostic characters……………………….........…....…………… 3 Key to major plant groups.......................................................... 5 Chapter 2: Basal Angiosperms............................................................ 7 Amborellales……………….............................................................. 8 Nymphaeales………………………..................................................... 9 Austrobaileyales………................................................................. 11 Chloranthales............................................................................. 13 Cannellales……………………………………………………….........…....……… 14 Piperales………………………………………………………............…………….. 16 Magnoliales………………………………………………….........…….…………. 21 Laurales………………………………………………………….........………..……. 28 Chapter 3: Monocots............................................................................ 36 Acorales………………………............................................................. 37 Alismatales……………………………................................................... 38 Petrosaviales……………................................................................. 47 Dioscoreales............................................................................... 48 Pandanales………………………………………………………...........………….. 51 Liliales……………………………………………………………..….........…………. 56 Asparagales…………………............................................................. 65 Arecales……….……………………….................................................... 86 Commelinales……………............................................................... 88 Zingiberales................................................................................ 93 Dasypogonaceae………………………………………..……….........…………. 100 Poales..……………………………………………………….……….........………… 101 Chapter 4: Basal Eudicots ................................................................... 116 Ceratophyllales……………............................................................. 117 Ranunculales………………………..................................................... 118 Sabiaceae…………………................................................................ 130 Proteales.................................................................................... 131 Trochodendrales…………………………………………………...........………. 136 Buxales………………………………………………………………..........………… 137 Gunnerales………………….............................................................. 139 Dilleniaceae………………………….................................................... 141 Chapter 5: Rosids ……………………………………........................................... 142 Saxifragales…………………............................................................. 143 Vitales…….…………………………….................................................... 153 Zygophyllales..…………................................................................. 154 Fabales....................................................................................... 156 CHAPTER 2: BASAL ANGIOSPERMS Basal angiosperms are the earliest diverging lineages of flowering plants and the most commonly encountered are the laurels, magnolias and waterlilies. The group often exhibit ‘primitive’ or ancestral characters such as an open flower organisation, designed for large insect (e.g. beetle) pollination, and little distinction between the sepals and petals. They generally lack ‘advanced’ or derived characters such as fused petals and zygomorphy. Basal angiosperms are generally woody plants with notable herbaceous exceptions such as many Piperales and the aquatic families of the Nymphaeales. The leaves are simple and often aromatic containing ethereal oils. The flowers are spirally arranged or 3-merous and often have many perianth parts, stamens and carpels. The carpels are often free and the styles and stigmas are poorly developed. The stamens are usually laminar shaped with poorly differentiated anthers. 130. DAPHNIPHYLLACEAE Trees or shrubs; branchlets with leaf scars and lenticels. Leaves simple, alternate or rarely opposite, often clustered at branch ends; margins entire; petioles present; stipules absent. Inflorescences axillary racemes; bracts conspicuous. Flowers unisexual (plants dioecious), actinomorphic. Sepals free, ±imbricate or sometimes absent. Petals absent. Male flowers: stamen filaments often shorter than anthers; anthers basifixed. Female flowers: ovaries superior; carpels fused; locules 2; ovules (1-)2 per locule; placentation axile or apical; stigmas decurrent; sometimes staminodes. Fruits drupes. Genera 1/species ca. 30; Daphniphyllum. Distribution:India to Australia and East Asia. Floral formula: K(0-)3-6 C0 A5-14 G2(-4) References: Endress & Igersheim 1999; Fishbein et al. 2001, 2004; Huang 1965, 1966; Kubitzki 2007; Min & Kubitzki 2004. © KENPEI/ CC-BY-SA-3.0 © Steve Law/ CC-BY-2.0 Male flowers of Daphniphyllum Daphniphyllum macropodum 131. ITEACEAE teijsmannii Trees or shrubs, sometimes climbing (Itea). Leaves simple, alternate; margins often spiny toothed to entire; petioles present; stipules tiny or absent. Inflorescences racemes or panicles (Itea), or cymes to corymbs (Pterostemon). Flowers bisexual or rarely polygamous, actinomorphic, hypanthium. Sepals free or basally fused, valvate, persistent. Petals free, persistent, clawed (Pterostemon). Stamens alternating with petals (Itea) or sepals (Pterostemon); anthers dorsifixed, introrse; staminodes 5 (Pterostemon). Ovaries superior to part-inferior (Itea) or inferior (Pterostemon); carpels fused; locules 2 or 5; ovules 4-6 (Pterostemon) or many (Itea) per locule; placentation axile. Fruits capsules. Genera 2/species ca. 18. Distribution: Tropical to northern temperate regions. Floral formula: Itea K5 C5 A5 Ĝ2 Pterostemon K5 C5 A5+5° Ĝ5 Notes: Both genera were previously associated with Escalloniaceae. References: Bohm et al. 1999; Kubitzki 2007a, 2007b; Fishbein et al. 2001, 2004; Generic synopsis • Itea (southeast Asia to western Malesia, eastern North America, East © Wouter Hagens/ CC-BY-SA-3.0 © Imc/ CC-BY-SA-3.0 to South Africa; ca. 16 spp.). Itea virginica Itea ilicifolia • Pterostemon (much-branched shrubs; Oaxaca [Mexico]; ca. 2 spp.). 132. GROSSULARIACEAE GOOSEBERRY FAMILY Shrubs, sometimes climbing, often spiny; often glandular hairs. Leaves aromatic, usually deciduous, simple, alternate; venation pinnate to 3-palmate; margins lobed or toothed; petioles present; stipules usually present or absent. Inflorescences racemes, usually on short-shoots; sometimes bracteate, hairy. Flowers bisexual or rarely unisexual (plants dioecious, e.g. R. diacanthum), actinomorphic, hypanthium well-developed (and lobed) and petaloid; green, white, yellow or red; sometimes bracteolate. Sepals fused, persistent. Petals rarely absent or free, imbricate, sometimes interpretated as staminodes, smaller than sepals. Stamens opposite the sepals; anthers basifixed. Ovaries inferior to part inferior; carpels fused; locule 1; ovules 4-many; placentation parietal; style 2. Fruits berries, with persistent perianth. Genus 1/species ca. 150; Ribes. Distribution: Temperate northern hemisphere and Andes to southern South America. Floral formula: K(3-)5(-9) C(0-3-)5(-9) A4-5 Ĝ2 Useful species: R. nigrum (black currants); R. rubrum (red currants); R. uva-crispa (gooseberry). Notes: Ribes speciosum is 4-merous. References: Fishbein et al. 2001, 2004; Messinger et al. 1999; Morin 2009; Senters & Soltis 2003; Schultheis & Donoghue 2004; Weigend Ribes glutinosum Ribes tenue 2006; Weigend et al. 2002. 157. FAGACEAE BEECH, CHESTNUT & OAK FAMILY Trees or shrubs. Leaves ±gland-dotted, deciduous (Castanea, some Castanopsis, Fagus, some Quercus), simple, alternate or whorled in 3’s (some Trigonobalanus); margins entire, toothed or deeply lobed (e.g. Quercus); petioles present, swollen at base; stipules often triangular, deciduous. Flowers unisexual (plants monoecious or rarely dioecious), actinomorphic. Perianth fused or free, bract-like. Male flowers in heads or catkins; stamen filaments free or rarely basally fused; sometimes pistillode. Female flowers in © KENPEI/ CC-BY-SA-3.0 spikes or solitary, each flower surrounded by bracts (= cupule); staminodes absent or Lithocarpus edulis 6-12; ovaries inferior; carpels fused; locules 2-6(-9); ovules 2 per locule; placentation axile to apical; styles 3-6. Fruits nuts with subtending bracts (= cupule). Genera 8/species ca. 925. Distribution: Widely distributed. ©Böhringer Friedrich/ CC BY-SA 3.0 AT Fruit of Quercus robur Floral formula: P6(-9) A(4-)6-12(-many) Ĝ(2-)3-6(-15) Useful species: Quercus suber (bark used for cork); Castanea (chestnuts). Notes: The flowers are usually wind pollinated or less often insect pollinated (Castanea). References: Huang et al. 1999; Kubitzki 1993; Manos et al. 1993, 2002, 2006; Manos & Stanford 2001; Nixon 1993; Nixon & Crepet 1989; Oh & Manos 2006; WCSP 2013. Generic synopsis Fruit a solitary nut (= acorns) and round in cross-section • Lithocarpus (male inflorescences erect; India to China and New Guinea; ca. 335 spp.). Castanea sativa • Notholithocarpus (cupule scales reflexed and hooked at tip; western USA; 1 sp., N. densiflorus). • Quercus (oaks; male inflorescences pendulous; cupule scales rarely reflexed; temperate northern hemisphere to Malesia and Colombia; ca. 430 spp.). Fruit 1-several nuts and angled (often 3-angled) in cross-section